This is the accepted version of the article: Abella, Juan; Martín Perea, David M.; Valenciano, Alberto; [et al.]. «Copro- lites in natural traps : direct evidence of bone-eating carnivorans from the Late Miocene Batallones-3 site, Madrid, Spain». Lethaia, (July 2021). DOI 10.1111/let.12438 This version is available at https://ddd.uab.cat/record/250383 under the terms of the license 1   Coprolites in natural traps: direct evidence of bone eating carnivorans from the 2   Late Miocene site of Batallones-3 (Madrid, Spain) 3   4   JUAN ABELLA, DAVID M. MARTÍN-PEREA, ALBERTO VALENCIANO, 5   DANIEL HONTECILLAS, PLINI MONTOYA AND JORGE MORALES 6   7   We describe two carnivoran coprolites found in the pseudokarst natural carnivore trap 8   of Batallones-3, from the Late Miocene of Spain. The larger one, comprising multiple 9   indistinguishable fragments of broken and corroded bones, indicates that the producer 10   of the dropping might have been highly capable of crushing the softer parts of large 11   bones. On the other hand, the smaller one shows several relatively larger and more 12   complete bone fragments, thus exhibiting a greater capacity to break and swallow large 13   portions of bone. The external morphology of the large coprolite is similar to that of 14   extant bears, whereas the smaller one more closely resembles that of the living 15   insectivorous hyaenid Proteles in morphology, on one hand, and that of the viverrid 16   Genetta in size, on the other. We hypothesize that the amphicyonid Magerycion anceps 17   was the producer of the large coprolite and the jackal-sized basal hyaenid 18   Protictitherium crassum excreted the smaller one. Thus, we present the first direct 19   evidence of a bone durophagous diet in the carnivorans of Batallones. 20   21   Juan Abella [juan.abella@icp.cat] Institut Català de Paleontologia Miquel Crusafont, 22   Universitat Autònoma de Barcelona, c/ Columnes s/n, Campus de la UAB, 08193 23   Cerdanyola del Vallès, Barcelona (Spain); Juan Abella [juan.abella@icp.cat] Instituto 24   Nacional de Biodiversidad, Pje. Rumipamba N. 341 y Av. de los Shyris (Parque La 25   Carolina) Quito (Ecuador); David M. Martín-Perea [davidmam@ucm.es] Museo 26   Nacional de Ciencias Naturales (MNCN-CSIC), José Gutiérrez Abascal 2, 28006 27   Madrid, Spain; David M. Martín-Perea [davidmam@ucm.es] Geodynamics, 28   Stratigraphy and Palaeontology Department, Faculty of Geology, Complutense 29   University of Madrid, Madrid, Spain; David M. Martín-Perea [davidmam@ucm.es] 30   Institute of Evolution in Africa – IDEA, Madrid, Spain; Alberto Valenciano 31   [a.valenciano@unizar.es] Departamento de Ciencias de la Tierra, Area de 32   Paleontología. Universidad de Zaragoza, C/ Pedro Cerbuna, 12, 50009, Zaragoza, 33   Spain; Daniel Hontecillas [daniel.hontecillastamayo@gmail.com] Museo Nacional de 34   Ciencias Naturales (MNCN-CSIC), José Gutiérrez Abascal 2, 28006 Madrid, Spain; 35   Plini Montoya [pmontoya@uv.es] Departament de Geologia, Àrea de Paleontologia, 36   Universitat de València, Doctor Moliner 50, 46100 Burjassot, València, Spain; Jorge 37   Morales [mcnm166@mncn.csic.es] Museo Nacional de Ciencias Naturales (MNCN- 38   CSIC), José Gutiérrez Abascal 2, 28006 Madrid, Spain 39   40   * Corresponding author. juan.abella@icp.cat 41   42   43   Keywords: Iberian Peninsula, Late Miocene, Pseudokarst, fossil faeces, Carnivora 44   45   46   47   48   49   50   51   Introduction 52   Locality and age. The Cerro de los Batallones (late Miocene [MN10]) is a pseudokarst 53   complex of 9 fossil sites located 30 km south of Madrid (Figure 1). It represents one of 54   the richest and best-preserved Neogene mammalian assemblages on the Iberian 55   Peninsula (Morales et al. 2004, 2008). The fossiliferous deposits are embedded in marls 56   deposited in hourglass-shaped cavities with upper openings (Pozo et al. 2004; Calvo et 57   al. 2013) in which two types of assemblages can be found: (1) carnivoran-dominated 58   assemblages in the lower part of the hourglass structure (e.g., Batallones-1 and BAT-3), 59   interpreted as carnivore traps (Domingo et al. 2011, 2013a, 2013b) and (2) herbivore- 60   dominated assemblages in the upper parts (e.g., Batallones-4 and Batallones-10), 61   interpreted as herbivore traps (Calvo et al. 2013). The coprolites studied in the present 62   paper were found in one of these lower, carnivoran-dominated assemblages (BAT-3). 63   Furthermore, these upper and lower deposits have been found to contain several discrete 64   fossiliferous levels (Martín-Perea et al. 2020). 65   The fauna of the Cerro de los Batallones localities has yielded a rich and diverse 66   assemblage of vertebrate fossils including freshwater fish, amphibians and sauropsids 67   (testudines, squamata) and predatory birds (Morales et al. 2008; Pérez-García & 68   Murelaga 2013; Pérez-García & Vlachos 2014; Morales 2017; Villa et al. 2018), 69   micromammals (López-Antoñanzas et al. 2010, 2014; Álvarez-Sierra et al. 2017; 70   Medina-Chavarrías et al. 2019), equids, bovids, mosquids, giraffes, rhinoceroses, 71   proboscidians, suids (Morales et al. 2008; Sánchez et al. 2009, 2011; Pickford 2015; 72   Ríos et al. 2017; Romano et al. 2017; Sanisidro 2017; Morales 2017; Domingo et al. 73   2018; Ríos & Morales 2019), and carnivorans, which include the most diverse sample. 74   The last group comprises a large amount of skulls, mandibles and almost complete 75   skeletons of two species of sabre-tooth cats, a hyaenid, two species of derived 76   amphicyonids, an ailuropod ursid, a large ailurid, mephitids and mustelids (e.g., Abella 77   et al. 2013, 2015; Antón et al. 2004, 2020; Peigné et al. 2005, 2008; Salesa et al. 2006, 78   2008, 2010, 2012, 2019; Domingo et al. 2013a, 2016; Monescillo et al. 2014; Siliceo et 79   al. 2014, 2015, 2017, 2020; Fabre et al. 2015; Valenciano et al. 2015, 2020; Fraile 80   2016, 2017; Abella & Valenciano 2017; Morales et al. 2017; Valenciano 2017a, 81   2017b). This fauna, alongside the micromammal remains, enabled all the sites to be 82   assigned to the late Miocene (Vallesian, c. 9.6-9.3 Ma) (Peláez Campomanes et al. 83   2017). However, there are minor differences in both micro- and macromammal 84   taxonomic composition among the different localities that have been attributed to slight 85   temporal differences. Indications are that BAT-10 is older than BAT-1, and that BAT-1 86   is older than BAT-3 (López-Antoñanzas et al. 2010; Siliceo et al. 2014)), whereas the 87   difference in age between Batallones-2, 3 and 5 remains unknown (Valenciano et al. 88   2020). 89   Furthermore, the carnivoran guild of BAT-3 is the most diverse of the whole complex. 90   It includes two machairodont felids Machairodus aphanistus (Kaup, 1832) and 91   Promegantereon ogygia (Kaup, 1832) (Monescillo et al. 2014; Siliceo et al. 2014), as 92   well as the small feline Leptofelis vallesiensis (Salesa, Antón, Morales & Peigné, 2012), 93   the plesiomorphic hyaenid Protictitherium crassum (Filhol, 1883) (Fraile 2016, 2017), 94   the ailuropod ursid Indarctos arctoides (Depéret, 1895) (Abella 2011; Abella et al. 95   2012, 2013, 2015; Abella & Valenciano 2017a), two amphicyonids Magericyon anceps 96   Peigné, Salesa, Antón, Morales, 2008 (Siliceo et al. 2017) and the thaumastocyonine 97   Ammitocyon kainos Morales, Abella, Sanisidro, Valenciano (Morales et al. in press), the 98   giant mustelid Eomellivora piveteaui Ozansoy, 1965 (Valenciano et al. 2015; 99   Valenciano 2017a, 2017b), the small size hypercarnivorous mustelid Circamustela 100   peignei Valenciano, Pérez-Ramos, Abella & Morales, 2020 (Valenciano 2020a), the 101   badger Adroverictis sp., and two mephitids Promephitis sp., nov., and Mephitidae gen. 102   nov and sp. nov (Valenciano 2017b). 103   104   Coprolite generalities. Coprolites are among the most important ichnofossils from the 105   fossil record (Hunt & Lucas 2020) and can be found in a diversity of taphonomical, 106   ecological, and geological contexts (Qvarnström et al. 2016). As trace fossils, coprolites 107   represent specific moments in time, providing individual windows into the evolution of 108   ecological interactions such as predation, herbivory, and parasitism, and can contain 109   palaeoecological proxies spanning from thousands to millions of years in the past 110   (Dentzien-Dias et al. 2013; Qvarnström et al. 2017; Barrios-de Pedro et al. 2018; Riley 111   2018). And although they might not give the general diet reconstruction of a species, 112   they can provide direct information about the last food income and feeding behaviours 113   of ancient vertebrates (e.g., Mellett 1974; Meng & Wyss 1997; Chin et al. 1998, 2003; 114   Prasad et al. 2005; Chin 2007; Backwell et al. 2009), serving as ichnological proxies for 115   the presence of animals in palaeoecosystems (Edwards & Yatkola 1974; Farlow et al. 116   2010; Hunt & Lucas 2007, 2010). 117   Coprolites may also provide information on the diagenetic history of the fossil remains 118   themselves (Chin 2007). Over time, following burial, they undergo taphonomic 119   processes, becoming permineralized or lithified and forming a cast or mould of the 120   original faecal matter (Bajdek et al. 2017; Mychajliw et al. 2020). In general, coprolites 121   from sites protected by geologic features such as caves and rock shelters exhibit the 122   highest level of preservation. In contrast, coprolites from open sites can become 123   severely degraded by these taphonomic processes (Reinhard et al. 2019). 124   Coprolites of herbivorous animals are generally scarcer in the fossil record than those of 125   carnivores, because the phosphate content of the latter from the soft tissue and bones of 126   prey animals predisposes them to mineralization (Thulborn 1991; Chin 2002; Hunt & 127   Lucas 2019). On the other hand, coprolites of bone-crushing hyenas are 128   disproportionately represented in the fossil record, not only because of their chemical 129   composition, but also due to: (1) their social and denning behaviour, resulting in large 130   accumulations; (2) their inhabiting caves, which enhances the potential for preservation; 131   and (3) the fact that hyena coprolites are resistant, and can undergo taphonomic 132   resedimentation (Hunt & Lucas 2019). Bones as a supplementary food source 133   necessarily constitute a net gain if the cost of processing them (both ingestion and 134   digestion) can be managed. The average compact bone consists of approximately 30% 135   organic matrix (Eastoe & Eastoe 1954; Guyton & Hall 2006), mostly collagen fibres. 136   The organic component (nutritional value) is even greater if the marrow in the 137   cancellous bone is also considered. Living spotted hyenas (Crocuta crocuta Erxleben 138   1777) in Africa possess morphological and physiological adaptations enabling efficient 139   consumption of bones; accordingly, they are known to consume the entire carcass 140   (freshly killed or scavenged), leaving no bones behind (Kruuk 1972). 141   142   Carnivoran bone cracker adaptations. Living hyenas are known to crush the bones of 143   their prey to extract the nutritious marrow within. This feeding ability is rare today, and 144   both African and Asian hyenas, particularly the spotted hyena, are the only true ‘bone- 145   crackers’ in our modern ecosystems. Bone-crackers (modern and extinct) play an 146   important ecological role as apex predators and providers of free organic material 147   needed for decomposition, essential with regard to maintaining a healthy ecosystem 148   (Wang et al. 2018). 149   In general, hyaenids present the most advanced dentition for crushing bones, whilst 150   retaining the basic feliform dental plan, which involves highly reduced grinding 151   components (in M1, m1 talonid and m2), which allows room for enlargement of the 152   premolars (Wang et al. 2018). The composition of hyena droppings is quite distinctive, 153   enabling them to be differentiated from those of most other carnivorans, a consequence 154   of their high bone content diet (Larkin et al. 2000). The gastrointestinal system of 155   hyenas has apparently evolved to process large quantities of bones. Hyaenid faeces, 156   particularly those of the spotted hyena, are known to contain well-digested calcium 157   phosphates in the form of white powders and bone residues (Estes 1991). To a lesser 158   extent, the scat of striped hyenas Hyaena hyaena (Linnaeus 1758) is also white or light 159   grey (Macdonald 1978; Hulsman et al. 2010). These white powders consist of calcium 160   and phosphate salts, Ca3(PO4)2 1.5Ca(OH)2, similar to hydroxyapatite, the main 161   inorganic component in bones (Kruuk 1972; Wang et al. 2018). Supposing that the 162   common ancestor of Crocuta Kaup 1828 and Hyaena Brisson 1762 already possessed 163   this bone-dissolving gastrointestinal system, then this must have occurred over 8.6 Ma 164   (Koepfli et al. 2006; Wang et al. 2018). 165   Although the vast majority of Miocene coprolites have either been described as being 166   excreted by bone-cracking large Hyaenidae or by the Borophaginae canids (e.g 167   Pesquero et al. 2011; Wang et al. 2018 and references therein), other fossil coprolites 168   have been described as belonging to other vertebrates, including crocodilians, 169   notoungulates, sirenians, rodents, as well as undetermined carnivorans (Dentzien-Dias 170   et al. 2018); some of them even reveal shark bite marks (Godfrey & Smith 2010). 171   Batallones-3 does not present any typically large durophagous bone-cracker species; 172   hence, study of these coprolites constitutes a very interesting approach to bone 173   consumption in other dietary groups of carnivorans. The main aim of the present 174   research paper involves analysing the morphology and bone content of the only two 175   coprolites unearthed in the whole Cerro de los Batallones site complex in order to 176   obtain information on the possible producer of the scat and to more clearly elucidate 177   their dietary ecology. 178   179   Materials and methods 180   Abbreviations. BAT: Batallones fossil site from Cerro de los Batallones fossil site 181   complex. 182   MNCN- Museo Nacional de Ciencias Naturales, Madrid, Spain. 183   184   Studied material. The fossil sample of coprolites of BAT-3 comprises the following two 185   specimens: BAT-3’9.178 and BAT-3’10.153. 186   187   Methodology. We followed the method proposed by Diedrich (2012) and modified by 188   Wang et al. (2018) to characterise the external morphology of hyaenid coprolite 189   aggregate pellets (Figure 2). 190   191   3-D models. Coprolites BAT-3’9.178 and BAT-3’10.153 were CT-scanned at the 192   “Servicio de Técnicas No Destructivas: Microscopía Electrónica y Confocal y 193   Espectroscopía” of the MNCN-CSIC in Madrid, Spain. The scanned data were imported 194   into FIJI 1.52 (National Institutes of Health, USA) for artefact removal and contrast 195   enhancement. The data were finally segmented, rendered and analysed in Avizo 7.1.0 196   (VSG, Burlington, MA, USA) in order to generate 3D models and to perform metrical 197   and morphological analyses. We measured the trend and plunge of elongated bone 198   fragments with a clearly longer axis, using the coprolites’ long axis as the pole (0º trend, 199   90º plunge) and its corresponding plane (0º trend, 0º plunge) with a theoretical fixed 200   north bearing to measure the trend. Trend and plunge data were plotted on an equal area 201   Schmidt stereographic projection with inverse area smoothing. 202   Following Esteban-Nadal et al. (2010), all the fragments were measured in length and 203   volume (Supplementary material 1). However, only the ones exhibiting a more 204   complete morphology are presented in the present paper (TABLE 1). The label number 205   was given in the order of segmentation and has no specific meaning whatsoever. 206   207   Results 208   Bone fragments inside the coprolites showed preferential orientations very similar to 209   that of their major axis (0/360º trend, 90º plunge). This is particularly so in the large 210   one, with a mean trend and plunge closely resembling the coprolite’s major axis (308.0º 211   trend, 87.7º plunge; Figure 3), whereas in the small one, bones present a smaller, albeit 212   still steep (93.9º trend, 64.4º plunge) mean plunge compared to that of the coprolite’s 213   major axis (Figure 3). 214   215   Description and interpretation. 216   BAT-3’9.178: Consists of a large fragment of an almost cylindrical coprolite 116.3 mm 217   in length, 52.5 mm in width, 34.7 mm in height and a volume of 83.5738 cm3 (Figure 218   4). It shows no apparent torsion and consists of one massive portion of scat, with no 219   internal or external divisions. The total number of bone fragments presenting a volume 220   greater than 4.21 mm3 is 134, although most of them are exclusively small fragments of 221   trabecular bone. In consequence, few fragments are of sufficient size or have 222   sufficiently intact surfaces to be considered for description. Hence, we mainly focused 223   upon fragments with some preserved portion of cortical bone, and on others displaying 224   any outstanding morphological trait. 225   226   The main bones found in the coprolite are (3D images as supplementary files, each one 227   with its label number): 228   229   9: Two associated fragments of bone (21.13 mm), one of these constituting a large piece 230   of cancellous bone and the other a relatively flat fragment of triangular cortical bone. 231   The latter presents a rounded depression that might represent a partial tooth mark. 232   233   29: A relatively large and flat portion of cancellous bone (24.19 mm). Although its 234   shape is not recognizable, its size and morphology are worth highlighting. 235   236   104: A thick portion of cortical and cancellous bone (14.28 mm) possibly belonging to 237   the surface of a bone epiphysis or vertebra. 238   239   107: A large portion of cancellous bone attached to a thick portion of curved cortical 240   bone (27.61 mm). Possibly also from the side of a bone epiphysis or vertebra. 241   242   108: A large and robust portion of cortical bone (19.38 mm). It has a marked crest along 243   the centre, this constitutes one of the only distinctive structures recovered in this 244   sample. 245   246   116: Fragment of curved bone surface (18.92 mm), which may represent the largest and 247   most distinct portion of bone surface in the whole sample. However, its overall size and 248   shape are insufficient for a precise anatomical determination. 249   250   BAT-3’10.153: It is much smaller in size than the other coprolite, measuring 51.6 mm 251   in length, 22.3 mm in width and 21.8 mm in height (Figure 4). It consists of three 252   clumped fragments, which show a slight torsion-like morphology. It is relatively thick 253   and although it is thinner at both ends it does not present an elongated tip. 254   The main bones found in the coprolite are (3D images as supplementary files #2): 255   256   15: An elongated fragment (8 mm) of cortical bone, which shows a foramen half-way 257   along its length. It could be part of a larger bone. 258   16: A relatively large fragment of an undetermined bone (21.02 mm). It represents a 259   portion of the curved shaft of a long bone, with cancellous bone in its inner part. 260   261   17: A relatively long and flat bone fragment (17.83 mm) with several longitudinal crests 262   dividing the surface into several flat-to-concave areas. It could be part of the epiphysis 263   of a long bone or a fragment of the spine of a scapula. 264   265   21: An elongated and triangular bone fragment (20.97 mm). This bone is one of the 266   most complete remains; it appears to be almost complete, only missing the two 267   epiphyses. It also presents a foramen towards the central part of the shaft. 268   269   22: A large and almost semi-circular bone fragment (17.60 mm). It might be one half of 270   a proximal epiphysis, likely belonging to a femur. There is a small cavity interpreted 271   herein as the fovea capitis, which serves for the attachment area for the ligamentum 272   teres. It also presents a rounded puncture and a cutting chip that could be interpreted as 273   a tooth mark. 274   275   26: An elongated and curved bone fragment (4.98 mm). There are two rounded 276   structures in the middle of the bone, one larger than the other, forming a line running 277   parallel to both sides of the bone. This could represent a mandibular or maxillary 278   fragment with tentatively two teeth. 279   280   27, 29 and 30: These three elongated bone fragments (3.03 mm, 5.24 mm and 8.37 mm, 281   respectively) could belong to one single larger fragment which, due to its morphology, 282   could be interpreted as a small vertebrate mandible. It has some sulci and foramina 283   similar to those found in reptile bones, but this is difficult to assert due to the corrosion 284   and general state of preservation of the specimen. 285   286   33: A compact small cylindrical fragment of bone (2.68 mm) with a foramen on one of 287   its sides. It could be identified as a fragment of a large long bone, but its broken nature 288   hinders a more specific determination. 289   290   43: A robust bone structure (19.96 mm) that appears to be part of a larger, more 291   complex structure, due to its morphology and overall inner structure. Although it cannot 292   be completely proven due to its state of preservation, this structure might well represent 293   a partial skull of a small undetermined vertebrate. 294   295   Discussion 296   297   Following Diedrich (2012), BAT-3’9.178 is a long oval portion of scat similar in shape 298   to those of Ursus arctos (Chame 2003) (Figure 5). The large amount of broken bone 299   material in the coprolite appears to indicate a high consumption rate of bone matter; 300   however, this is still far from that found in modern hyaenas (Estes 1991). Although 301   some of the remains still possess some cortical bone, most of the fragments consist 302   mainly of cancellous bone. This leads to the interpretation that only parts of the 303   epiphysis of long bones or vertebrae were gnawed and consumed by the producer of the 304   coprolite, similar to what is mostly consumed by modern canidae (Mech 1970; Munthe, 305   1989; Sillero-Zubiri 2009). 306   The size of this coprolite fragment means it can only have been produced by one of the 307   four large-sized carnivorans present in BAT-3 (TABLE 2): the sabre-tooth cat 308   Machairodus aphanistus, the amphicyonids Ammitocyon kainos and Magericyon 309   anceps or the bear Indarctos arctoides. The first two are large hypercarnivorous 310   carnivorans, [Machairodus aphanistus 117-285 kg following Domingo et al. (2013a) 311   and Ammitocyon kainos 231 kg according to Morales et al. (in press)] with an extreme 312   specialized dentition for cutting meat which would have prevented them from eating all 313   but the most easily accessible portions of flesh on a carcass, leaving large quantities of 314   food (including bones) for other carnivorans and scavengers (Ewer 1954, 1998; Turner 315   1992; Pesquero et al. 2011). 316   At the other end of the diet spectrum is the hypocarnivorous Indarctos arctoides, a 317   member of the Ursidae family related to modern giant pandas. Its estimated body mass 318   ranges from 137 to 266 kg (Abella 2011; Abella et al. 2013; Domingo et al. 2016). It 319   has been suggested that this bear might have had a more omnivorous diet than that of 320   giant pandas (Abella et al. 2012; Abella et al. 2011; Abella et al. 2014), but also that 321   their diet would have included abundant plant material (Abella et al. 2011; Monescillo 322   et al. 2014; Domingo et al. 2016). Consequently, as frequently occurs with other bear 323   scats (Hewitt & Robbins 1996), this would have hindered the fossilization process. 324   However, the dentition of I. arctoides shows a certain capacity for durophagy, similar to 325   that of smaller species of related bears (de Bonis et al. 2017). This would have allowed 326   this species to shift from eating tough plant material to gnawing bone with some degree 327   of success, especially in individuals challenged with finding their favourite food 328   sources, e.g. the young. For this reason, this species cannot be completely ruled out as 329   the producer of this particular coprolite. 330   Finally, Magericyon anceps was a large amphicyonid (Peigné et al. 2008) present in 331   two of the Batallones fossil traps (BAT-1 and BAT-3; Siliceo et al. 2015, 2017, 2020), 332   with an estimated body mass of 175-195 Kg (Domingo et al. 2013a; Siliceo et al. 2015). 333   The described dentition is similar to other larger Amphicyonidae, but slightly more 334   trenchant, thus indicating a dentition less capable of crushing (Peigné et al. 2008). 335   However, the wear pattern observed in the carnassial teeth of M. anceps is quite similar 336   to that of other large amphicyonines, implying a similar occlusal mechanism, which 337   probably helped this animal to gnaw on the epiphysis (much softer than the diaphysis) 338   of long bones, using its P4-M2 and m1-m2 teeth in the same way as extant canidae. 339   Moreover, although in BAT-3 there are not abundant remains of this species, the same 340   mortality rate as in BAT-1 is to be expected, because the majority of the individuals are 341   young adults and juveniles (Peigné et al. 2008). As previously mentioned, this age 342   group might well have consumed a slightly different diet than their more adult 343   counterparts. We therefore hypothesize that this species represents the most likely 344   producer of this coprolite. 345   In this specific case we do not coincide with Lofgren et al. (2017), who consider that 346   the coprolite was likely produced by the more abundant of the two species, since the 347   reconstructed diet of M. anceps more closely resembles the crushed bone fragments 348   found inside the coprolite. 349   350   BAT-3´10.153 constitutes an irregular portion of scat following Diedrich (2012). It is 351   similar in shape to the scats of the living aardwolf Proteles cristatus (Sparrman 1783) 352   (Stuart & Stuart 1994) or the common genet Genetta genetta (Linnaeus 1758), and 353   similar in size to the latter (Chame 2003) (Figure 5). Isolated bones account for 35.79% 354   of the total volume of the coprolite (approximately 26.13% were individualised), but a 355   large amount thereof could consist of digested bone matter or powder as described in 356   Estes (1991). Most of the bone fragments are amorphous and unidentifiable, and just a 357   few of them can be partly identified anatomically, but not taxonomically. In other 358   published articles, the percentage of unidentifiable (both anatomically and 359   taxonomically) bone fragments from fossil coprolites and extant scats from canids 360   ranges from 80% to 95% (Fosse et al. 2012; Wang et al. 2018) 361   362   The three individual sections of the coprolite can be characterised by containing bone 363   fragments presenting different sizes and morphologies, especially on comparing the 364   ones located at both ends. The assemblage in one of these sections could represent the 365   remains of a small vertebrate (perhaps a reptile), due to the abundance of small, dense 366   individual bones (around 15), together with a large mass of broken cancellous bone 367   matter. On the other hand, the other outer section mainly consists of relatively large 368   cortical bones or fragments of long bones arranged subparallel to the long axis of the 369   coprolite (see figure 4). Indeed, the relative size of these fragments is noteworthy in 370   relation to the size of the coprolite, even more so when compared with the larger one 371   (Figure 6). The size and morphology of this coprolite appear to indicate that several 372   carnivorans that could have produced it. The first and most probable carnivoran is 373   Protictitherium crassum, a small-medium sized carnivoran situated at the base of the 374   Hyaenidae (Fraile 2016, 2017; Morales et al. 2019). This coprolite morphology is 375   similar to that of small hyenas such as Proteles cristatus (Stuart & Stuart 1994) and its 376   size fits well with the ones produced by medium-to-large viverrids (Chame 2003). 377   Furthermore, the fact that the coprolite has relatively large broken cortical bone 378   fragments might suggest that this individual was already performing “bone-cracking” 379   feeding, but did not have the enzymatic adaptation of extant hyenas to almost 380   completely dissolve bones, reducing them to powder during digestion (Estes 1991). 381   This fits well with an early member of the hyaenid family, such as P. crassum, which 382   has slightly enlarged premolars but does not yet present reduced postcarnasssial molars 383   like the more derived hyaenids, thus distributing the crushing dental apparatus between 384   the premolars and the molars (Fraile 2016, 2017). 385   Another species that could have produced this coprolite is the medium-sized feline 386   Pristifelis Salesa, Siliceo, Antón, Peigné, Morales 2012. However, this species is not 387   described in BAT-3, and as occurred with the larger coprolite, these felids are 388   interpreted as hypercarnivorous animals, and might not have been able to break bones to 389   feed upon them. Furthermore, the shape of the coprolite does not fit well with those of 390   this group, since felid scats are more elongated (Chame 2003). Gilmour & Skinner 391   (2011), however, on studying the scats of small felids, concluded that they have 392   disproportionately large bone fragments in their scat compared with canids, whose scats 393   contain relatively more abundant and smaller bone fragments. Furthermore, a broad 394   musteloid sample is present in BAT-3, comprising relatives of the living martens, 395   badgers, skunks, and honey badgers, of which the badger-sized Adroverictis sp. 396   Ginsburg & Morales, 1996 and the giant sized Eomellivorini Eomellivora piveteaui 397   (Valenciano et al. 2015; Valenciano & Govender 2020b) might have been the producers 398   of the coprolite. Eomellivora may have had a clearly durophagous diet, based on dental 399   morphology. However, its large size rules out its assignment to this coprolite. Although 400   the size could fit well with a mustelid the size of Adroverictis, the reconstructed diet 401   (which was likely more omnivorous) together with the morphology of the scat also 402   brings us to rule out this species with a certain degree of certainty. 403   As previously mentioned, the carnivoran guild of BAT-3 is the most complex of all the 404   Batallones fossil sites, presenting many different body sizes and dietary groups. 405   Moreover, the Batallones carnivoran assemblage is unique compared to sites of similar 406   age on the Iberian Peninsula due to the complete absence of large or medium hyenas. 407   The lack of these forms, present both before and after this period in this region (Fraile et 408   al. 1997; Morales et al. 2015), is difficult to assess without conducting a detailed 409   analysis comparing the ecomorphologies of the carnivoran guilds of the Vallesian, in 410   order to provide a broader view of how the ecological roles of these niches evolved 411   throughout this time period; similar to those already performed by Morlo et al. (2020) 412   across the Middle/Late Miocene boundary in Germany. 413   Finally, a taphonomical issue is raised by the presence of these coprolites in BAT-3. As 414   previously mentioned, the lower carnivoran-dominated assemblages, such as BAT-3 415   and BAT-1 have been interpreted as natural traps (Domingo et al. 2013a, 2013b; 416   Domingo et al. 2016; Domingo et al. 2011). In BAT-1 for example, none of the large 417   bones within the trap were found to be eaten or gnawed (Domingo et al. 2013), although 418   some Moschidae remains present corrosion surfaces that could be interpreted as 419   resulting from digestion (Sánchez et al. 2011). Future taphonomical analyses for 420   Batallones-3 should address whether or not the presence of coprolites suggest the site 421   acted as a den rather than a natural trap, since the latter is likely to present fewer 422   coprolites. 423   424   Conclusions 425   Two coprolites were discovered in one of the nine sites of the Batallones complex. 426   Although these fossils are rarely preserved, the particular genesis of these carnivoran 427   natural traps in cavities would theoretically have helped to conserve them. 428   The larger coprolite could be described as a “long oval” portion of a scat. It mainly 429   presents broken parts possibly involving an epiphysis of a long bone or a vertebra. This 430   fact would indicate that the producer would have been able to gnaw on a bone, 431   Magericyon anceps being the most likely species or, to a lesser extent, Indarctos 432   arctoides. 433   The small coprolite could be defined as an almost terminal “irregular portion” of a scat 434   with the size and morphology of those produced by large viverrids and small hyaenas. It 435   presents two differentiated parts: one that appears to result from eating a complete and 436   unidentified small vertebrate, and the second produced by swallowing the remains of a 437   relatively large broken bone. The species that produced this coprolite could have been 438   possibly either Protictitherium crassum or, less likely, Adroverictis sp. 439   Finally, study of these coprolites provides direct evidence of bone consumption 440   amongst carnivorans in the trophic network of the feeding complex of the Batallones 441   palaeocommunity. In the case of Magericyon, this bone consumption appears to confirm 442   that large Amphicyonidae display adaptations related to generalist feeding; nonetheless, 443   one of the members of the group, Magericyon anceps seems to have developed a 444   tendency towards hypercarnivorism. 445   446   447   Acknowledgements 448   This study was supported by the Spanish Research Projects CGL 2015-68333-P, 449   CGL2016-76431-P and PGC2018-094122-B100 (AEI/ FEDER, UE), the BSCH- 450   UCM910607 of the Universidad Complutense de Madrid, the Research Groups CSIC 451   64 1538 and CAM-UCM 910607 and PVC-GIUV, and the Generalitat de Catalunya 452   (CERCA Programme, and Beatriu de Pinós contract 2017 BP 00223 from AGAUR to 453   JA). We thank Alejandro Serrano for his work on the CT-images as an ICP technician 454   and Alejandro Pérez-Ramos for his help dealing with the files. The “Juan de la Cierva 455   Formación” program (FJC2018-036669-I), from the Spanish Ministry of Science, 456   Innovation, and Universities funded also AV. DMMP was funded by a FPI Predoctoral 457   grant BES-2016-079460. We thank Cormac De Brun for the thorough language review. 458   The editor B. Britt and the reviewers Dr. de Bonis and Dr. Morlo are also acknowledged 459   for valuable comments that helped to improve the final version of this manuscript. 460   References 461   Abella, J. 2011 (Thesis). Indarctos arctoides Depéret, 1895 (Carnivora, Mammalia) del 462   yacimiento vallesiense de Batallones 3 (cuenca de Madrid). pp. 406. 463   Departamento de Biología, Facultad de Ciencias, Universidad Autónoma de 464   Madrid. 465   Abella, J. & Valenciano, A. 2017: Indarctos arctoides: Los ancestros de los grandes 466   osos Panda, in: Morales, J. (Ed.), La colina de los Tigres Dientes de Sable. 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B, 931   Location of Batallones-1 and Batallones-3 sites, west of Valdemoro. 932   933   Figure 2 934   Nomenclature of individual pellets in a complete assemblage of faeces from a single 935   dropping event by the extant spotted hyena, Crocuta crocuta. Horizontal axis indicates 936   the orientation of the dropping. After Diedrich (2012): Figure 4 and Wang et al. (2018): 937   Figure 1. 938   939   Figure 3 940   Stereographic projections of the elongated axis of bone fragments in the coprolites. 941   Mean trend and plunge represented by plus sign. A, BAT-3’9.178 coprolite. B, BAT- 942   3’10.153 coprolite. 943   944   Figure 4 945   View of the disposition of the bone fragments present in A, Transparent outline of the 946   coprolite BAT-3-’10.153 with the fragments in colours. B, Inner view of coprolite 947   BAT-3-’10.153 with amorphous bone matter in grey and fragments in colours. C, Inner 948   view of coprolite BAT-3-’10.153 only showing fragments in colours. 949   D, Transparent outline of the coprolite BAT-3’9.178 with the fragments in colours. E, 950   Inner view of coprolite BAT-3’9.178 only showing fragments in colours. Scale bar 1 951   cm. 952   953   Figure 5 954   Figure comparing extant dropping outline morphology from Chame (2003) to the 955   coprolites BAT-3’9.178 and BAT-3-’10.153 956   957   Figure 6 958   View of the disposition of the bone fragments present at the coprolites. A, BAT-3’9.178 959   and B, BAT-3-’10.153. Same size, not to scale. 960   961   Table captions 962   Table 1 963   Measurements of the Maximum length and second maximum length (in mm) and the 964   volume (in mm3) of the described bone fragments found inside the coprolites BAT- 965   3’9.178 and BAT-3’10.153 from BAT-3, Spain. 966   967   Table 2 968   Estimated body mass (in Kg) of the BAT-3 carnivorans discussed in this work and both 969   the average body mass (in Kg) and the coprolite length (in cm) of the extant carnivorans 970   used for comparison. References: 1.-­‐  Domingo et al. 2013a, 2016; 2.- Fraile 2016; 3.- 971   Garshelis 2009; 4.- Holekamp and Kolowski 2009; 5.- Larivière S. & Calzada, J. 2001; 972   6.-Lariviére & Jennings 2009; 7.- Nowak 2005; 8.- Sillero -Zubiri 2009; 9.- Sunquist & 973   Sunquist 2009; 10.-Valenciano 2017; 11.- Wade-Smith & Verts 1982; 12.- This study, 974   approximated based on similar-sized living relatives. 975   976   Supplementary data captions 977   978   Supplementary material 1 979   Measurements of the Maximum length and second maximum length (in mm) and the 980   volume (in mm3) of all the bone fragments found inside the coprolites BAT-3’9.178 and 981   BAT-3’10.153 from BAT-3, Spain. The numbers only reflect the order in which the 982   images were segmented from the original CTscan files. 983   984   Other supplementary material 985   Supplementary 3D files of the described bone fragments found inside the coprolites 986   BAT-3’9.178 and BAT-3’10.153 from BAT-3, Spain. 987   988   989   990   991   992   993   994   995   996   997   998   Table  1     Second Maximum maximum Volume Coprolite label Number length (mm) length (mm) (mm3) BAT-3’9.178 #9 21.13 16.53 507.07 #29 24.19 22.05 577.15 #104 15.27 9.63 193.45 #107 27.61 19.33 1088.14 #108 19.38 9.5 232.48 #116 18.92 8.49 206.44 BAT-3’10.153 #15 8 1.99 9.841 #16 21.02 12.4 645.641 #17 17.83 5.07 107.562 #21 20.97 8.91 270.119 #22 17.6 16.19 902.955 #26 4.98 0.77 2.036 #27 3.03 1.46 1.763 #29 5.25 1.56 3.871 #30 8.37 2.15 12.228 #33 2.68 1.89 3.911 #43 19.96 8.83 355.843   Table  2   Estimated body Average body Coprolite Fossil Species mass (Kg) Ref. Extant Species mass (Kg) Ref. length (cm) Machairodus aphanistus 117-285 1 Panthera leo 120-225 9 15 Magericyon anceps 175-195 1 Ursus americanus 40-225 3 8-11 Thaumastocyon sp. 170-321 1 Canis lupus 18- 80 8 16 Indarctos arctoides 137-266 1 Promegantereon ogygia 28-97 1 Panthera onca 31- 121 9 2.6- 11.2 Eomellivora piveteaui 35.8-45.5 10 Puma concolor 34- 72 9 3.7- 6.1 Leptofelis Parahyaena vallesiensis 7–9 12 brunnea 28- 47.5 4 5 Protictitherium crassum 6- 7 2 Gulo gulo 6- 18 6 13 Circamustela peignei 3-5 12 Mellivora capensis 7- 13 7 6.8 Adroverictis sp. 8-12 12 Taxidea taxus 6.3- 8.7 6 3.4- 4.9 Promephitis sp. 1-5 12 Genetta genetta 1- 3 5 5.5 Mephitidae Gen. nov and sp. nov 1-2.5 12 Mephitis mephitis 1.2-5.3 11 3-4.4