Ichnos, 11:371–384, 2004Copyright © Taylor & Francis Inc.ISSN: 1042-0940 print / 1563-5236 onlineDOI: 10.1080/10420940490444951

Carnivore Trackways from the Miocene Site of Salinasde Anana (Alava, Spain)

Mauricio AntonDepartamento de Paleobiologıa, Museo Nacional de Ciencias Naturales, Madrid, Spain

Gregorio Lopez and Robert SantamariaDepartament de Geologia (Paleontologia), Facultat de Ciencies, Universitat Autonoma de Barcelona,Spain

A fossil tracksite of Lower Miocene age discovered near Salinasde Anana, Alava (Spain), has rendered an exceptionally well-preserved assemblage of vertebrate ichnites. The site shows a highproportion of carnivore tracks (3 out of 5 mammal ichnospecies)and a high number of individual trackways (15), some includingover 50 consecutive footprints. The carnivore ichnites are classifiedas Felipeda lynxi Panin & Avram, 1962, Felipeda parvula ichnosp.nov. and Canipeda longigriffa Panin & Avram, 1962, and they are at-tributed to a felid, an undetermined small aeluroid, and a herpestid,respectively. The long trackways allow determination of gaits,which include lateral sequence walks and diagonal sequence trots,and of speed, which ranges from 0.4 to 1.4 m/s. Froude numbersrange between 0.1 and 0.8, agreeing with gait interpretations andspeed calculations. The felid trackways provide the first known evi-dence of group traveling in fossil cats. The herpestid footprints showmodern-grade adaptations for terrestrial locomotion and digging.

Keywords Mammalia, ichnites, trackways, carnivores, cats, mon-goose, Miocene, Spain

INTRODUCTIONA lower Miocene tracksite discovered near Salinas de Anana

(Alava, Spain) (Laumen, 1989; Anton et al., 1993 and 1999)shows abundant, exceptionally preserved vertebrate trackways(Fig. 1). About 60 square meters of the site have been excavateduntil now, yielding about 700 footprints, many of them in cleartrackways (Figs. 2 and 3).

The site includes at least 15 carnivore trackways, some of ex-ceptional length with over 50 consecutive footprints, and with anexquisite preservation of morphological detail. Apart from thecarnivore trails, there is another clear mammal trackway (marked

Address correspondence to Mauricio Anton, Departamento dePaleobiologıa, Museo Nacional de Ciencias Naturales, C/Jose GutierrezAbascal 2, Madrid, 28006, Spain.

“U” in Fig. 2) corresponding to a small ruminant, although thereare many footprints of a small artiodactyl that cannot be eas-ily grouped into individual trackways and are attributed to amember of the family Cainotheridae (Anton et al., 1993 and1999). This abundance of carnivore trackways is remarkable,considering that carnivore ichnites are rare in the fossil record,so that even in exceptional tracksites with long trackways ofother mammals only isolated footprints of carnivores are usu-ally found (Robertson and Sternberg, 1942; Alf, 1959; Scrivnerand Bottjer, 1986; Perez-Lorente et al., 1999). Also, and in therare cases where trackways are available, preservation of mor-phological detail is usually poor (Alf, 1966; Leakey and Harris,1987). The tracks from Salinas allow the most detailed study todate of the locomotion of Miocene carnivores, as well as mor-phological comparisons that reveal the phylogenetic affinities ofthe trackmakers.

In addition to the mammalian trackways, the site preservestwo bird ichnospecies. All the tracks are preserved in a fine-grained limestone laid on an ancient lakeshore. The remarkabledetail of the footprints suggests that they were produced near theshoreline, within the strandline zone (Cohen et al., 1991, 1993).In this zone the water content of the sediment allows excellentpreservation of small animal footprints, but high bioturbationtends to destroy them quickly, so burial must occur within daysof ichnite production to allow preservation. These data pointto a lacustrine basin subject to sudden, seasonal floods, one ofwhich buried the tracks and led to their preservation. This con-clusion is also supported by detailed sedimentologic studies andregional palaeogeographic reconstructions on our studied area(Dreikluft, 1996; Laumen, 1989), that consistently report thepresence of braided rivers and ephemeral floodplain lakes dur-ing the Oligocene and lowermost Miocene in this region. TheSalinas de Anana section that includes the track-bearing bedsis sedimentologically characterized by an alternation of sand-stones and silt beds with load casts related to the high original

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FIG. 1. Geographic situation of the Salinas de Anana tracksite.

water content of the sediment, some fine-grained limestoneswith charophytes, and Pulmonata gastropods related to lake fa-cies. This stratigraphic sequence clearly corresponds to the distalpart of a fluvial system close to some lakes (facies associationFA VII of Dreikluft, 1996).

The sediments are dated as Burdigalian (= Aragonian) in pre-vious geological studies (IGME, 1978; Dreikluft, 1996) and theichnites are consistent with this age (Anton et al., 1993, 1999).The mongoose tracks are significant because herpestid fossilsare first recorded in Europe by MN4, in the early Aragonian(Schmidt-Kittler, 1987; Ginsburg, 1990). The abundant tracksof a minute artiodactyl attributed to the family Cainotheridae(Anton et al., 1993, 1999) are also important because this fam-ily is last recorded in biochronological zone MN 6 (de Bruijnet al., 1992). According to the biochronological scheme pro-posed by Mein (1979), the above data date the site between thebeginning of MN4 (20 Ma) and the end of MN 6 (15 Ma).

MATERIALS AND METHODSThe tracks were traced on sheets of Perspex directly over

the sediment. All 15 carnivore trackways were measured, butin this paper we only present analyses of seven, which displaythe best preservation of locomotor information, and are repre-sentative of the rest. Footprint dimensions, stride length andglenoacetabular distance were measured directly on the track-ways, following the methodology of Leonardi (1987). Glenoac-etabular distance (Fig. 4) is defined as the distance between twopoints, which are the theoretical projection on the ground of the

shoulder and hip joints, approximating the living animal’s bodylength. Methods for estimating glenoacetabular distance varyaccording to the gait of the trackmaker. In trackways indicatinga trot or a walk in diagonal sequence, the distance measured isthe segment above the midline that unites the intersection pointsbetween the midline and two lines: the one joining the referencepoints of the manus and the one joining the reference points ofthe pes (Leonardi, 1987, pl.8-c). An alternative, slightly differentmethod for estimating glenoacetabular length is to measure thedistance between the midpoint of the segment uniting the twoconsecutive hind footprints and the midpoint of the segmentjoining the next pair of consecutive fore footprints (Moratallaet al., 1989). This method yielded more consistent estimates innarrow trackways, so we favored it when measuring the Salinastrails that corresponded to diagonal sequence gaits (Fig. 5, leftand center). In the case of trackways corresponding to a walkin lateral sequence, the distance measured is the length of thesegment of the midline that joins the two following points: 1, theprojection of the reference point of the more advanced forefootand 2, the intersection of the midline with the line that joins thereference point of the two hind feet of the other two sets (Fig. 5,right). As stressed by Leonardi (1987, p. 48), there are no abso-lute rules to determine which hypothesis to choose, so the optioncan only be taken after careful examination of each trackway.In the case of the Salinas tracks analyzed in this study, the greatlength of most of the trackways facilitated the determination ofgait, because many successive series can be measured to test theconsistency of the hypotheses.

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FIG. 2. Trajectories of the main trackways. Tracks named R are Felipeda lynxi, K is for Felipeda parvula and I for Canipeda longigriffa, U is a small artiodactyleand A are bird tracks. Numerous tracks of a very small artiodactyle cross the site in all directions, but have been omitted in this drawing to avoid confusion. Noteparallel felid trackways R6, R7, R9 and R10.

Determining the gait was done through examination of thetrackway configuration using the methodology of Halfpenny(1986) and Leonardi (1987).

The speed of the trackmakers was calculated using the for-mula of Alexander, V = 0.25 × g 0.6 ×λ1.67× h−1.17(Alexander,1976). This method in turn requires an estimate of limb height(femur plus tibia plus metatarsal plus 9%) for each trackmaker.To obtain such estimates, we reviewed the body proportionsof comparable carnivores both living and extinct. Using pho-tographs of living animals and published figures for hind-limbbone lengths (Walker, 1985) and for footprint length (Stuart andStuart, 1994), we calculated the following ratios for extant car-nivores:

1) Ratio of limb height to footprint length.2) Ratio of limb height to glenoacetabular length.

The first ratio (Table 1) was calculated from published dataof five species of extant African small to medium-sizedaeluroids: serval cat, Felis serval (Schreber, 1776); leopard,Panthera pardus (Linnaeus, 1758), common genet, Genettagenetta (Linnaeus, 1758); african civet, Civettictis civetta(Schreber, 1776); and egyptian mongoose, Herpestes ichneumon(Linnaeus, 1758). The latter species is regrettably the only her-pestid for which we found data on both footprint size and skeletalmeasurements to match, but since Herpestes (Illiger, 1811) is arather generalized herpestid and likely a reasonable model for

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FIG. 3. Photograph of a section of the Salinas tracksite. On the left side there is a trackway of the ichnospecies Felipeda lynxi and tracks of a small ungulate.Another Felipeda lynxi trail crosses from the top of the picture toward the right bottom corner, and broadly parallel below it runs a trail of Felipeda parvula. Atrackway of Canipeda longigriffa (mongoose tracks) runs across the right bottom corner.

the more primitive, early members of the family, we consider thatthe data provide a valid approximation. In addition, our reviewof other herpestids, not included here because only part of theset of measurements could be taken on each species, indicates abroad similarity in proportions within the family.

Although the leopard is considerably larger than the othercarnivores in the list, it was included because its short-limbedproportions are more likely to resemble those of the lowerMiocene felids than the extremely long-limbed, but more con-

FIG. 4. Outline drawing traced from a photograph of a banded mongoose, showing the positions of the main limb bones as inferred from the situation of relevantpoints of the external anatomy (Done et al., 1999). Vertical broken lines indicate the projections on the ground level of the glenoid of the scapula and acetabulumof the pelvis, which in turn define the glenoacetabular distance (a-b).

veniently sized, serval cat (Ginsburg, 1961a). The results ofthis review show small to medium aeluroids with retractableclaws averaging 9.23 (average of the values for serval, leop-ard, civet and genet shown in Table 1), while African Her-pestes, displays a much lower value at 5.8. It thus appears thatherpestids have longer footprints (excluding the claws) rela-tive to hind limb dimensions. We estimated the height of thelimb of the Salinas carnivores, which have retractable clawsusing the average value for modern cats and viverrins (9.23),

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FIG. 5. Diagrams of carnivore trackways recognized at Salinas. Left, K3 (Felipeda parvula); center, I1 (Canipeda longigriffa); right, R2 (Felipeda lynxi). Hindfootprints are filled in black. Segment a-b = glenoacetabular distance; M, midline; Sf, step of forelimb; Sh, step of hind limb; St, Stride. Scale bars = 5 cms.

while for the Salinas herpestid we used the value of modernHerpestes.

The second ratio was calculated from the analysis of pho-tographs of living animals in lateral view (Fig. 4). To estimatethe relative lengths of the hind-limb bones and the glenoacetab-ular distance, we inferred the position of the articulations using

TABLE 1Average footprint length (l), Hind-limb height, equaling femur + tibia + metatarsus + 9% (h) and h/l ratio in five extant African

aeluroid carnivores. Footprint data from Stuart and Stuart (1994), and limb bone data from Walker (1985).

Serval Leopard Genet Civet Mongoose

Average footprint length (l) 45 90 21 48 42Limb height (h) 495 670 237 344 245h/l 11 7.4 11.3 7.2 5.8

reference points in the external surface of the body, as defined byDone et al. (1997). After reviewing hundreds of photographs, wefound that only a few of them showed the animals in side viewwith body proportions clear enough and with their entire limbsvisible. We analyzed 11 photographs in side view correspondingto 7 species of small aeluroid carnivores that were considered

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acceptable analogues of the Salinas trackmakers because of theirdigitigrade stance and generalized body proportions. The speciesanalyzed were: domestic cat, Felis catus (Linnaeus, 1758) (n =2); serval cat (n = 1); common genet (n = 3); African civet(n = 2); Asian ground civet, Viverra tungalunga (n = 1);white tailed mongoose, Ichneumia albicaudia (Cuvier, 1788)(n = 1); and banded mongoose, Mungos mungo (Gmelin, 1788)(n = 1). Our review shows that felids, viverrids, and herpestidsare rather homogenous, averaging a ratio of 0.93 with a minimumof 0.85 and a maximum of 1,03. Variation within the sample ofone species was comparable to that observed between species,which makes us confident that the ratio is fairly constant amongsmall digitigrade members of the Aeluroidea. The average ratioof 0.93 is thus used for the estimates of limb height offered inTable 2.

The average ratios obtained for modern species were usedto estimate limb height from known track dimensions (foot-print length and glenoacetabular length) in the Salinas track-ways. Finally, we averaged the values obtained with both meth-ods, so that the possible biases in each one are compensated(Table 2).

A review of the body proportions of small and medium sizedlower Miocene aeluroid carnivores shows that they generally re-semble their extant relatives. Larger members of the Aeluroideatend to evolve more specialized body proportions, some with ex-ceedingly robust, short limbs, others with very elongated limbsand in some cases shortened backs, but the smaller species arein general more conservative (Ewer, 1973). Fossil evidence in-dicates that lowermost Miocene cats had relatively short limbs,resembling the more scansorial among extant members of thecat family.

We estimated the Froude number using the method of Alexan-der (1991). This number (v2/gl) relates speed to limb height,

TABLE 2Measurements and estimated limb height, speed and Froude numbers in representative carnivore trackways from Salinas. See

Fig. 2 for trackway keys.

Trackway R1 R2 R4 R6 I1 K3 KP

Footprint length (mm) Average 48 44 51 48 42 22 23n 36 24 10 26 58

Gleno acetabular Extremes 394/440 368/388 465/482 360/397 245/292 194/234distance (mm) Average 420 380 474 381 269 220 223Stride (mm) = λ Extremes 486/570 470/518 495/577 672/676 450/590 380/530 250/233

Average 520 500 540 735 520 440 240Limb height Ratio 1 Fl × 9.23 Fl × 9.23 Fl × 9.23 Fl × 9.23 Fl × 5.8 Fl × 9.23 Fl × 9.23

(mm) = h Results 1 443 406 470 443 244 203 212Ratio 2 Gd × 0.93 Gd × 0.93 Gd × 0.93 Gd × 0.939 Gd × 0.93 Gd × 0.93 Gd × 0.93Results 2 391 353 441 354 250 205 223Average 417 380 456 399 247 204 218

Speed (m/s) 0.731 0.763 0.701 1.371 1.348 1.276 0.429Froude number 0.131 0.156 0.110 0.481 0.751 0.814 0.086Gait Walk Walk Walk Trot Trot Trot Walk

illustrating the relative effort of an animal during locomotion.As the Froude number increases, that is, as the speed increasesrelative to the animal’s limb-height, it becomes more energeti-cally efficient for animals to “change gears” and turn from walkto trot and from trot to gallop. Ideally, the Froude number andthe speed estimations should fit with the gait reconstruction asinferred from the trackway measurements.

Error MarginsThe incorporation of average values of limb height into the

speed calculations introduces a certain margin of error. For in-stance, if we would estimate the limb height of trackway R2(Table 2) using the ratio of footprint length to limb height of theleopard (7.4), instead of the average value for carnivores with re-tractable claws, then the speed estimate would change from 0.76m/s to 0.87 m/s, and the Froude number would be 0.23 insteadof 0.16. But, if we had used the ratio of the genet (11.3) then thespeed would have been 0.66 m/s and the Froude number wouldbecome 0.66. In fact, the limb height estimates based on foot-print length ratios are consistently higher than estimates basedon glenoacetabular length in the Felipeda lynxi Panin & Avram,1962 trackways, while they are more similar in the other species,making us suspect that the trackmaker might have been robustand short-limbed like the extant leopard, so it would be temptingto drop the averages and use that ratio instead. But we have pre-ferred to use ratios as uniform as possible for the trackmakerswith retractable claws, rather than having to choose in a moreor less arbitrary way for each particular case. Thus, the speedestimates for R2 could be expressed as 0.76 m/s ± 0.1 m/s. Suchan error margin seems tolerable because both the speed estimateand the Froude number in each case would still be broadly inaccordance with the gait inferences and with other parametersof the trackway, such as footprint length-to-stride ratios.

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ClassificationIn morphological terms, the carnivore tracks from Salinas are

similar enough to previously described ichnites from other sitesas to be included in already existing ichnogenera, although inall cases the ichnites from Salinas display a much better preser-vation of detail, and the samples are much larger. In some casesthere are reasons to think that the ichnites were not producedby animals belonging to the same genus or even to the samefamily as the presumed trackmakers of the tracks that were thebasis of morphologically related ichnogenera, but such problemsare intrinsic to the method of ichnological parataxonomy. Whenstudying well-preserved ichnites that can be attributed with rea-sonable confidence to extant families of mammals, as is the casewith the Salinas tracks, it may be legitimate to object to the useof a parataxonomy, which may add more confusion than clarityto the interpretation of the fossil evidence. While many previ-ous studies of Neogene mammal tracksites have classified therespective ichnites in ichnogenera and ichnospecies (Lockleyand Meyer, 2000 and references therein), the opposite approachis exemplified in the study of vertebrate tracks from the AfricanPliocene tracksite of Laetoli (Leakey and Harris, 1987). Ear-lier studies on Salinas de Anana footprints were preliminary inadopting an informal nomenclature of the ichnites (Anton et al.,1993 and 1999). In the present study, and for the sake of easierreference, we have adopted the generally accepted criteria fornaming ichnites, assigning the Salinas carnivore tracks to twodifferent ichnogenera. The exquisite preservation of detail in theSalinas tracks warrants assignment down to the ichnospecieslevel, according to the criteria specified by Leonardi (1987).

Besides the ichnotaxonomic classification of the tracks, weattempt the attribution to known fossil taxa of the lowest possibletaxonomic level through a review of the fossil record of theCarnivora in the Miocene of Western Europe.

ReconstructionPutting together the information on track measurements, gait,

inferred body length and limb height, and hypothetical attribu-tion of the trackways to fossil taxa, we attempt a reconstructionof the trackmakers. The reconstructed animal is shown on topof a selected trackway segment, moving in the correspondinggait. The reconstruction is drawn to the same scale as the tracks,allowing us to check if the inferred body proportions and gait areconsistent with the track measurements. An incorrect inferencemight result in an impossibly or incoherently proportioned ani-mal. The classic photographic sequences of Muybridge (1957)were especially useful for checking the relationship between gaitand limb posture in walking and trotting carnivores

SYSTEMATIC ICHNOLOGY

Ichnogenus Felipeda Panin and Avram, 1962Felipeda lynxi Panin and Avram, 1962

Figs. 3, 5, 6, 7,8,12 and 13, Table 2

FIG. 6. Photograph of a section of a trackway of Felipeda lynxi. Notice a taildrag mark, which coincides with the change in direction of the trail. Parallel tothis track to the right is a Felipeda parvula trackway.

* 1962 Felipeda lynxi n. gen., n. sp.; Panin and Avram, p. 461,lam. II, fig. 15.

v 1993 Carnıvoro grande; Anton, Lopez and Santamarıa p. 23-24, figs. 2a and 2b.

v 1999 Felido R3; Anton, Lopez and Santamarıa p. 153, lam.I, figs. 1 and 2.

Number of tracks: There are ten trackways attributed to thisichnospecies, which cross the excavated area in all directions(Figs. 2, 3, 6 and 7).

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FIG. 7. Close up photograph of two footprints of Felipeda lynxi.

Description: These tracks are indistinguishable in size andmorphology from those originally described as Felipeda lynxiby Panin and Avram (1962), from the lower Miocene of theVrancea region in Romania, although the preservation of theSalinas tracks is much more detailed and of course the samplesize is incomparably larger.

The footprints average about 50 mm in length and 40 mmin width, they are paraxonic, with four digits, fused interdig-ital pads, and retracted claws (Anton et al., 1993 and 1999);the interdigital pad has a slight anterior indentation and dou-ble posterior indentation typical of extant aeluroid footprints(Fig. 8a).

Discussion: These footprints correspond to an animal aboutthe size of an extant Spanish lynx, Lynx pardinus (Temminck,1827), and generally resemble modern felid tracks, althoughthere are several morphological differences with the latter,including:

1. A relatively longer interdigital pad, meaning a less dig-itigrade stance; this difference is due to the reduction, inmodern cats, of interdigital pads 2 and 4, still well devel-oped and projecting posterior from the main pad in theSalinas tracks (Fig. 8a).

2. A greater asymmetry of some of the footprints, pointingto a more supinated posture of the feet. In this trait, theSalinas trackmaker resembles some living arboreal viver-rids such as Nandinia binotata (Reinhard, 1830), whichalso produces asymmetrical footprints (Liebenberg, 1992;Taylor, 1970, 1974).

On purely morphological grounds, these tracks could corre-spond to either a viverrid or an early hyaenid. The possibilitythat they correspond to a viverrid is currently rejected on thebasis of size. The only viverrids with retractable claws knownto have inhabited Europe are members of the subfamily Viver-rinae, such as genets or civets. No fossil or living genet everattained such large size as the species that left the larger carni-vore tracks from Salinas, and Miocene civets were also relativelysmall animals (Ginsburg, 1961). A size problem also applies forthe fossil hyaenids. The earliest members of the hyaenid fam-ily retained some degree of claw retractability, which was lostin the course of evolution, so a hyaenid of early Miocene agecould possibly make tracks morphologically similar to theseones. However, fossil hyenids of that age were far smaller thanthe lynx-sized animal that left these tracks, and their skeletonsbetray a more complete adaptation to terrestrial locomotion thanthat seen in early cats (Beaumont and Mein, 1972; Hunt andSolounias, 1991; Werdelin and Solounias, 1991) so we can ex-pect that their footprints would resemble more those of cursorialcarnivores, displaying more reduced interdigital pads.

On the other hand, footprint morphology of the Salinas track-maker matches skeletal features in the limbs of PseudaelurusGervais, 1850, the felid genus typical of the early Miocene,which retained several arboreal adaptations from some viverrid-like ancestor (Ginsburg, 1961a, b). The feet of Pseudaeluruswere less fully digitigrade than those of modern felids, and sev-eral features of the limb bones indicate a greater ability to rotatethe feet. Felid footprints of later age, such as those from theHemphillian of Death Valley in California, described as severalspecies of Bestiopeda (Felipeda) by Scrivner and Bottjer (1986),show a modern pattern, with interdigital pads 2 and 4 reducedand pressed tightly against pad 3, as in living cats.

All these data point to a small species of Pseudaelurus as theprobable trackmaker. The estimated dimensions of the limbs ofthe Salinas trackmaker are in between those of the wildcat-sizedPsedaelurus turnauensis and the lynx-sized P. lorteti. There isconsiderable size variation within, and even overlap between,the two species, so the Salinas tracks could correspond either toa large variety of the small species or a small-bodied populationof the larger taxon. (Viret, 1951; Heizmann, 1973; Ginsburget al., 1981; Ginsburg, 1999).

Felipeda parvula n. ichnosp.Figs. 3, 5, 6, 8, 9 and 12, Table 2

v 1993 Carnıvoro pequeno; Anton, Lopez and Santamarıap. 24, fig. 2h.

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FIG. 8. Pad patterns of the Salinas carnivores with reconstruction of the autopods (top row) compared to those of living carnivores (bottom row). a) The largerfelid from Salinas, Felipeda lynxi (i) has a longer interdigital pad compared to the living serval (ii). 2, 3 and 4: interdigital pad numbers. b) The herpestid footprintsfrom Salinas, Canipeda longigriffa (i) are very similar to those of the living gray mongoose (ii). c) The undetermined small aeluroid from Salinas, Felipeda parvula(i) has a shorter and more simplified interdigital pad than the living genet (ii).

Holotype: Footprint Number MCNA 1240-KP1 (Fig. 9),which is preserved as a resin cast at the Museo de CienciasNaturales de Alava (Vitoria, Spain), and provisionally exposedat the town hall of Salinas de Anana. Rubber and sand bedsin the footprint site at Salinas de Anana protect the originalspecimen.

Hypodigm: is the set of ichnites of trackway KP (Fig. 9).This set is preserved in the same resin cast (MCNA 1240) as theholotype and is also housed at Museo de Ciencias Naturales deAlava (Vitoria, Spain).

Locality: Lower Miocene (Burdigalian, = Aragonian) se-quence at Salinas de Anana, Alava (Spain).

Diagnosis: Paraxonic tracks, averaging 23 mm in length, and24 mm in width, with four toe pads and no claw marks, simpli-fied, sub-triangular, and wider than long interdigital pad, madeup of three fused pads, with slight anterior indentation. Differsfrom Felipeda lynxi in being absolutely smaller and in havinga relatively smaller, shorter interdigital pad, with pads 2 and 5projecting less posteriorly (Figs. 8c and 9).

Derivatio Nominis: parvula meaning small, for the minutesize of the ichnites.

Number of tracks: There are four trackways of this smallcarnivore, one of which (named KP) is a natural cast; the originalcorresponding trackway was never found in the site.

Discussion: Ichnites are about the size of a modern genet(Figs. 3, 6 and 9). They resemble the footprints of livingviverrines and cats (Anton et al., 1993 and 1999), according todata of Stuart and Stuart (1994), but a third possibility is thatthey correspond to an early hyenid. Living hyenids are largecarnivores with non-retractile claws, but the early Miocenehyenids were small animals resembling modern viverrines, withprimitively long bodies and partly retractile claws. These traitsare evident in the Salinas tracks, which lack claw marks andwhere footprint length is about one tenth of the glenoacetabularlength, implying small feet and a long body (Pilgrim, 1931;Hunt and Solounias, 1991).

The difficulties in attributing these tracks at the family levelcan be summarized as follows:

1. The tracks match the size and body proportions of ex-tant and fossil genets (Fig. 8c), but the reduction of theinterdigital pads suggests a more evolved, cursorial ani-mal than any living genet. The known limb anatomy ofSemigenetta repelini (Helbing, 1927), a genet of approxi-mately the same geological age as the Salinas ichnites, isremarkably similar to that of extant genets and indicatesthat the footprints would also be similar. Other species ofthe genus Semigenetta are known from partial remains, all

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FIG. 9. Photograph of trackway KP of Felipeda parvula n. ichnosp. Holotype(MCNA 1240-KP1) and Hypodigm of the ichnospecies.

indicating a gait and structure similar to that of moderngenets (Ginsburg, 1999).

2. The configuration of the footpads matches that of someliving felids, and some species of Miocene cats suchas Pseudaelurus turnauensis are almost as small as thistrackmaker. The foot anatomy of those small cats is notwell known and it is possible that they were more terres-trially adapted than their larger relatives, and thus mighthave produced footprints like these.

3. The tracks fit well with the osteology of such ictitherinehyaenids as Plioviverrops Kretzoi and Protictitherium,which include species of the right size and age (deBeaumont and Mein,1972) but ictitherines are an extinctgroup and we can only make inferences about their foot-pad structure. In brief, while the cursorial adaptations ev-idenced by the footpad reduction in these footprints maypoint to an early hyaenid as the possible trackmaker, asmall felid appears just as feasible.

Ichnogenus Canipeda Panin and Avram, 1962Canipeda longigriffa Panin and Avram, 1962

Figs. 2, 3 and 11

* 1962 Canipeda longigrifa n. gen., n. sp.; Panin and Avram,p. 461–462, lam. II, fig. 16.

v 1993 Carnıvoro mediano; Anton, Lopez and Santamarıap. 24, fig. 2c.

Number of tracks: There is one, very long trackway of thisspecies (Figs. 2, 3 and 11).

Description: Paraxonic tracks averaging 48 mm in length and33 mm in width, with four parallel oriented digit prints; simpli-fied, sub-trapezoidal, relatively large interdigital pad made upof three fused pads, with slight anterior indentation; claw marksindicating large, non retracted claws which are longer in the forethan in the hind feet (Anton et al., 1993 and 1999). Occasionallythere may be a faint print of the claw of digit 1 (Figs. 8b and 10).

Discussion: The tracks are indistinguishable in size and shapefrom a footprint described by Panin and Avram (1962) underthe name Canipedia longigriffa. While adopting the generic andspecific designations coined by those authors, we feel that thegeneric name is strongly misleading, because it unambiguouslypoints to attribution of the tracks to a member of the canid family.Such an attribution was based on the observation of superficialsimilarities, but the large size of the interdigital pad relativeto the toe pads, as well as the detailed morphology of the in-terdigital pad, clearly distinguish these tracks from true canidfootprints, and indicate that they were produced by a mongoose,an attribution which also makes sense in biogeographic terms,as commented below. So, while adopting the original denomina-tion for formal reasons, we want to emphasize that the attributionof both the Romanian and Spanish tracks to a herpestid appearsto us unambiguous.

The difference in length between the claws of the fore andhind feet evidenced in this trackway (Fig. 10) is normal in

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FIG. 10. Close-up photograph realized on two footprints of Canipeda longi-griffa. Top, left fore foot, bottom, left hind foot.

modern carnivores that have digging habits. The footprints(Fig. 8b) are remarkably similar in size and shape to those ofthe living Egyptian mongoose, Herpestes ichneumon, as clearlyobserved on data from Stuart and Stuart (1994).

Herpestid footprints cannot be confused with those of anyother aeluroid carnivore. They superficially resemble the tracksof hyenas because both have claw marks, but in the latter thetoe pads are much more closely pressed to the interdigital pad,which in turn is proportionally much smaller than in herpestids.These are to our knowledge the first data on the foot morphologyof Miocene herpestids, which hitherto were known only on thebasis of teeth and fragmentary cranial material. The foot fea-tures displayed by the Salinas tracks seem to be derived traitsamong herpestids and aeluroids in general. According to Ewer(1973), the primitive herpestid foot morphology is retained inthe living galidine Galidia elegans (Geoffroy), a semiarborealmongoose from Madagascar, with fan-like splaying digits, shortclaws, and well-separated, non-reduced interdigital pads. Lepto-plesictis, the only known herpestid genus from that age has a veryprimitive dentition, similar to that of Galidia (Petter, 1974), andsuch a primitive dentition might lead one to believe that lowerMiocene herpestids would have a similarly primitive foot pat-tern. Contrary to such expectations, the Salinas ichnites, whichvery likely correspond to Leptoplesictis, show feet of moderngrade, well adapted to terrestrial locomotion and digging.

From a palaeobiogeographical point of view, these footprintsconfirm the presence of herpestids of basically modern grade

in the lower Miocene of Europe, and strengthen the case for aPalaearctic origin of the group (Petter, 1974). Another exam-ple of the presence of herpestid tracks in the early Miocene ofEurope is a footprint from Romania, attributed to a canid and de-scribed as Canipeda longigriffa by Panin and Avram (1962). Thefootprints of canids differ strikingly from those of herpestids ascommented above, and members of the family canidae evolvedin North America and are unknown in Europe until the Turolian(Late Miocene) (Agustı and Anton, 2002).

LOCOMOTIONThe configurations of the trackways (Fig. 5) correspond with

symmetrical gaits, trots and walks in diagonal and lateral se-quences (Halfpenny, 1986). The glenoacetabular length is fairlyconstant in the long trackways (Table 2) and this indicates thatthere was little or no dorsoventral flexion of the spine duringlocomotion, as is normal in walking and trotting carnivores(English, 1980). Stride length is small to moderate in all thetrackways (Table 2), corresponding with the slow speeds of walkand trot. In three of the tracks, R6 (Felipeda lynxi) I1 (Canipedalongigriffa), and K3 (Felipeda parvula) the stride length ap-proximates a dimension twice that of the estimated limb height,agreeing with the diagonal sequence-trotting gait indicated bytrackway configuration. Froude numbers of the various track-ways are low and consistent with the gaits inferred form track-way measurements, and once again the highest figures are thoseof the three trotting trackways, R6, K3 and I1. According toAlexander (1976) mammals change gaits from walk to trot orrun at Froude numbers around 0.6, and while the figure for R8is somewhat lower, those of I1and K3 are above that limit. Allthe studied trackways from Salinas yield Froude numbers wellbelow the figure 2.6 given by Alexander (1991) for the transitionto gallop in modern mammals.

RECONSTRUCTIONThe reconstruction of the body outlines of the trackmakers

drawn over the trackways allows us to test the consistency of ourinferences about body proportions and gait (Fig. 12). Felipedalynxi is reconstructed on the basis of the anatomy of the fos-sil felid genus Pseudaelurus; Felipeda parvula was drawn afterthe proportions of the fossil hyaenid genera Plioviverrops andProtictitherium. Canipeda longigriffa was drawn after the bodyproportions of modern Herpestes because the pos-cranial skele-ton of Miocene herpestids is virtually unknown. The dental andmandibular remains of the fossil herpestid Leptoplesictis allowat least an estimate of body size, which was used as a guide forthe reconstruction of Canipeda longigriffa.

The consistency of the reconstructions with the measure-ments of the trackways is remarkable, and serves as an addi-tional test for the gait inferences and taxonomic attributions. Ascommented above, the relationship between body proportionsand footprint dimensions among the aeluroid carnivores withretractable claws displays similar variations across families, so

382 M. ANTON ET AL.

FIG. 11. Photograph of a section of trackway I1 of Canipeda longigriffa.

FIG. 12. Reconstructions of three species of carnivore trackmakers fromSalinas, drawn to the same scale relative to each other and to the trackwaysegments. Top, Felipeda lynxi, with a segment of trackway R1; center, Fe-lipeda parvula, with a segment of trackway K3; bottom Canipeda longigriffa,with a segment of trackway I1. On each trackway segment, the footprints thatcorrespond to the feet touching the ground in the reconstruction are shaded inblack.

it remains feasible that Felipeda parvula could be reconstructedas a small cat.

On the other hand, the relatively long, clawed feet of her-pestids leave little doubt as to the identity of the Canipeda longi-griffa tracks.

A further step is to attempt a realistic restoration of the track-makers. As shown for Felipeda lynxi in Fig. 13.

ECOLOGY AND BEHAVIORThe abundance of carnivores, both in number of trackways

(15) and of species (60% of the recorded mammal species),is remarkable. Given the brief time span between ichnite pro-duction and burial, it is clear that much carnivore activity wasconcentrated in a small area (about 60 m2) during a shorttime. Considering the territorial behavior and foraging habits ofcomparable modern carnivores (Ewer, 1973; Kitchener, 1991;Wemmer, 1977), and their relative scarcity in modern ecosys-tems, it seems likely that something was exerting attraction onthese animals, perhaps the presence of a carcass in the vicinity.

MIOCENE TRACKWAYS: SPAIN 383

FIG. 13. Reconstruction of trackmaker of trackway R1, Felipeda lynxi, based on the track measurements and on the anatomy of the fossil felid Pseudaelurus,with data from Ginsburg (1961a), and Turner and Anton (1997).

The need of drinking water would likely have attracted her-bivores and carnivores in a more statistical proportion (Cohenet al., 1991).

The presence of four parallel felid trackways that cross thesite in a Northwest direction (Fig. 2) is the first known exampleof group traveling in fossil felids. There are several trackwaysof other carnivores and ungulates, crossing randomly with theseparallel trails, and implying that there was no natural obsta-cle, such as the lake shore, forcing the animals to travel in thesame direction. The most conservative interpretation would bethat three adult-sized cubs were walking alongside their mother.This would indicate that, as in modern cats, cubs remained withtheir mother at least until reaching nearly adult size (Kitchener,1991).

ACKNOWLEDGEMENTSWe thank Jesus Alonso and Carmelo Corral from the Museo

de Ciencias Naturales de Alava for their support and helpduring the excavation works. Financial support on fieldworkwas provided by Museo de Ciencias Naturales de Alava. We aregrateful to Spencer Lucas and Joanna Wright for their usefulcomments on the manuscript. Lourdes Casanovas told two ofthe authors (G.L. and R.S.) about the chance finding of the siteby Petra Laumen.

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