Felis lynxi, (Panin and Avram, 1962)

Abbassi, Nasrollah, Khoshyar, Masoumeh, Lucas, Spencer G. & Esmaeili, Farid, 2021, Extensive vertebrate tracksite from the Upper Red Formation (middle-late Miocene), west Zanjan, northwestern Iran, Fossil Record 24 (1), pp. 101-116 : 104-106

Felipeda lynxi ( Panin and Avram, 1962) .

Discussion

Felipeda is known as four-digit imprints of carnivore footprints, which are distinguished by their round digit imprints without claw impressions. The type specimen of Felipeda lynxi is incomplete, and it does not include complete sole imprints ( Panin and Avram, 1962, fig. 15). Thus, the assessment of Felipeda as plantigrade or digitigrade is important. Vialov (1965) introduced Bestiopeda as imprints of four digits, the rear pad 53 mm wide and 62 mm long in B. bestia (type ichnospecies), each digit imprint an elongate oval, digits close together and almost contiguous, imprint of first digit has a noticeably narrower anterior edge, and digits II and III project anteriorly in front of digits I and IV ( Sarjeant et al., 2002). Aramayo and Manera de Bianco (1987), on the other hand, described felid footprints as digitigrade and medium sized with imprints of a subtrapezoidal to subtriangular palmar pad or plant and four digital pads. They named these footprints Pumaeichnum and attributed them to the living puma ( Felis concolor ) based on size and morphology. The diagnosis of Pumaeichnum can be emended to include plantigrade and semi-plantigrade to digitigrade, obligate quadruped tetradactyl feloid footprint impressions exhibiting four digits ( II – V), each with a spheroidal to ovoidal digital pad forming a single semicircular arc in front of the interdigital pad, sole and heel impressions; digital pads are of equal or similar size, and impressions of claw tips are usually absent ( Remeika, 2001). There are no significant differences between this diagnosis and the diagnosis of Felipeda by Sarjeant et al. (2002). Thus, Pumaeichnum has been considered a junior synonym of the ichnogenus Felipeda ( Agnolin et al., 2019). Sarjeant et al. (2002) considered Pumaeichnum and Bestiopeda as junior synonyms of Felipeda.

Pantheraichnus is a large plantigrade felid footprint from the Late Pleistocene in Germany. It has four round-oval digit II – V pads imprinted in front of the large pad. In the best preserved imprint, this large pad is oval-shaped in the manus and more kidney-shaped or rectangular in the pes ( Diedrich, 2011). Nevertheless, there are no morphological differences between Pantheraichnus and Felipeda, so it is another junior synonym of Felipeda.

Mitsupes Rodríguez-de la Rosa et GuzmAEnGutiØrrez (2012) has been introduced for medium-sized felid footprints with asymmetrical metatarsal imprints. This asymmetry distinguishes this ichnogenus from the bilaterally symmetrical Felipeda.

Quiritipes Sarjeant et al. (2002) from the Eocene in Wyoming was named for semi-digitigrade to semi-plantigrade mammalian footprints of ovoidal to lanceolate outline and a small to moderate size. The pes is somewhat larger than the manus. Four digits are imprinted; all are acuminate (though not acute) without claws. The lack of claw imprints suggests a non-canid trackmaker and differs from Felipeda by size and the age of the footprints, which are older than the first occurrence of Felidae in the Oligocene. The earliest felids appeared sometime between ca. 35 Ma (age of the sister group) and 28.5 Ma (minimum age of the earliest fossils) ( Werdelin et al., 2010). The name Felipeda, however, has been used for Eocene carnivore footprints as a subichnogenus of Bestiopeda (Felipeda) by Scrivner and Bottjer (1986). Felipeda differs from Pycnodactylopus Sarjeant et al. (2002) in lacking broadly ovoid and broad digital impressions with two phalangeal pads.

Abbassi and Shakeri (2005) reported felid footprints from the URF of the Mushampa area and for the first time from Iran and assigned them to Bestiopeda isp. In contrast, Abbassi and Amini (2008) reported poorly preserved felid footprints from the URF of the Eyvanekey section in eastern Tehran as Chelipus isp. Based on morphology and the lack of claw imprints, these reports are revised as Felipeda isp.

Ichnospecies Felipeda lynxi ( Panin and Avram, 1962) Fig. 5

Specimen

Numerous footprints on the large slab, studied in the field and traced on transparent paper. There are 27 footprints arranged in six trackways measured on the lower part of the slab using transparent paper.

Diagnosis

Felidae footprint with four oval digit imprints separate from sole imprints. The footprint is 52 mm wide, and footprint length cannot be measured because the trace is not complete. The digit II and III imprints are closer to each other and have a frontal position in contrast to the lateral digits I and IV, so digit III is 3 mm forward of digit IV, and digit II is 2–3 mm forward of digit I. Digit imprints are 17 × 12 mm in dimensions. Claw imprints are absent, suggesting that they were retractable. Footprints of Felis chaus and Lynx caracal are the closest analogies to the studied footprints (from Panin and Avram, 1962; translated from Romanian).

Description

These plantigrade footprints are Felidae footprints; fourdigital impressions are spheroidal to ovoid in shape, arranged in front of the metacarpal and metatarsal pad imprints. Metacarpal and metatarsal pads show conspicuous posterior indentations. A gap separates digits II – V from the metacarpal and metatarsal pads. The imprints of digits III – IV are larger and closer to each other. No claw imprints are visible. Table 1 shows dimensions of these footprints.

Discussion

Shape of the digit imprints, morphology and size of metatarsal and/or metacarpal imprints, span between digits and between metatarsal and/or metacarpal imprints, size comparison with living felids, and symmetry or asymmetry of the footprints are the main characters for ichnospecies identification of the ichnogenus Felipeda. Based on the new combination of Bestiopeda and Pumaeichnum as Felipeda, and the newly reassigned ichnospecies, there are eight valid ichnospecies of Felipeda: F. lynxi Panin 1965 , F. biancoi Aramayo and Manera de Bianco, 1987 , F. felis Panin 1965 , F. milleri Remeika 1999 , F. scrivneri Sarjeant et al. 2002 , F. bottjeri Sarjeant et al. 2002 , F. stouti (= Pumaeichnum stouti ) ( Remeika, 1999) and F. paryula ( Anton et al., 2016) . Kordos (1985) introduced Bestiopeda maxima for mammalian carnivore footprints of great size with the sole impression proximodistally flattened with five separate toe prints that are oval in plain view. Sarjeant et al. (2002) considered this as Felipeda maxima , a new combination. Indeed, this footprint comprises five digit imprints and it should be assigned to Felipeda.

Like this combination, Sarjeant et al. (2002) reclassified Bestiopeda sanguinolenta as Felipeda sanguinolenta , but Melchor et al. (2019) recombined it as Canipeda sanguinolenta .

Bestiopeda gracilis was considered as the type ichnospecies of Chelipus by Sarjeant and Langston (1994) and attributed to the Canidae . Vialov (1966) described B. gracilis as a small tetradactyl track, 35 mm long, 29 mm wide, with digits not in contact but fanned out, and with wide digits that are sharply pointed (clawed), and he identified it as the tracks of a small canid ( Lucas, 2007). Pehuencoichnum gracilis Aramayo and Manera de Bianco (1987) has been combined with Chelipus ( Remeika, 2001) because it has a footprint structure strikingly like that of a canid rather than of a felid, as originally assigned, and it is certainly referable to Chelipus as a member of this ichnogenus. A reexamination of photos of Pehuencoichnum shows that it has claw imprints and should be assigned to Canipeda ( Melchor et al., 2019) .

Sarjeant et al. (2002) considered Bestiopeda gracilis as Felipeda gracilis in a new combination. The photo of the holotype of B. gracilis shows exactly tapered digit imprints with a small metatarsal and/or metapodium and a gap between the digit and metatarsal and/or metapodium pad imprints. Felipeda lacks claw imprints or tapered digit imprints, so B. gracilis should not be combined with Felipeda.

In the last report of vertebrate footprints from the lower part of the slab studied here, 25 of the footprints were identified as Bestiopeda isp. ( Khoshyar et al., 2016a). These footprints have spheroidal digital and metatarsal and/or metapodium pad impressions without claw imprints, so we assign them to Felipeda lynxi .

Figures

Figure 5. (a–d) Ichnospecies Felipeda lynxi photographed in the lower part of the slab of the tracksite, preserved as convex hyporelief. Scale equals millimeters (mm) in (a) and centimeters (cm) in (b)–(d).