Iberian lynx (Lynx pardinus)

Fig 1. An adult Iberian LynxTerms of use: This image is licensed under a Creative Commons Attribution 3.0 Spain License. The image is attributed to Lynx ex situ and is unedited.

Fig 1. An adult Iberian Lynx

Terms of use: This image is licensed under a Creative Commons Attribution 3.0 Spain License. The image is attributed to Lynx ex situ and is unedited.

Taxonomy

Lynx pardinus, commonly known as Iberian lynx, is a species of felid (Family Felidae) that belongs to the genus Lynx – the genus to which all the extant and extinct lynx species belong (1,2). Although initially thought to be a subspecies of Eurasian lynx (Lynx lynx), nowadays, due to morphological and molecular evidence, the Iberian lynx is considered as a distinct species (1,2). There are 2 traditionally recognized subspecies of Iberian lynx – L. p. pardinus, the extant subspecies, and L. p. spelaeus, the extinct Middle-Late Pleistocene subspecies of Iberian lynx, commonly referred to as the “cave lynx” – based on morphological characteristics (3). However, a recent study based on morphological characteristics seems to show that L. p. spelaeus is a junior synonym of L. p. pardinus (4). This hypothesis is further supported by ancient DNA data that shows that specimens previously classified as L. p. spelaeus were of the same haplotype as L. p. pardinus (5). Due to the heavily debated nature of the taxonomy of this species, we’ll use Lynx pardinus to refer to all the fossils assigned to the Iberian lynx (including L. p. spelaeus).

The putative ancestor for all the members of the Lynx lineage is the Issoire lynx (Lynx issoidorensis), which originated in Africa during the Early Pliocene (4 Mya), having subsequently spread across Eurasia (3,5,6,7). The origin of the Iberian lynx dates back to the Middle-Late Villafranchian faunal turnover (1.7-1.6 Mya), when the European population of Issoire lynx was separated due to a glacial pulse, which led to it eventually taking refuge in the Iberian Peninsula, where the Iberian lynx evolved (5,7). Despite this event being within the range established by molecular studies (7), due to the unreliable classification of the Iberian lynx fossils of the latest Villafranchian-Epivillafranchian, we cannot evaluate the impact of another event, the Early-Middle Pleistocene Transition (EMPT) (1.2-0.4 Mya) (6). The EMPT, also known as the “Mid-Pleistocene Revolution”, was a major climatic event that strongly influenced the fauna and flora, especially in the Northern Hemisphere (6,17). In his classic and comprehensive study of the evolution of lynxes, Werdelin (1981) suggested the phylogenetic lineage L. i. issoidorensisL. i. valdarnensisL. p. spelaeusL. p. pardinus as a long anagenetic (or gradual) evolutionary lineage, characterized by the decrease in size and a relative lengthening of the lower carnassial (the large upper and lower molar teeth of a carnivore, adapted for shearing flesh) (3,5,6,7). However, as it will be discussed in the Morphology and Ecology section, this hypothetical lineage is nowadays regarded as inaccurate (6).

Distribution and Age

The earliest putative remains of the Iberian lynx date back to the Early Pleistocene (1.7-1.6 Mya) to a fossil site in Spain, which is around the time suggested by molecular studies (5,6,7). Between 1.6 and 0.6 Mya, the record becomes fragmentary for members of the genus Lynx, in Southwestern Europe. In the Middle Pleistocene (0.6 Mya), fossil remains attributed to L. pardinus start to reappear in fossil sites across the Southwest of Europe (6).

Currently, the Iberian lynx can only be found in Portugal, where, since 2014, some individuals have been reintroduced, and in Spain, where the last two wild populations exist: the Doñana-Aljarafe population, in the coastal plains in the West side of the lower Guadalquivir River; and the Andújar-Cardeña population, in the East side of Sierra Morena (Fig. 2) (1,2). But, in the Pleistocene, the story was much different. The palaeodistribution of the Iberian lynx ranged from the Iberian Peninsula to the Italic Peninsula (Fig. 2) (6).

Despite having been considered Critically Endangered by the International Union for the Conservation of Nature (IUCN) for a long time, in 2015, the Iberian lynx was given the Endangered status (1,2). This historic step in the conservation of endangered felines was due to, among other things, the increase in population size in the two wild populations to around 156 mature individuals, between 2002 and 2012 (1). As of 2020, between Portugal and Spain, the population size of the Iberian lynx reached 1111 individuals, a 30% increase compared to 2019 (18), and a sign that this iconic carnivore may one day return to being a common sight in the Iberian Peninsula.

Fig 2. The extant range (Green) and estimated fossil range (Red) based on fossil remains (Ammonites) of Lynx pardinus  from Pleistocene sites (6, 19).

Fig 2. The extant range (Green) and estimated fossil range (Red) based on fossil remains (Ammonites) of Lynx pardinus from Pleistocene sites (6, 19).

Morphology and Ecology

The Iberian lynx's appearance is quite different from that of the other lynx species. The lynx species whose appearance is most similar to that of the Iberian lynx is the Eurasian lynx, which is the member of the Lynx genus genetically closest to the Iberian lynx (2). The Iberian lynx differs from the Eurasian lynx by its smaller size, and by the presence of the characteristic and prominent facial ruff, which can reach up to 12 cm, on both sexes (2). This ruff has a black fringe and is crisp white under the chin (2). Another feature that distinguishes the Iberian lynx from other lynx species is its fur. The fur of the Iberian lynx, which can range in color from this tawny-grey to reddish-brown, is the most spotted of all lynx species, with all individuals being distinctly spotted (2). However, the pattern of spots can be quite variable (2).

The extant populations of Iberian lynx are made up of individuals that weight on average 9 kg, for females, and 13 kg, for males (6). However, the extinct populations of L. pardinus appear to have had average sizes equal to or greater than those of the largest extant individuals.  The biggest specimen of L. pardinus dates back to 40 000 years ago and was found in the site of Ingarano, in Italy (6). These lynxes may have reached 25 kg, almost twice the size of the average extant Iberian lynx male (6). As stated earlier in this essay, Werdelin (1981) described a phylogenetic lineage that started with L. issoidorensis and ended with L. pardinus, which was characterized by the decrease in size and the increase in relative length of the lower carnassial (3,5,6,7). But a recent paper showed that there is no decrease in intraspecific size over time (6). For example, individuals retrieved from the fossil site of L’Escale (Middle Pleistocene - 0.6 Mya), in France, have an estimated mass of 16.9-22.0 kg, and individuals retrieved from the fossil site of Ingarano (Meghalayan Holocene - 40 000 ya), in Italy, have an estimated mass of 23.7-25.0 kg, showing that there was no decrease in size over time (6).

Much like the other two small-sized lynx species – the Canadian lynx (Lynx canadensis) and the bobcat (Lynx rufus) – the Iberian lynx is a lagomorph specialist hunter (2,6). (The bobcat is not a specialized lagomorph hunter being capable of taking down large prey such as adult white-tailed deer (Odocoileus virginianus) (2). Notwithstanding, lagomorphs can represent up to 90% of the bobcat’s diet (2,6).) The European rabbit (Oryctolagus cuniculus) is a lagomorph (Order Lagomorpha, Family Leporidae) (8,9) that evolved in the Iberian Peninsula during the Middle Pleistocene (0.6 Mya), when it starts to appear in the fossil record of Southern Spain (9). The relationship between the Iberian lynx and European rabbit is an ancient and deeply-rooted one. There is evidence that shows a close correlation of the evolution of the lynx and the evolution of the rabbit, whose geographical distributions followed the same contraction dynamics according to Pleistocene glacial-interglacial oscillations (6,7). However, we cannot assume, due to lack of data, that the Iberian lynx was always a lagomorph specialist hunter (6). Since the extant Iberian lynx populations occasionally feed on juvenile ungulates such as red deer (Cervus elaphus), fallow deer (Dama dama), and European mouflon (Ovis orientalis musimon) (10), and some size estimates point towards large body masses in some individuals, it is not unreasonable to think that the Iberian lynx could feed on larger prey during the Pleistocene (6). This is especially true if we consider the fact that these large prey overlapped with the currently known palaeodistribution of the Iberian lynx (11,12,13).

The Iberian lynx has antagonistic relationships with many small- to medium-sized carnivores, being known to often kill other predators such as red foxes (Vulpes vulpes) (10,14,15), European wildcats (Felis silvestris) (16), domestic cats (Felis catus) (10,16), domestic dogs (Canis lupus familiaris), common genets (Genetta genetta), Egyptian mongooses (Herpestes ichneumon), and Eurasian otters (Lutra lutra) (10), due to intraguild predation and/or competition. Where they overlap, the grey wolf (Canis lupus) and the Iberian lynx also compete to some extent. During the Pleistocene, it is hypothesized that the Iberian lynx and the Eurasian lynx would compete over resources in areas where they occurred in sympatry (3).

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Citations

1.       Rodríguez, A. & Calzada, J. (2015). Lynx pardinus (errata version published in 2020). The IUCN Red List of Threatened Species 2015: e.T12520A174111773. https://dx.doi.org/10.2305/IUCN.UK.2015-2.RLTS.T12520A174111773.en

2.       Hunter, L. & Barrett, P. (2015). Wild cats of the World. Bloomsbury.

3.       Werdelin, L. (1981). The evolution of lynxes. Annales Zoologici Fennici 18(1), 37-71. Retrieved August 20, 2021, from http://www.jstor.org/stable/23734102

4.       Boscaini, A., Alba, D. M., Beltrán, J. F., Moyà-Solà, S., & Madurell-Malapeira, J. (2016). Latest early Pleistocene remains of Lynx pardinus (Carnivora, Felidae) from the Iberian Peninsula: Taxonomy and evolutionary implications. Quaternary Science Reviews 143, 96–106. https://doi.org/10.1016/j.quascirev.2016.05.015

5.       Rodrı́guez-Varela, R., Tagliacozzo, A., Ureña, I., Garcı́a, N., Crégut-Bonnoure, E., Mannino, M. A., Arsuaga, J. L., Valdiosera, C. (2015). Ancient DNA evidence of Iberian lynx palaeoendemism. Quaternary Science Reviews 112, 172–180. https://doi.org/10.1016/j.quascirev.2015.01.009

6.       Mecozzi, B., Sardella, R., Boscaini, A., Cherin, M., Costeur, L., Madurell-Malapeira, J., Pavia, M., Profico, A., Iurino, D. A. (2021). The Tale of a Short-Tailed Cat: New Outstanding Late Pleistocene Fossils of Lynx pardinus from Southern Italy. Quaternary Science Reviews 262, 106840. https://doi.org/10.1016/j.quascirev.2021.106840

7.       Boscaini, A., Madurell-Malapeira, J., Llenas, M., Martı́nez-Navarro, B. (2015). The Origin of the Critically Endangered Iberian Lynx: Speciation, Diet and Adaptive Changes. Quaternary Science Reviews 123, 247-53. https://doi.org/10.1016/j.quascirev.2015.07.001

8.       Villafuerte, R. & Delibes-Mateos, M. (2019). Oryctolagus cuniculus (errata version published in 2020). The IUCN Red List of Threatened Species 2019: e.T41291A170619657. https://dx.doi.org/10.2305/IUCN.UK.2019-3.RLTS.T41291A170619657.en

9.       Lopez-Martinez, N. (2008). The lagomorph fossil record and the origin of the European rabbit. In Lagomorph Biology (pp. 27–46). Springer Berlin Heidelberg. https://doi.org/10.1007/978-3-540-72446-9_3

10.   Delibes, M., Rodrıguez, A., & Ferreras, P. (2000). Action Plan for the Conservation of the Iberian lynx (Lynx pardinus) in Europe, no. 111. Council of Europe Publishing. Nature and Environment, Strasbourg, France.

11.   Sommer, R. S., Fahlke, J. M., Schmölcke, U., Benecke, N., & Zachos, F. E. (2009). Quaternary history of the European roe deer Capreolus capreolus. Mammal Review 39(1), 1–16. https://doi.org/10.1111/j.1365-2907.2008.00137.x

12.   Sommer, R. S., Zachos, F. E., Street, M., Jöris, O., Skog, A., & Benecke, N. (2008). Late Quaternary distribution dynamics and Phylogeography of the red deer (Cervus elaphus) in Europe. Quaternary Science Reviews 27(7-8), 714–733. https://doi.org/10.1016/j.quascirev.2007.11.016

13.   Stefano, G. D., & Petronio, C. (1997). Origin and evolution of the European fallow deer (Dama, Pleistocene). Neues Jahrbuch für Geologie Und Paläontologie - Abhandlungen 203(1), 57–75. https://doi.org/10.1127/njgpa/203/1997/57

14.   Fedriani, J. M., Palomares, F., & Delibes, M. (1999). Niche relations among three sympatric Mediterranean carnivores. Oecologia 121(1), 138–148. https://doi.org/10.1007/s004420050915

15.   Sarmento, P., Bandeira, V., Gomes, P., Carrapato, C., Eira, C., & Fonseca, C. (2021). Adapt or perish: How the Iberian lynx reintroduction affects fox abundance and behaviour. Hystrix, the Italian Journal of Mammalogy 32(1), 48–54. https://doi.org/https://doi.org/10.4404/hystrix-00372-2020

16.   Nájera, F., Sánchez-Cuerda, S., López, G., Rey-Wamba, T. D., Rueda, C., Vallverdú-Coll, N., Panadero, J., Palacios, M. J., López-Bao, J. V. & Jiménez, J. (2019). Lynx eats cat: Disease risk assessment during an Iberian lynx intraguild predation. European Journal of Wildlife Research 65(3). https://doi.org/10.1007/s10344-019-1275-5

17.   Head, M. J., & Gibbard, P. L. (2005). Early-Middle Pleistocene transitions: An overview and recommendation for the defining boundary. Geological Society, London, Special Publications 247(1), 1–18. https://doi.org/10.1144/gsl.sp.2005.247.01.01

18.   Lusa. (2021, May 28). Pela primeira vez em 20 anos, há mais de 1000 linces-ibéricos. Público. https://www.publico.pt/2021/05/28/p3/noticia/primeira-20-anos-ha-1000-lincesibericos-1964416

19. Palaeobiology Database. (2021). Lynx pardinus. https://paleobiodb.org