Taxonomic Notes
This species was described as Lacerta lepida, and this name has remained widely used (including in legislation and the Bern Convention), although it has included in Timon by a number of authors since the late 20th century (e.g. Mayer and Bischoff 1996; Fu 1998, 2000; Harris et al. 1998; Harris and Carretero 2003).
Paulo (2001, 2008) suggested that the subspecies Timon lepidus nevadensis warrants recognition as a distinct species, and it was formally recognized as a distinct species by Miraldo et al. (2013) on the basis of genetic and morphological evidence and the low frequency of hybrids. Subsequent genetic and morphological research (summarized by Speybroeck et al. 2020) supports this arrangement.
Three subspecies are recognized, based on morphological, allozyme or chromosomal data.
Justification
Global and European regional assessment: Least Concern (LC)
EU 27 regional assessment: Least Concern (LC)
This European endemic species is listed as Least Concern both globally and for the EU27 Member States on the basis that, although this species has undergone range-wide declines over several decades and a number of local extinctions have been recorded, this is believed to be at a rate substantially below 30% in a 3-generation period (estimated to be approximately 12-15 years) in most of the range and the species remains abundant in the Iberian Peninsula, the core of its distribution. This species is nevertheless subject to an extremely complex suite of threats both through direct pressures on the species and its habitat and through declines in rabbit populations on which it may rely, and potentially in food resources. It is therefore important to establish monitoring, management and habitat recovery in areas where the species is at risk to ensure that it won't qualify for listing in future.
Geographic Range Information
This species is endemic to southern Europe. It is widely distributed in Portugal and Spain, being absent only from the range of the Sierra Nevada Ocellated Lizard in the southeast Iberian Peninsula, and occurring through southern France (and disjunctly along the Atlantic coast of this country) to extreme northwest Italy (Speybroeck et al. 2016). It is known in Andorra from a site in the south of the country (Amat and Roig 2003). It formerly occurred in Monaco and Gibraltar, and while now extinct in both (having been lost from Gibraltar during the 20th Century) it can still be found close to the borders (Mateo 2015). It also occurs on some Atlantic islands along the Spanish and Portuguese coasts. It ranges from as low as sea level; Speybroeck et al. (2016) report that it is "Mostly" found below 1,000 m asl, although it reaches more than 2,000 m asl in the Pyrenees (Mateo 2015).
Population Information
This species remains abundant in much of the southern Iberian Peninsula, with maximum densities in continental areas of around 60 individuals/ha in Spain and Portugal (Mateo 2015). A density as high as 16 individuals/1,200 m2, equating to 3.3 individuals/ha (Berlenga Island; Paulo 1988), is however regarded as "impressive" by Grillet et al. (2010), and pre-decline densities in the Crau area of southern France were 6.2 individuals/ha (Grillet et al. 2011). This may reflect differences in survey methods, as visual transect surveys have been found to underestimate abundance substantially and are deemed "not very useful" by Mateo (2015). The highest recorded densities are on islands: those as high as 136-208 individuals/ha have been recorded on Spanish islands (Galán 2003). The Berlenga subpopulation has since declined and Grillet et al. (2010) indicated that it had been lost before 2000, however, more recently Cheylan (2016) indicated that it survives on this island, but is on the verge of extinction.
Declines have been recorded or inferred in four range states: Portugal (Paulo 2008), Spain (Mateo 2015), France (Cheylan and Grillet 2005), and Italy (Salvidio et al. 2004). Salvidio et al. (2004) concluded from an analysis of habitat suitability that 60% of the Italian range remained suitable for the lizard. Although it remains abundant in the mainland Iberian Peninsula and several Galician islands there is a "fairly general impression" that it has undergone significant, widespread declines in abundance throughout this area over several decades, though at a lower rate than in France or Italy (summarized in Mateo 2015), and it has become very rare in several areas of Iberia (Mateo 2015). Most subpopulations appear to have been strongly declining since at least the start of the 21st Century (Grillet et al. 2010) and several northern subpopulations have been lost (Thirion et al. 2002, Cheylan and Grillet 2005), but in at least parts of the range the species has been in longer-term decline. These latter authors summarize evidence for declines and local extinctions in France that have taken place over the preceding century, and at least one French island population was lost as early as Mourgue (1930). By 2016 it survived on only two French islands: although it was abundant on Porquerolles, until at least the end of the 19th Century it was rare by the start of the 1980s (Cheylan 2016). The last confirmed record from Porquerolles was taken in 2000, and Cheylan (2016) reported its extinction following the failure to record it in an intensive two-year survey. It consequently now survives on only one French island, Oléron Island. This subpopulation has crashed in parallel with the island's rabbit population (Grillet et al. 2010). Experimental installation of artificial burrows following this population crash in 2005 appears to have successfully prevented further decline, but four years of subsequent surveys suggest that this measure has not succeeded in increasing lizard abundance (Grillet et al. 2010). A crash of over 70% between 1992 and 2009 was reported from a 30 ha study site in the Crau area of southern France even where suitable shelter sites were abundant (Grillet et al. 2010); Tatin et al. (2016) caution that this could have been a decline following a density artificially elevated by the provision of artificial shelters, which were removed shortly before the 1992 survey, and speculate that natural densities in this area are likely to have been low. These authors report on more systematic, widespread monitoring in the Crau region since 2011, recording a density of 85 adults/km2 while cautioning that their data are insufficient to determine population trends.
Habitat and Ecology Information
This species is found in open areas of woodland, scrubland, olive groves, vineyards, meadows, arable areas and sandy or rocky sites, including areas almost devoid of vegetation. It occurs on older dunes in coastal areas (Speybroeck et al. 2016) and on Pyrenean peaks, but these seem to be rare outliers rather than typical habitat for this species (Mateo 2015). In most of its range, it is a generalist, found in a range of habitats and climates with annual precipitation between 300 and more than 1,500 mm and a wide range of temperatures (Mateo 2015). Within this broad range of climates and macrostructure, it consistently appears to exhibit an association with structural complexity in vegetation and/or rocks at small scales, areas that include both sun-exposed spots for basking and retreat sites (Mateo 2015).
It is generally present in areas that have refuges such as bushes, stone walls, rabbit burrows and other holes. The ocellated lizard does not dig its own burrows (other than nests for egg laying) and due to its large size has specific refuge requirements for large, deep burrows (which are nonetheless not large enough to allow ingress for predators), within an open habitat matrix that allows sufficient foraging, basking and shade-providing herbaceous vegetation (Grillet et al. 2010). This renders it reliant on existing mammal burrows, and particularly those of animals like rabbits that make use of its favoured grassland habitats, for shelter from environmental extremes (especially in the colder north of the range) and predation pressure (Grillet et al. 2010). Its natural distribution, including available evidence of historical range expansion and contraction, closely matches that of the rabbit, suggesting that this has been an important relationship throughout the lizard's evolutionary history (Grillet et al. 2010). Individual ocellated lizards exhibit high fidelity to specific burrows (Grillet et al. 2010). It preys on insects (Grillet et al. 2010); its main prey group is beetles, but orthopterans and hymenopterans are also part of the diet (Mateo 2015, Tatin et al. 2016). It actively hunts prey and it has been suggested that dung beetles are a particularly important part of the diet, however, no particular bias towards specific beetle families seems to exist (Mateo 2015). The females lay clutches of 5 to 24 eggs between May and July (Speybroeck et al. 2016).
It has a recorded age in captivity of 14 years and 5 months (Slavens and Slavens 1992, 1993), with a report of a still-living (in 2012) zoo specimen having reached 28 years of age (Jesus 2012); wild specimens of up to 11 years have been recorded (Mateo 2015). On average, adults appear to be much shorter-lived, with the majority of the population being at most five years of age (Mateo 2015). In an unpublished study, this author recorded only 18 individuals of 389 with verifiable ages greater than 5 years, and no female lived longer than six years. There may be regional variation, with average maximum ages of 5-7 years recorded in different areas (summarized in Mateo 2015). In general sexual maturity is reached at 32-33 months, but younger ages have been reported and in one subpopulation most individuals are mature at 21 months (Mateo 2015).
Threats Information
Habitat destruction and fragmentation have been implicated in long-term declines in this species, particularly development of the Mediterranean coastline and agricultural intensification (Cheylan and Grillet 2004, Grillet et al. 2006, Mateo 2015). The latter has replaced large areas of once-suitable habitat for this lizard over the last century, including traditional plantations such as olive and almond groves with vineyards and the replacement of oak groves and Mediterranean shrubland with pine plantations (Mateo 2015). Conversely, the abandonment of traditional cattle pasture and other older forms of agricultural use has resulted in afforestation and increases in tree and shrub cover that favour the replacement of this species with the western green lizard (Mateo 2015). This has been associated with declines in Hérault and Porquelles (the latter an island where the species now appears to be extinct) in France, and Aigüesmoll de l'Emporda in Spain (Mateo 2015, Cheylan 2016). Pheasants introduced to Porquelles Island for hunting are also likely to have preyed on juvenile lizards (Cheylan 2016).
Predation by dogs is a possible impact in areas where the use of sheepdogs has increased following wolf reintroduction, such as the Crau region (Tatin et al. 2016) to which the species spread naturally following releases in the Italian Alps (J.M. Pleguezuelos pers. comm. 2022). Predation by introduced cats and dogs has been implicated in the species' decline on Berlenga Island (Portugal) and extinction on Ratonneau Island (France) (Cheylan and Grillet 2005). The Berlenga decline has also been correlated with rabbit declines (Grillet et al. 2010), and as the invasive mammals presumably preyed on both species it is unclear whether the lizard responded most strongly to the impacts of direct predation or the decline in rabbits. Egyptian Mongoose have been implicated in a significant decline in lizard abundance in parts of the Sierra Morena (Moreno 2015), although the mongoose is thought to be native to the Iberian Peninsula (Gaubert et al. 2011, Palomares 2017).
More recent, rapid declines appear to have been driven in large part by declines in populations of the common rabbit. The impact of declining rabbit populations in the north of this species' range over the decade prior to 2010 was "disastrous" for local subpopulations of this lizard, which is heavily reliant on rabbit burrows as refuges (Grillet et al. 2010). These authors suggest that, in addition to being the main provider of burrows with the specific characteristics this lizard needs, rabbit grazing may increase the suitability of grazed areas for dung beetle prey and that the mammal's decline may therefore have reduced food availability. While they expect that the lizard-rabbit association is most important in the north of the species' range, due to the shelter provided from harsh winters, the species appears to exhibit an association with rabbits throughout its range (Grillet et al. 2010). Rabbit declines are being exacerbated by agricultural intensification (Grillet et al. 2010). The massive decline recorded at one site in the Crau area of southern France has been attributed to the use of antiparasitic livestock medicine which resulted in the loss of dung beetles from this site (Grillet et al. 2010, Cheylan 2016). This interpretation has however been called into question (Tatin et al. 2016), due both to the low systematic use of these chemicals in the region and the lizard's ability to adapt to more abundant prey insects such as grasshoppers (although the latter form an irregular part of the diet and seem only to be important at high densities; Mateo 2015). The use of these chemicals has also been implicated in the decline of dung beetles in southern Spain, and would be also responsible for the decline of the Ocellated Lizard, as in this region grasshoppers are not of continuous presence along the seasons (Lobo 2001).
It is subject to persecution by local hunters and farmers based on a mistaken belief that these essentially insectivorous lizards prey on bees, partridge eggs and young rabbits (Grillet et al. 2010, Mateo 2015). This reputation resulted in a deliberate eradication campaign in the 1970s, and delayed measures to protect it in Spain (Mateo 2015). While legal persecution has ceased, in several areas poisoning and shooting is still used to kill lizards (Mateo 2015).
Climate modelling (including both this species and the Sierra Nevada ocellated lizard, since elevated to species status) suggests a contraction in the species' current potential distribution between 30 and 32% between 2041 and 2070 (Araújo et al. 2011). A model correlating current climate trends with the species' extant distribution found that, assuming no dispersal, the species would lose 8-39% of its distribution in the west and south between 2020 and 2050, although assuming dispersal it could colonize a comparable area to the north of the current range (Ceia-Hasse et al. 2014).
Use and Trade Information
This species is included in the pet trade and has been one of the most popular lizards among European hobbyists since the late 19th Century (Mateo 2015). Legal trade is presently only permitted in captive-bred specimens. Illegal collection was ongoing at least until the 1990s, and if not presently ongoing the resumption of this activity is plausible in future (Tatin et al. 2016). The species was heavily harvested for scientific collections as recently as the 1970s, but it is unlikely that this is ongoing in any significant numbers (Mateo 2015). The species has been sporadically harvested for food; although probably once widespread this is a quite localised and limited activity restricted to Extremadura, where the population is abundant enough to sustain current, low, levels of offtake (Mateo 2015).
Conservation Actions Information
Although this may be the best-studied lizard in Europe, further studies are needed in order to better-understand the scale of population declines in this species. It is a protected species in Spain, Portugal, Italy and France, but is not included in the EU Habitats Directive (Mateo 2015). It is listed on Annex II of the Bern Convention and is present in a number of protected areas, though it has nearly disappeared from a number of these. The most effective general protection measures for this species are the maintenance and restoration of structurally complex habitats (Mateo 2015). The Galician island subpopulations, which represent the endemic subspecies Timon lepidus oteroi, would benefit for management of thicket, control of cats, and the restoration of walls (Mateo 2015).
Trials using artificially constructed burrows between 2005 and 2009 found that most were rapidly occupied by lizards, particularly juveniles, indicating that this is an effective tool for improving habitat quality in areas from which rabbits have been lost or are in strong decline (Grillet et al. 2010). Further research is needed to determine whether this is a sufficient measure to prevent population decline, e.g. by investigating whether the surrounding habitat provides sufficient food resources to support viable lizard subpopulations, and whether this is effective at stemming declines over longer timescales (Grillet et al. 2010). It is likely that, at least, it is a cost-effective measure that can buffer subpopulations against short-term fluctuations in rabbit numbers (Grillet et al. 2010). Anecdotal evidence suggests that the maintenance of open, herbaceous areas in protected areas can support thriving subpopulations of both rabbits and ocellated lizards over a scale of decades, even when isolated from other parts of the range (Grillet et al. 2010). These authors suggest that mechanical site maintenance in combination with the creation of artificial burrows could be an effective, simple management technique in the absence of rabbits, potentially combined with supplementation by faecal pellets from herbivores to support beetle populations (Grillet et al. 2010). The Oléron subpopulation is subject to an established monitoring programme that conducts a census every three years to inform management and evaluate extinction risk (Grillet et al. 2011).
The species has been the target of a national action plan in France, the range state where declines appear to have been most pronounced (Cheylan 2016), since 2012. Tatin et al. (2016) stress the value of the high-profile Crau subpopulation as a model system for investigating population trends and the existence and impacts of potential threats in order to produce better-informed conservation assessments.