Escaping behavior and morphological correlates in two Liolaemus species of Central Chile (Lacertilia: Iguanidae)

Summary Two central Chile lizards differ in their response to the approach of potential predators: Liolaemus lemniscatus escapes from rocky patches into grassy patches, and Liolaemus fuscus runs away from a rock onto another within the same rocky patch. These escaping responses are associated with concealing coloration and comparatively shorter legs in L. lemniscatus, as compared to L. fuscus. The adequacy of the behavior of L. lemniscatus is discussed.     

Variability in the Assessment of Snake Predation Risk by Liolaemus Lizards

The ability to assess and respond to predation risk is a strong selective force. Detection of predators is carried out by one or more sensory modalities, but the use of chemoreception has significant advantages. This study examines the chemosensorial assessment of snake predation risk and corresponding behaviours in different species and populations of Liolaemus lizards naturally exposed to different levels of snake predation pressure. The species studied were sympatric (Liolaemus lemniscatus), parapatric (L. nigroviridis) and allopatric (L. fitzgeraldi) to the saurophagous snake, Philodryas chamissonis. Additionally, two populations of L. lemniscatus from areas differing in snake densities were compared. Chemo‐assessment of predation risk was determined by comparing the number of tongue‐flicks (TF) and the behavioural responses of lizards submitted to three treatments (with semiochemicals of snake, conspecifics, and without semiochemicals – control). The results suggest that Liolaemus lizards can chemo‐assess snake predation risk, but this was modulated by the predation pressure experienced by lizards in their natural habitats. When exposed to snake semiochemicals, the sympatric prey showed less chemical exploratory behaviour (i.e. lower number of TF), a higher frequency of antipredator behaviours that would reduce its detection by a predator, and did not show the behaviour triggered by conspecific semiochemicals. The parapatric prey showed similar number of TF across different treatments, suggesting an absence of recognition of snake semiochemicals; however, it did show antipredatory behaviours when confronted with snake semiochemicals. The allopatric prey showed similar behaviour in all treatments. Both populations of the sympatric species, L. lemniscatus, showed a similar ability to detect predation risk when confronted with snake semiochemicals (i.e. similar number of TF), but antipredatory behaviours were diminished, and marking behaviours were present in the population subject to lower predation pressure. Relaxed predation pressure from a predator that releases and detects semiochemicals had similar consequences at species and population levels.     

Reciprocal trophic niche shifts in native and invasive fish: salmonids and galaxiids in Patagonian lakes

1. Rainbow (Oncorhynchus mykiss) and brown trout (Salmo trutta) are widespread and invasive salmonids with important lethal effects as predators, although indirect effects are also possible. We used stable isotope analyses (δ¹⁵N, δ¹³C) to explore how the density of invasive trout in 25 Patagonian lakes alters the trophic niche (TN) of a widespread native fish, Galaxias platei (Galaxiidae). We also explored how the density of the galaxiid influences the TN of invasive trout. 2. We quantified two aspects of the TN: (i) the proportion of littoral carbon (PL) and (ii) trophic height (TH) (i.e. the ‘height’ at which the fish feeds in the food web). We related these measures of TN in a given species to the density of other species (as estimated by catch‐per‐unit‐effort). 3. As G. platei body size increased, their PL increased (increasing littoral feeding) in several lakes. However, none of the fish species investigated showed changes in PL with increasing density of the other fish species. TH increased with body size in all three species. In addition, the TH of large G. platei declined with increasing trout density and, reciprocally, the TH of large S. trutta decreased with decreasing G. platei density. 4. The reciprocal effects of native and the invasive fish on TH were as large as a shift of one trophic level. This pattern is consistent with an exhaustion of galaxiid prey for both piscivorous G. platei and S. trutta in lakes with high trout density. 5. These finding support the suggested management strategy of culling trout from overpopulated lakes, which should simultaneously protect native fish and enhance a lucrative sport fishery for large trout.     

Cannibalism in the Andean lizard Liolaemus orientalis

Cannibalism is a recurrent behavior across the animal kingdom, with important ecological and evolutionary consequences due to its potential trade-offs on the fitness of involved individuals and demography dynamics. Cannibalism has been reported in eight species of several phylogenetic lineages within Liolaemus, a highly diverse genus of Neotropical lizards. Within the Liolaemus montanus series, two species are reported to eat juveniles in captivity. We observed a female Liolaemus orientalis, another L. montanus series member, which had ingested a juvenile of its own species in the wild. As it generally happens with other lizards, cannibalism in Liolaemus has been suggested to be performed by the larger sex, with L. chiliensis being the only other case observed until now in which the cannibalistic individual was a female. The fact that females are slightly smaller than males in L. orientalis means that our observation is an exception to previously observed trends. We suggest that there is a relationship between this behavior and the presence of a masculine trait in the cannibalistic female: developed precloacal pores. We also discuss how refuge availability may influence demography and the probability of dispersing juveniles encountering adults, increasing the chances for cannibalism to play some role in population density regulation.     

Microhabitat shifts of lizards under different contexts of sympatry: a case study with South American Liolaemus

Summary The Iguanid lizard Liolaemus tenuis is shown to be a rock and trunk dweller (apparently preferring perches between 0–30 cm height) in a central Chilean locality where it coexists with a single ground-dwelling congener. In its southern distributional ranges L. tenuis is sympatric with another tree-dweller, L. pictus. Habitat shift is demonstrated in this latter case by L. tenuis concentrating on tree trunks, and at modal heights 30–60 cm. Liolaemus pictus occupies lower (apparently more favorable) perches, actively interferring with its congener.     

Phylogeographic history of Patagonian lizards of the Liolaemus elongatus complex (Iguania: Liolaemini) based on mitochondrial and nuclear DNA sequences

In this study, we present a phylogeographic analysis of a group of lizards distributed in north‐western Patagonia, the Liolaemus elongatus complex. We sequenced 581 individuals for one mitochondrial gene (cytochrome‐b), and for a subset, we sequenced another mitochondrial gene (12S rRNA) and two nuclear genes: kinesin family member 24 (KIF24) and the anonymous nuclear locus LDAB1D. We estimated gene trees, mitochondrial and nuclear haploytpe networks, standard molecular diversity indices, genetic distances between lineages and Bayesian skyline plots. Our results provide evidence for recognition of seven species previously described within the L. elongatus complex: Liolaemus antumalguen, Liolaemus chillanensis, Liolaemus carlosgarini, Liolaemus burmeisteri, Liolaemus smaug, Liolaemus elongatus and Liolaemus crandalli, but we did not find sufficient evidence to support Liolaemus choique, Liolaemus shitan or Liolaemus sp. 6 as distinct species. We identified four candidate species (Liolaemus sp. 1, Liolaemus sp. 2, Liolaemus sp. 3 and Liolaemus sp. 7), and we discuss evolutionary processes that may have contributed to the origin of these lineages and their taxonomic and conservation implications.     

Molecular evidence for a species complex in the patagonian lizard Liolaemus bibronii and phylogeography of the closely related Liolaemus gracilis (Squamata: Liolaemini)

The lizard genus Liolaemus is endemic to temperate South America and includes 190 species. Liolaemus bibronii has a large geographic distribution and inhabits a great diversity of habitats, including the Monte, Steppe, and high Andean grassland environments. Liolaemus gracilis has a similar body size and shape to L. bibronii; the two are parapatrically distributed, and L. gracilis is also widely distributed. Here we use the mtDNA cytb sequence data of these two species to investigate lizard phylogeographic patterns in southern South America. L. bibronii is paraphyletic with respect to L. gracilis, Liolaemus ramirezae, Liolaemus robertmertensi and Liolaemus saxatilis; it is composed of many genetically different allopatric haploclades, some of which are reciprocally monophyletic. We also found evidence for introgression between L. bibronii and L. gracilis in the same area that introgression was hypothesized in the Liolaemus darwinii complex. We discuss the distribution of the major haploclades with inferences of their population histories, the concordance of these clades' distributions and histories with other lizard complexes studied with the same markers and methods, and taxonomic implications of these results.     

Multilocus phylogeography of the Patagonian lizard complex Liolaemus kriegi (Iguania: Liolaemini)

This study presents a detailed phylogeographical analysis of one of the most conspicuous groups of lizards in northwestern Patagonia, the Liolaemus kriegi complex. This region is geographically very complex as a result of Andean orogeny and subsequent volcanism coupled with a long history of glaciations and climatic changes. For 247 individuals we sequenced one mitochondrial gene (cytochrome b) and for a subset we sequenced another mitochondrial gene [12S ribosomal RNA (12S)] and two nuclear fragments [kinesin family member 24 (KIF24) and BA3 ribosomal RNA (BA3)]. We obtained gene trees and mitochondrial and nuclear haploytpe networks, and estimated genetic distances between the main lineages and basic molecular diversity indices. We also performed spatial analysis of molecular variance (SAMOVA) and Bayesian Skyline Plot (BSP) analyses, and concordant patterns from different lines of evidence permitted delimitation of seven lineages: two described species, Liolaemus buergeri and Liolaemus tregenzai; four candidate species, Liolaemus sp. A, Liolaemus sp. B, Liolaemus sp. C, and Liolaemus sp. D; and one lineage that includes all individuals from the geographical range of Liolaemus ceii and L. kriegi, referred to as L. kriegi + L. ceii. We discuss the evolutionary processes that may contribute to the origin of these lineages and their taxonomic and conservation implications. © 2014 The Linnean Society of London, Biological Journal of the Linnean Society, 2014, 113 , 256–269.     

Lizard assemblages in a fragmented landscape of central Chile

We studied lizard assemblages assessing abundance, richness, and nestedness in a fragmented landscape of central Chile including native temperate forest, forest fragments, and commercial pine plantations. Fragmentation and plantations increase the availability of edge habitats triggering both the support of additional lizard species, absent at the continuous forest, and the nestedness of lizard assemblages, where interior habitats of forest and plantations are nested subsets of habitat edges. A vulnerable lizard (Liolaemus tenuis) thrives at fragments in abundance similar to the continuous forest. Therefore, remnants ought to be considered in the conservation of lizard assemblages.     

Phylogeography between valleys and mountains: the history of populations of Liolaemus koslowskyi (Squamata,Liolaemini)

The lizard genus Liolaemus is endemic to temperate South America and includes approximately 200 species. Liolaemus koslowskyi occurs in north‐western Argentina, where it is confined to a system of interior basins and valleys. This topographically complex region is now viewed as different enough that it has been suggested for recognition as a separate zoogeographical region: The Monte Desert of Mountains and Isolated Valleys. Here we use the mtDNA cytochrome b sequence data to investigate the phylogeographical pattern of L. koslowskyi and its relationships with other species of the darwinii group. Liolaemus koslowskyi is monophyletic with respect to all the species of the darwinii group included in this analysis. Three main clades were recovered within L. koslowskyi and we hypothesized that at least one of these, which shows 7% genetic divergence, is a candidate species. We discuss the phylogeographical patterns in association with the geological history of the region. The highly structured L. koslowskyi clade suggests that it has a relatively ancient history in a topographically rich, terrestrial archipelago of habitat and tectonic islands that are themselves relictual mountains and valleys.     

Differential prey utilization by the generalist predator Coleomegilla maculata lengi according to prey size and species

Prey utilization by the generalist predator Coleomegilla maculata lengi Timb. (Coleoptera: Coccinellidae) of three sympatric lepidopterous species was quantified in relation with prey size (age) and prey species. Based on optimal foraging theory, we argued that costs associated with the utilization of small and large prey are higher than those of intermediate prey size. As a result, we expected a higher prey consumption rate on intermediate prey size leading to a convex prey consumption curve. Laboratory experiments showed that, within a given prey instar, Coleomegilla maculata lengi preyed more on Plutella xylostella (L.) compared to Artogeia rapae (L.) and Trichoplusia ni (Hübner). Generally, prey consumption rate by Coleomegilla maculata lengi on the three prey species decreased with increasing immature prey size (age). The predation efficacy of Coleomegilla maculata lengi adults and fourth instar larvae was higher compared to younger coccinellids (L2). Although, Coleomegilla maculata lengi showed a higher level of predation on smaller immature prey, we demonstrated that it is not the optimal size range for this predator. As predicted, prey weight consumption rate by Coleomegilla maculata lengi was higher at intermediate prey size leading to a convex prey utilization curve. The beneficial impact of Coleomegilla maculata lengi predation on the host plant was also estimated by using a Protection Index that considers the differential predation caused by the coccinellids and the relative importance of each pest species in terms of plant injury. Coleomegilla maculata lengi has a more significant beneficial impact when it preys on T. ni immatures.     

Body proportions,microhabitat selection,and adaptive radiation of Liolaemus lizards in central Chile

Summary A biometric analysis of body proportions with presumably functional meaning for microhabitat selection was made on 12 species of Liolaemus lizards in central Chile. Characters studied were forelimb length, hindlimb length, tail length (all standardized by the corresponding snout-vent length), and the ratio forelimb/hindlimb length. It is shown that irrespective of terrestrial, saxicolous, or arboreal habits, Liolaemus species are remarkably similar in body proportions. The only exceptions are: L. lemniscatus, an open ground-dweller which exhibits significantly shorter limbs; and L. chiliensis and L. schroederi, both shrub-climbers which exhibit significantly longer tail. It is concluded that the adaptive radiation of Liolaemus lizards in central Chile has been accomplished mainly by diversification of activity time, food size, and microhabitat type. Morphological divergence in body proportions seems to have played an unimportant role.     

Differences between European native and American invasive populations of Lythrum salicaria

Abstract. We compared the demographic characteristics of native central-European and invasive USA-populations of Lythrum salicaria growing in similar habitats. Based on the ‘Evolution of increased competitive ability’ (EICA) hypothesis, we predicted that shoot density, height, and biomass, fertility, and fecundity would be less in the European populations due to greater loss of plant material caused by exposure to control agents. Shoot density was significantly greater in most USA-populations, but shoot height and biomass were similar in invasive and native populations growing in similar habitats, especially in growing seasons with greater water availability. The number of fertile shoots was greater for invasive populations, except for those growing in sandy, nutrient-poor substrates, while percent fertility did not differ between populations growing under similar field conditions. Fecundity was also similar for populations growing in nutrient-poor and intermediate habitats, but was significantly greater in USA-populations (89–103 seeds per fruit), compared to European populations (58–64 seeds per fruit), growing in nutrient-rich habitats; seed predators were found in these European populations only. Log-linear analysis of transition frequency matrices showed that the growth of USA-populations is different from European populations, but that habitat effect was strong. Population dynamics were similar for populations growing in nutrient-poor habitats; location was of marginal importance only. USA-populations responded differently from European populations growing in intermediate and nutrient-rich habitats; differences were most pronounced between invasive (center of its North American distribution) and native populations growing in nutrient-rich habitats. The use of insect herbivores as biological control agents in North America will work best against L. salicaria populations growing in nutrient-rich habitats in the center of its invasive distribution, but will be less effective against populations growing in other habitats or portions of its range. Other factors, including nutrient- or water availability, and climate, may be as important as herbivory in affecting invasive populations; these factors interact to control L. salicaria in a more complex manner than thought previously.     

Observaciones autoecológicas en Liolaemus nitidus (Lacertilia: Iguanidae)

In an attempt to provide background information that may lead to a more comprehensive understanding of the genus Liolaemus, the paper documents some basic features of the autecology of Liolaemus nitidus. The study was carried out at two typical localities where L. nitidus is found: at the coast north of Santiago de Chile, and at the Andes ranges east of Santiago. The following parameters were investigated: microhabitat utilization, preferred body temperatures, diet, weight of gonads and of fat bodies, and frequency of regenerated tails. The main findings reported are:

1. Liolaemus nitidus is preferentially a rock‐dweller, although, when available, it can also be found perching on branches of the bromeliad Puya chilensis.

2. Preferred field body temperatures, evaluated with a quick‐registering Schultheis thermometer, indicate that L. nitidus has a preferendum of about 35.4°C. This mean preferred body temperature is not statistically different from the laboratory preferendum reported in the literature.

3. The diet of L. nitidus was found to vary with body size. Small individuals tend to eat mostly insects, whereas larger individuals tend to have more plant material in their guts. Total gut content is nevertheless relatively heavier for small than for large individuals. These facts are interpreted in the light of previous results obtained with Liolaemus species, and in relation to food availability. Regarding food availability, the above results are consistent with the hypothesis that arthropods are relatively scarce in the xerophytic areas where L. nitidus lives, and therefore these lizards complement their diet with plant material. Arthropod content of the guts was analyzed by size and taxonomic order. The findings are that mean prey size, and size of the largest and of the smallest prey in each stomach tend to be positively correlated with body size of the lizard, thus suggesting prey selectivity. On the other side, the orders most preyed upon by L. nitidus are Coleopterans, Hymenopterans (fundamentally ants), and Homopterans.

4. Correlations between either relative gonad weight or relative weight of the fat bodies, versus dry weight of the lizards are significant and positive. These results suggest that L. nitidus is a relatively K‐selected species, where there is a premium for somewhat delayed reproduction and perhaps larger clutches. Heavier fat bodies in the mountain population are interpreted in terms of length of the growing season and different needs to accumulate energy reserves for the unfavorable season.

5. Frequency of regenerated tails is greater in adults than in juveniles, but does not differ between sexes, suggesting that risk of predation would be more associated with the span of time the lizard is exposed to the environment than with differential behavior of males and females.

Results are interpreted and compared, when available, with data for other Liolaemus species.

These comparisons suggest that L. nigromaculatus and L. chiliensis, two species of similar size and ecological requirements as to L. nitidus, could be among the potentially severest competitors in limiting the geographic distribution of this species.     

On the identity of Liolaemus nigromaculatus Wiegmann,?1834 (Iguania,Liolaemidae) and?correction?of its type locality

In the current study, we review the taxonomic status of Liolaemus nigromaculatus. Despite being the nominal species of the nigromaculatus group and being the second species of the genus Liolaemus that was described, this species is of uncertain type locality and its true identification is a matter of discussion. After carefully analyzing several digital pictures of the holotype (juvenile male), reviewing all of the literature concerning the issue, examining specimens of nearly all recognized species of the nigromaculatus group, and determining the locations visited by the specimen collector, we are able to point out the following: 1) Liolaemus nigromaculatus was collected between Puerto Viejo and Copiapó of the Atacama region in Chile, and not in Huasco 2) Liolaemus bisignatus is a nomen nudum, and populations attributed to Liolaemus bisignatus should be referred to as Liolaemus nigromaculatus. 3) There is agreement that Liolaemus copiapoensis is indistinguishable from populations currently referred to as Liolaemus bisignatus (= Liolaemus nigromaculatus), 4) Populations found in Huasco (currently considered the type locality of Liolaemus nigromaculatus) are very similar to those found in Caldera (currently considered Liolaemus bisignatus) and should be designated as Liolaemus nigromaculatus, and 5) Liolaemus oxycephalus and Liolaemus inconspicuus are not synonymous with Liolaemus nigromaculatus, although their true identities are difficult to determine. We also detail several characteristic based on the holotype of Liolaemus nigromaculatus, in addition to drawing diagnostic comparisons between this species and others belonging to the nigromaculatus group.     

How robust are Palearctic migrants to habitat loss and degradation in the Sahel?

The Sahel region of West Africa is an important wintering ground for Palearctic migrants. It is, however, undergoing significant anthropogenic degradation that is likely to reduce its value in this regard. Despite there being evidence that the populations of several Palearctic species may be determined by conditions on their Sahelian wintering grounds, there are almost no baseline data on their habitat use there and so the effects that habitat change might have on their future density and distribution cannot be determined. This study measured which habitat variables were associated with the presence of Common Whitethroat Sylvia communis, Lesser Whitethroat S. curruca and Subalpine Warbler S. cantillans by carrying out point counts and habitat measurements at 1861 points at 16 sites in northern Nigeria during the winters of 2001 and 2002. Common Whitethroats were most common at intermediate tree heights and densities of Salvadora persica (with this effect strongest in the pre‐migratory period), and increased as Balanites aegyptiaca density increased initially, although this effect levelled off at higher Balanites densities. Subalpine Warblers were most common at intermediate tree heights and Balanites densities, increased as Acacia and Salvadora density increased, decreased with tree density and were least common at intermediate shrub densities. Lesser Whitethroats were most common at intermediate tree heights, Acacia and Salvadora densities, increased with Balanites density, and decreased with tree density and diversity. The shallow slopes of the significant relationships found between Palearctic warbler species and habitat parameters suggest that the group may be robust to habitat degradation, with densities remaining relatively constant across a wide range of habitat types. Common Whitethroats, in particular, appear to be able to survive in extremely degraded habitats, yet may be vulnerable to the disappearance of Salvadora, the fruit of which may allow pre‐migratory fattening.     

Thermal relationships between body temperature and environment conditions set upper distributional limits on oviparous species

We determined the thermal biology of the oviparous Liolaemus boulengeri and the viviparous Liolaemus lineomaculatus populations localised at high and low latitude sites in Patagonia, Argentina. We present data of body temperatures in the field (T_b) and preferred temperature in the laboratory (T_pref), micro-environmental and operative temperatures and the effectiveness of thermoregulation. Liolaemus boulengeri and L. lineomaculatus choose different heat sources for active selection of suitable thermal micro-environments for thermoregulation, and the oviparous L. boulengeri is a more effective thermoregulator (E=0.55) than the viviparous L. lineomaculatus (E=0.43). Even when L. boulengeri is a better thermoregulator and both species show identical timing in the reproductive cycles, there are constraint factors that impose limitations on the southernmost distribution of the oviparous L. boulengeri.     

Chemical self-recognition in the lizard Liolaemus fitzgeraldi

Social–chemical recognition is exhibited by all the Liolaemus lizards tested to date, except Liolaemus fitzgeraldi, which during post-hibernation did not discriminate chemosignals of same-sex individuals from a control. To clarify if L. fitzgeraldi is unique among the studied Liolaemus in lacking social–chemical recognition or if this was previously undetected, we recorded behavior during pre- and post-hibernation when confronted with chemosignals of conspecifics and from themselves. L fitzgeraldi showed self-recognition and seasonal changes in two exploratory behaviors. Potentially, conspecific recognition in L fitzgeraldi was undetected due to seasonality, but this species may rely comparatively less on chemical communication than congeners.