Neftalí Sillero

Unravelling biodiversity, evolution and threats to conservation in the Sahara‐Sahel

Deserts and arid regions are generally perceived as bare and rather homogeneous areas of low diversity. The Sahara is the largest warm desert in the world and together with the arid Sahel displays high topographical and climatic heterogeneity, and has experienced recent and strong climatic oscillations that have greatly shifted biodiversity distribution and community composition. The large size, remoteness and long‐term political instability of the Sahara‐Sahel, have limited knowledge on its biodiversity. However, over the last decade, there have been an increasing number of published scientific studies based on modern geomatic and molecular tools, and broad sampling of taxa of these regions. This review tracks trends in knowledge about biodiversity patterns, processes and threats across the Sahara‐Sahel, and anticipates needs for biodiversity research and conservation. Recent studies are changing completely the perception of regional biodiversity patterns. Instead of relatively low species diversity with distribution covering most of the region, studies now suggest a high rate of endemism and larger number of species, with much narrower and fragmented ranges, frequently limited to micro‐hotspots of biodiversity. Molecular‐based studies are also unravelling cryptic diversity associated with mountains, which together with recent distribution atlases, allows identifying integrative biogeographic patterns in biodiversity distribution. Mapping of multivariate environmental variation (at 1 km × 1 km resolution) of the region illustrates main biogeographical features of the Sahara‐Sahel and supports recently hypothesised dispersal corridors and refugia. Micro‐scale water‐features present mostly in mountains have been associated with local biodiversity hotspots. However, the distribution of available data on vertebrates highlights current knowledge gaps that still apply to a large proportion of the Sahara‐Sahel. Current research is providing insights into key evolutionary and ecological processes, including causes and timing of radiation and divergence for multiple taxa, and associating the onset of the Sahara with diversification processes for low‐mobility vertebrates. Examples of phylogeographic patterns are showing the importance of allopatric speciation in the Sahara‐Sahel, and this review presents a synthetic overview of the most commonly hypothesised diversification mechanisms. Studies are also stressing that biodiversity is threatened by increasing human activities in the region, including overhunting and natural resources prospection, and in the future by predicted global warming. A representation of areas of conflict, landmines, and natural resources extraction illustrates how human activities and regional insecurity are hampering biodiversity research and conservation. Although there are still numerous knowledge gaps for the optimised conservation of biodiversity in the region, a set of research priorities is provided to identify the framework data needed to support regional conservation planning.

Biogeographical patterns derived from remote sensing variables: the amphibians and reptiles of the Iberian Peninsula

The biogeographic patterns in species density of herptiles were analysed in the Iberian Peninsula. Geoclimatic regions were identified using a PCA. Individual habitat suitability (HS) models for 23 amphibians and 35 reptiles at 10 × 10 km scale were calculated with ENFA, using 12 environmental factors established with Remote Sensing (RS) techniques. The species presence proportion in each geoclimatic region was calculated through a cross-tabulation between each potential occurrence model and the geoclimatic regions. Species chorotypes were determined through Hierarchical Cluster Analysis using Jaccards index as association measure and by the analysis of marginality and tolerance factors from individual HS models. Predicted species density maps were calculated for each geoclimatic region. Probable under-sampled areas were estimated through differences between the predicted species density maps and observed (Gap analysis). The selected PCA components divided the Iberian Peninsula in two major geoclimatic regions largely corresponding to the Atlantic and Mediterranean climates. The Jaccards index clustered herptiles in two main taxonomic groups, with distribution similar to the Atlantic and Mediterranean geoclimatic regions (7 amphibian + 13 reptile species in three Atlantic subgroups and 16 amphibian + 22 reptile species in four Mediterranean subgroups). Marginality and tolerance factor scores identified species groups of herptile specialists and generalists. The highest observed and predicted species density areas were broadly located in identical regions. Predicted gaps are located in north-western, north-east and central Iberia. RS is a useful tool for biogeographical studies, as it provides consistent environmental data from large areas with high accuracy.

Amphibian mortality levels on Spanish country roads: descriptive and spatial analysis

Road-kills are the greatest source of direct human-induced wildlife mortality, especially in amphibians. Country roads could act as the most important source of mortality when main roads act as strong barriers hampering the migration movements of some species. Mortality patterns of amphibians on country roads (1380 km) were studied in Salamanca (Spain) in order to quantify the mortality levels, to test the effects of sex and age factors on road-kills, to determine the spatial distribution patterns of road-kills, and to identify routes of migration through a friction map and hotspots of road-kills. From a total of 819 records of amphibians, 38.1% were road-killed and 61.9% were live. Fourteen amphibian species were recorded during the surveys (10 anurans and four urodeles). The species more affected by road-kills were the anurans Bufo calamita, Pelobates cultripes and B. bufo (38.5, 23.4 and 11.9%, respectively). Females had higher incidence of road-kills than males, due to the differential activity patterns of both sexes during the reproductive period. Adults were the most common age period and also the most road-killed. The spatial distribution patterns of live and road-killed records were clustered. On the sampled roads, there were 0.23 road-kills per kilometre and 52 hotspots of road-kills. The friction map showed that most of the road-killed and live specimens were located on migration routes crossing suitable habitats. Conservation measures should be implemented in these areas, as these mortality patterns may be causing significant negative impacts at the population level.

The pond network: can structural connectivity reflect on (amphibian) biodiversity patterns?

Landscape connectivity is a very recurrent theme in landscape ecology as it is considered pivotal for the long term conservation of any organism’s populations. Nevertheless, this complex concept is still surrounded by uncertainty and confusion, largely due to the separation between structural and functional connectivity. Amphibians are the most threatened vertebrates around the globe, in Europe mostly due to habitat alteration, and to their particular life cycle. Pond breeding amphibians are considered to be organised in metapopulations, enhancing the importance of landscape connectivity in this group of animals. We sampled the amphibian species present in two pond groups in Central Western Spain. We applied the graph theory framework to these two pond networks in order to determine the importance of each pond for the entire network connectivity. We related the pond importance for connectivity with the species richness present in each pond. We tested if connectivity (partially) determined the presence of the amphibian species sampled using logistic regression. The results show that the structural connectivity of the pond network impacts on the amphibian species richness pattern and that the importance of the pond for the connectivity of the network is an important factor for the presence of some species. Our results, hence, attest the importance of (structural) landscape connectivity determining the pattern of amphibian (functional) colonization in discrete ponds.

Crocodiles in the Sahara Desert: An Update of Distribution, Habitats and Population Status for Conservation Planning in Mauritania

Background Relict populations of Crocodylus niloticus persist in Chad, Egypt and Mauritania. Although crocodiles were widespread throughout the Sahara until the early 20th century, increased aridity combined with human persecution led to local extinction. Knowledge on distribution, occupied habitats, population size and prey availability is scarce in most populations. This study evaluates the status of Saharan crocodiles and provides new data for Mauritania to assist conservation planning. Methodology/Principal Findings A series of surveys in Mauritania detected crocodile presence in 78 localities dispersed across 10 river basins and most tended to be isolated within river basins. Permanent gueltas and seasonal tâmoûrts were the most common occupied habitats. Crocodile encounters ranged from one to more than 20 individuals, but in most localities less than five crocodiles were observed. Larger numbers were observed after the rainy season and during night sampling. Crocodiles were found dead in between water points along dry river-beds suggesting the occurrence of dispersal. Conclusion/Significance Research priorities in Chad and Egypt should focus on quantifying population size and pressures exerted on habitats. The present study increased in by 35% the number of known crocodile localities in Mauritania. Gueltas are crucial for the persistence of mountain populations. Oscillations in water availability throughout the year and the small dimensions of gueltas affect biological traits, including activity and body size. Studies are needed to understand adaptation traits of desert populations. Molecular analyses are needed to quantify genetic variability, population sub-structuring and effective population size, and detect the occurrence of gene flow. Monitoring is needed to detect demographical and genetical trends in completely isolated populations. Crocodiles are apparently vulnerable during dispersal events. Awareness campaigns focusing on the vulnerability and relict value of crocodiles should be implemented. Classification of Mauritanian mountains as protected areas should be prioritised.

Spatial analysis of amphibian road mortality levels in northern Portugal country roads

Animal mortality caused by vehicle collisions is one of the main ecological impacts of roads. Amphibians are the most affected group and road fatalities have a significant impact on population dynamics and viability. Several studies on Iberian amphibians have shown the importance of country roads on amphibian road mortality, but still, little is known about the situation in northern Portugal. By being more permeable to amphibian passage, country roads represent a greater source of mortality than highways, which act as barriers. Thus, mitigation measures should be applied, but due to the extensive road network, the identification of precise locations (hotspots) and variables related to animal-vehicle collision is needed to plan these measures successfully. The aim of the study was to analyse the spatial occurrence and related factors linked to amphibian mortality on a number of country roads in northern Portugal, using spatial statistics implemented in GIS and applying a binary logistical regression. We surveyed 631 km of road corresponding to seven transects, and observed 404 individual amphibians: 74 (18.3%) alive and 330 (81.7%) road-killed. Bufo bufo represented 80% of the mortality records. Three transects showed clustered distribution of road-kills, and broadleaved forests and road ditches were the most important factors associated with hotspots of road-kill. Logistic regression models showed that habitat quality, Bufo bufo’s habitat preferences, and road ditches are positively associated with amphibians’ road mortality in northern Portugal, whereas average altitude and length of walls were negatively associated. This study is a useful tool to understand spatial occurrence of amphibian road-kills in the face of applying mitigation measures on country roads from northern Portugal. This study also considers the necessity of assessing the condition of amphibian local populations to understand their road-kills spatial patterns and the urgency to apply mitigation measures on country roads.

A place in the sun: interspecific interference affects thermoregulation in coexisting lizards

Solar radiation is in theory an unlimited resource on Earth, but can be locally limited. Heliothermic organisms use solar radiation to elevate their body temperatures, leading them to compete for sun-exposed areas, where interference for limited resource can occur. In coexisting lizard species, interference for basking sites could promote and direct interspecific interactions, which can be related to broader geographic species distribution patterns. We compared two competing lizard species (Podarcis muralis and Iberolacerta horvathi) that occupy sun-exposed gaps in a forested landscape in the Northern Dinaric Mountains (southern Europe). Both species exhibit an altitudinal segregation pattern with a high zone of overlap at middle altitudes. Using experimental thermal gradients, we tested for existence of interference competition. We used adult male lizards in three social contexts, alone as a measure of preferred body temperature (Tp) and in conspecific or heterospecific pairs as a measure of realized body temperature (Tr). I. horvathi achieved lower Tr compared to its Tp (alone), while Tr of P. muralis did not vary between social contexts. The evidence for interference found in I. horvathi could not be explained by a change in their general movement pattern in a thermal gradient. Our results suggest that interference was a result of direct agonistic interactions, assessed by the frequency of approaches and touches. I. horvathi showed higher interference-susceptibility than P. muralis indicating an asymmetric interaction between the two species. Sunlight (as a resource) is an important factor influencing distribution patterns in assemblages of heliothermic ectotherms where sun-exposed basking sites are limited.

Phylogeographic and environmental correlates support the cryptic function of the zigzag pattern in a European viper

Abstract The predator–prey relationship is a strong agent of natural selection on phenotype, and two evolutionary strategies derived from this antagonistic interaction are crypsis and aposematism. Although usually considered as opposites, both strategies could be ascribed to the dark zigzag pattern of European vipers (Vipera). Experiments using plasticine models demonstrated its aposematic role, and no evidence had been found regarding a possible cryptic function. We examined the possibility of a cryptic role by measuring five characters related to the zigzag size and shape in 465 Vipera latastei specimens from the Iberian Peninsula to assess geographic variation in these characters. This species shows genetic substructuring resulting from population isolation and occurs in strong environmental gradients, which allows testing whether historic and/or environmental (adaptive) factors explain this variation. Spatial interpolation of zigzag characters identified two major Iberian groups: the Western and the Eastern. The Western group was characterised by a larger zigzag extension and higher number of dorsal marks; specimens within this group were in granitic grounds and areas with higher rainfall and lower solar radiation than those of the Eastern group. The correlation of the zigzag shape and size with lithology and climatic variables suggested that dorsal pattern variation is driven by: (1) its cryptic role, as detectability might be influenced by the degree of contrast between the target and background lithology, or (2) its thermal role, as the larger zigzag may allow for faster heating in Western Iberian regions with limited thermal opportunities. A log-linear analysis using dorsal pattern groups, genetic lineages and lithological classes, showed significant interactions among the three variables. These results suggest that dorsal pattern variation of V. latastei resulted from genetic (i.e. historic) as well as environmental (i.e. adaptive) factors, first by population isolation in geographic refuges and further by local adaptation to particular environments.

Snakes on the Balearic Islands: An Invasion Tale with Implications for Native Biodiversity Conservation

Biological invasions are a major conservation threat for biodiversity worldwide. Islands are particularly vulnerable to invasive species, especially Mediterranean islands which have suffered human pressure since ancient times. In the Balearic archipelago, reptiles represent an outstanding case with more alien than native species. Moreover, in the last decade a new wave of alien snakes landed in the main islands of the archipelago, some of which were originally snake-free. The identification of the origin and colonization pathways of alien species, as well as the prediction of their expansion, is crucial to develop effective conservation strategies. In this study, we used molecular markers to assess the allochthonous status and the putative origin of the four introduced snake species (Hemorrhois hippocrepis, Malpolon monspessulanus, Macroprotodon mauritanicus and Rhinechis scalaris) as well as ecological niche models to infer their patterns of invasion and expansion based on current and future habitat suitability. For most species, DNA sequence data suggested the Iberian Peninsula as the potential origin of the allochthonous populations, although the shallow phylogeographic structure of these species prevented the identification of a restricted source-area. For all of them, the ecological niche models showed a current low habitat suitability in the Balearic, which is however predicted to increase significantly in the next few decades under climate change scenarios. Evidence from direct observations and spatial distribution of the first-occurrence records of alien snakes (but also lizards and worm lizards) suggest the nursery trade, and in particular olive tree importation from Iberian Peninsula, as the main pathway of introduction of alien reptiles in the Balearic islands. This trend has been reported also for recent invasions in NE Spain, thus showing that olive trees transplantation may be an effective vector for bioinvasion across the Mediterranean. The combination of molecular and ecological tools used in this study reveals a promising approach for the understanding of the complex invasion process, hence guiding conservation management actions.

Spatial Biodiversity Patterns of Madagascar's Amphibians and Reptiles.

Madagascar has become a model region for testing hypotheses of species diversification and biogeography, and many studies have focused on its diverse and highly endemic herpetofauna. Here we combine species distribution models of a near-complete set of species of reptiles and amphibians known from the island with body size data and a tabulation of herpetofaunal communities from field surveys, compiled up to 2008. Though taxonomic revisions and novel distributional records arose since compilation, we are confident that the data are appropriate for inferring and comparing biogeographic patterns among these groups of organisms. We observed species richness of both amphibians and reptiles was highest in the humid rainforest biome of eastern Madagascar, but reptiles also show areas of high richness in the dry and subarid western biomes. In several amphibian subclades, especially within the Mantellidae, species richness peaks in the central eastern geographic regions while in reptiles different subclades differ distinctly in their richness centers. A high proportion of clades and subclades of both amphibians and reptiles have a peak of local endemism in the topographically and bioclimatically diverse northern geographic regions. This northern area is roughly delimited by a diagonal spanning from 15.5°S on the east coast to ca. 15.0°S on the west coast. Amphibian diversity is highest at altitudes between 800–1200 m above sea-level whereas reptiles have their highest richness at low elevations, probably reflecting the comparatively large number of species specialized to the extended low-elevation areas in the dry and subarid biomes. We found that the range sizes of both amphibians and reptiles strongly correlated with body size, and differences between the two groups are explained by the larger body sizes of reptiles. However, snakes have larger range sizes than lizards which cannot be readily explained by their larger body sizes alone. Range filling, i.e., the amount of suitable habitat occupied by a species, is less expressed in amphibians than in reptiles, possibly reflecting their lower dispersal capacity. Taxonomic composition of communities assessed by field surveys is largely explained by bioclimatic regions, with communities from the dry and especially subarid biomes distinctly differing from humid and subhumid biomes.