Should I stay or move? Quantifying landscape of fear to enhance environmental management of road networks in a highly transformed landscape.

The development and expansion of road networks pose considerable threats to natural habitats and wildlife, fostering a landscape of fear. In addition to direct mortality caused by road collisions, road construction and maintenance often result in habitat fragmentation and loss, impeding animal movement and gene flow between populations. Mountain ungulates are already confined to fragmented habitat patches and roads can cause substantial disturbances to their critical ecological processes, such as dispersal and migration. In this study, we employed two key mountain ungulates, the wild goat (Capra aegagrus) and mouflon (Ovis gmelini), as functional models to examine how road networks impact the quantity and connectivity of natural habitats in southwestern Iran, where extensive road construction has led to significant landscape changes. We used the MaxEnt method to predict species distribution, the circuit theory to evaluate habitat connectivity, and the Spatial Road Disturbance Index (SPROADI) to assess road impacts. During the modeling process, we selected eleven important variables and employed a model parametrization strategy to identify the optimal configuration for the MaxEnt model. For SPROADI index we used three sub-indices, including traffic intensity, vicinity impact, and fragmentation grade. We then integrated the results of these analyses to identify areas with the most significant environmental impacts of roads on the coherency of the natural habitats. The findings indicate that suitable habitats for wild goats are widely distributed across the study area, while suitable habitats for mouflon are primarily concentrated in the northeastern region. Conservation gap analysis revealed that only 8% of wild goat habitats and 7% of mouflon habitats are covered by protected areas (PAs). The SPROADI map highlighted that 23% of the study area is negatively influenced by road networks. Moreover, 30.4% of highest-probability corridors for mouflon, and 25.7% for wild goat, were highly vulnerable to the impacts of roads. Our combined approach enabled us to quantitatively assess species-specific vulnerability to the impacts of heavy road networks. This study emphasizes the urgent need to address the negative effects of road networks on wildlife habitats and connectivity corridors. Our approach effectively identifies sensitive areas, which can help inform mitigation strategies and support more effective conservation planning in significantly transformed landscapes.

Body size and digestive system shape resource selection by ungulates: A cross-taxa test of the forage maturation hypothesis.

The forage maturation hypothesis (FMH) states that energy intake for ungulates is maximised when forage biomass is at intermediate levels. Nevertheless, metabolic allometry and different digestive systems suggest that resource selection should vary across ungulate species. By combining GPS relocations with remotely sensed data on forage characteristics and surface water, we quantified the effect of body size and digestive system in determining movements of 30 populations of hindgut fermenters (equids) and ruminants across biomes. Selection for intermediate forage biomass was negatively related to body size, regardless of digestive system. Selection for proximity to surface water was stronger for equids relative to ruminants, regardless of body size. To be more generalisable, we suggest that the FMH explicitly incorporate contingencies in body size and digestive system, with small-bodied ruminants selecting more strongly for potential energy intake, and hindgut fermenters selecting more strongly for surface water.

Endozoochorous dispersal by herbivores and omnivores is mediated by germination conditions.

BACKGROUND: Vertebrate-mediated seed dispersal is probably the main long distance dispersal mode. Through endozoochory, large mammals act as mobile links between habitats within and among forest patches. Along with other factors, their feeding regimes do affect their contribution as dispersal vectors. We conducted a cross-species comparative experiment involving two herbivores, red deer and roe deer; and two opportunistic omnivores, wild boar and brown bear, all occurring in the forest and steppe-forest ecotone habitats of the south-eastern Caspian region. We compared their role as endozoochorous seed dispersal agents by monitoring seedling emergence in their dungs under greenhouse and natural conditions. RESULTS: In total, 3078 seedlings, corresponding to 136 plant taxa sprouted from 445 paired dung sub-samples, under greenhouse and natural conditions. Only 336 seedlings, corresponding to 36 plant taxa, emerged under natural conditions, among which five taxa did not appear under greenhouse conditions. Graminoids and forbs composed 91% of the seedlings in the greenhouse whereas shrubs were more abundant under natural conditions, representing 55% of the emerged seedlings. Under greenhouse conditions, first red deer and then wild boar dispersed more species than the other two mammals, while under natural conditions brown bear was the most effective vector. We observed remarkably higher species richness and seedling abundance per dung sub-sample under buffered greenhouse conditions than we did under natural conditions. CONCLUSIONS: The four sympatric mammals studied provided different seed dispersal services, both in terms of seedling abundance and species richness and may therefore be regarded as complementary. Our results highlight a positive bias when only considering germination under buffered greenhouse conditions. This must be taken into account when planning management options to benefit plant biodiversity based on the dispersal services concluded from greenhouse experiments.

Bioaccumulation of heavy metals (Hg, Cd and Ni) by sentinel crab (Macrophthalmus depressus) from sediments of Mousa Bay, Persian Gulf.

The presence of heavy metals (HMs) in the environment can increase their risk of transfer to the food chain. The present study was conducted to investigate the bioaccumulation of HMs including Hg, Cd and Ni in soft and hard tissues of sentinel crab (Macrophthalmus depressus Rüppell, 1830) in sediments of Mousa Bay in northwest of the Persian Gulf. The average (mean ± SD) amounts of Hg, Cd and Ni in the sediments were 6.27 ± 3.66, 3.8 ± 1.48 and 71.6 ± 5.09 µg/g, whereas the relevant amounts for soft tissues were 6.16 ± 4.64, 3.3 ± 1.7 and 19.7 ± 3.96 µg/g, and for hard tissues were, 2.9 ± 1.67, 3.5 ± 1.35 and 10.44 ± 3.1 µg/g, respectively. Accordingly, soft tissues of the crab could be suitable bioindicators of Hg and Cd, whereas the hard tissues could be used as Cd bioindicator. Since the bioaccumulation of Ni was the most, it might have a higher risk among the metals to enter the food chain in the region.

Phylogeny and species delimitation of near Eastern Neurergus newts (Salamandridae) based on genome-wide RADseq data analysis.

We reconstruct the molecular phylogeny of Near Eastern mountain brook newts of the genus Neurergus (family Salamandridae) based on newly determined RADseq data, and compare the outcomes of concatenation-based phylogenetic reconstruction with species-tree inference. Furthermore, we test the current taxonomy of Neurergus (with four species: Neurergus strauchii, N. crocatus, N. kaiseri, and N. derjugini) against coalescent-based species-delimitation approaches of our genome-wide genetic data set. While the position of N. strauchii as sister species to all other Neurergus species was consistent in all of our analyses, the phylogenetic relationships between the three remaining species changed depending on the applied method. The concatenation approach, as well as quartet-based species-tree inference, supported a topology with N. kaiseri as the closest relative to N. derjugini, while full-coalescent species-tree inference approaches supported N. crocatus as sister species of N. derjugini. Investigating the individual signal of gene trees highlighted an extensive variation among gene histories, most likely resulting from incomplete lineage sorting. Coalescent-based species-delimitation models suggest that the current taxonomy might underestimate the species richness within Neurergus and supports seven species. Based on the current sampling, our analysis suggests that N. strauchii, N. derjugini and N. kaiseri might each be subdivided into further species. However, as amphibian species are known to be composed of deep conspecific lineages that do not always warrant species status, these results need to be cautiously interpreted in an integrative taxonomic framework. We hypothesize that the rather shallow divergences detected within N. kaiseri and N. derjugini likely reflect an ongoing speciation process and thus require further investigation. On the contrary, the much deeper genetic divergence found between the two morphologically and geographically differentiated subspecies of N. strauchii leads us to propose that N. s. barani should be considered a distinct species, Neurergus barani Öz, 1994.

Thermal vulnerability of sea turtle foraging grounds around the globe.

Anticipating and mitigating the impacts of climate change on biodiversity requires a comprehensive understanding on key habitats utilized by species. Yet, such information for high mobile marine megafauna species remains limited. Here, we compile a global database comprising published satellite tracking data (n = 1035 individuals) to spatially delineate foraging grounds for seven sea turtle species and assess their thermal stability. We identified 133 foraging areas distributed around the globe, of which only 2% of the total surface is enclosed within an existing protected area. One-third of the total coverage of foraging hotspots is situated in high seas, where conservation focus is often neglected. Our analyses revealed that more than two-thirds of these vital marine habitats will experience new sea surface temperature (SST) conditions by 2100, exposing sea turtles to potential thermal risks. Our findings underline the importance of global ocean conservation efforts, which can meet climate challenges even in remote environments.

MaxEnt brings comparable results when the input data are being completed; Model parameterization of four species distribution models.

Species distribution models (SDMs) are practical tools to assess the habitat suitability of species with numerous applications in environmental management and conservation planning. The manipulation of the input data to deal with their spatial bias is one of the advantageous methods to enhance the performance of SDMs. However, the development of a model parameterization approach covering different SDMs to achieve well-performing models has rarely been implemented. We integrated input data manipulation and model tuning for four commonly-used SDMs: generalized linear model (GLM), gradient boosted model (GBM), random forest (RF), and maximum entropy (MaxEnt), and compared their predictive performance to model geographically imbalanced-biased data of a rare species complex of mountain vipers. Models were tuned up based on a range of model-specific parameters considering two background selection methods: random and background weighting schemes. The performance of the fine-tuned models was assessed based on recently identified localities of the species. The results indicated that although the fine-tuned version of all models shows great performance in predicting training data (AUC > 0.9 and TSS > 0.5), they produce different results in classifying out-of-bag data. The GBM and RF with higher sensitivity of training data showed more different performances. The GLM, despite having high predictive performance for test data, showed lower specificity. It was only the MaxEnt model that showed high predictive performance and comparable results for identifying test data in both random and background weighting procedures. Our results highlight that while GBM and RF are prone to overfitting training data and GLM over-predict nonsampled areas MaxEnt is capable of producing results that are both predictable (extrapolative) and complex (interpolative). We discuss the assumptions of each model and conclude that MaxEnt could be considered as a practical method to cope with imbalanced-biased data in species distribution modeling approaches.

The influence of superstitions and emotions on villagers' attitudes towards striped hyena in southwestern Iran.

The intensity of human-carnivore conflict in socio-ecological systems may primarily be determined by people's attitudes and perceptions of carnivore-related threats. Direct or indirect threats posed by large carnivores to human interests may eventually lead to negative attitudes that can trigger retaliatory bahaviour against them. We studied local people's attitudes towards striped hyena (Hyaena hyaena), the nature and extent of the human-hyena conflict, and the socio-cultural drivers of the conflicts in 19 rural communities in southwestern Iran. We employed structural equation modelling to assess socio-cultural factors affecting attitudes towards striped hyenas. The findings of 300 interviews showed significant differences in local people's superstitious attitudes regarding gender, age, and education. More than 40% of the participants had encountered hyenas, and on average, each respondent lost 0.44 livestock in the past five years due to hyena attacks. However, livestock depredation by the hyena was low (13.3%) compared to the damage inflicted by all carnivores (73%). While the respondents indicated some degrees of fear, hatred to hyena was relatively low and they generally showed positive attitudes towards the species. Women and older people expressed the highest and respondents with higher education the least superstitious beliefs. Attitude score of respondents toward hyenas was correlated negatively with hatred for hyenas and positively with knowledge about them, but socio-demographics effects on attitudes towards hyenas were not statistically significant. Self-reported livestock loss was a relatively good predictor of hatred and fear. Herders who had not protected their livestock reported carnivore attacks at least once. We conclude that superstitions can potentially negatively affect hyena persistence, but can be reduced by improving the educational level of local people.

Ecological niche models reveal divergent habitat use of Pallas's cat in the Eurasian cold steppes.

Identifying the association between the patterns of niche occupation and phylogenetic relationships among sister clades and assisting conservation planning implications are of the most important applications of species distribution models (SDMs). However, most studies have been carried out regardless of within taxon genetic differentiation and the potential of local adaptation occurring within the species level. The Pallas's cat (Otocolobus manul) is a less-studied species with unknown biogeography and phylogenetic structure across a widespread yet isolated range from the Caucasus to eastern China. In the current study, by considering a previously proposed genetic structure and based on a cluster analysis on climatic variables, we supposed three clades for this species, including O. m. manul, O. m. ferrugineus, and O. m. nigripectus. We developed SDM for each clade separately and compared it with a general distribution model of the species to determine whether the hypothesized taxonomic resolution affects the predicted ecological niche of the within-species structures. We assessed the effect of climate change on the future distribution of the species to detect the most sensitive clades to global warming scenarios. Our results showed that for all clades' models, the AUC and TSS were greater than the general model. Access to the preferred prey of the Pallas's cat, that is, pika, had a significant effect on the distribution of O. m. manul and O. m. ferrugineus, whereas the most influential variable affecting O. m. nigripectus habitat suitability was terrain slope. Based on our future projections, we found that future climate change likely threatens the clades O. m. ferrugineus and O. m. nigripectus more than O. m. manul, findings that were hidden in the general model. Our results highlight the proficiency of SDMs in recognizing within-taxon habitat use of widespread species and the necessity of this procedure for implementing effective conservation planning of these species.

Phylogeographic and phenotypic divergence between two subspecies of Testudo graeca (T. g. buxtoni and T. g. zarudnyi) across their contact zone in Iran.

Contact zones are considered as windows into the evolutionary process, allowing identification of factors influencing the evolutionary forces. Here, we combined phylogenetic and morphometric analyses to explore the evolutionary process affecting the taxonomic pattern of two subspecies of Testudo graeca (T. g. buxtoni and T. g. zarudnyi) across their contact zone in Central Iran. Our results showed high levels of phylogeographic and phenotypic variation in the contact zone. Two monophyletic clades including, clade 1 (T. g. zarudnyi) and clade 2 (T. g. buxtoni) were identified. Furthermore, four distinct subclades were found in T. g. buxtoni, across a wide geographic range. Divergence time analysis suggests that the two subspecies diverged from one another after the uplifting of the Zagros Mountains during the early Pliocene. Using neutrality tests and mismatch distribution analysis, we found no evidence of recent population expansion. Morphological associations among geographical populations in the contact zone found more distinctions, with some significant adaptive and non-adaptive morphological variations in these populations. These distinctive morphological populations can be considered as management units (MUs) to conserve the evolutionary potential of this species. Finer scale evolutionary studies are required to address the southern part of the Zagros mountain range, where the overlapping of mitochondrial clades and subclades has occurred. Such information is essential for effective conservation of T. graeca populations, preventing translocation or mixing of individuals without comprehensive genetic and morphological assessment.

Species versus within-species niches: a multi-modelling approach to assess range size of a spring-dwelling amphibian.

Species Distribution Models (SDMs) can be used to estimate potential geographic ranges and derive indices to assess species conservation status. However, habitat-specialist species require fine-scale range estimates that reflect resource dependency. Furthermore, local adaptation of intraspecific lineages to distinct environmental conditions across ranges have frequently been neglected in SDMs. Here, we propose a multi-stage SDM approach to estimate the distributional range and potential area of occupancy (pAOO) of Neurergus kaiseri, a spring-dwelling amphibian with two climatically-divergent evolutionary lineages. We integrate both broad-scale climatic variables and fine-resolution environmental data to predict the species distribution while examining the performance of lineage-level versus species-level modelling on the estimated pAOO. Predictions of habitat suitability at the landscape scale differed considerably between evolutionary level models. At the landscape scale, spatial predictions derived from lineage-level models showed low overlap and recognised a larger amount of suitable habitats than species-level model. The variable dependency of lineages was different at the landscape scale, but similar at the local scale. Our results highlight the importance of considering fine-scale resolution approaches, as well as intraspecific genetic structure of taxa to estimate pAOO. The flexible procedure presented here can be used as a guideline for estimating pAOO of other similar species.

The legacy of Eastern Mediterranean mountain uplifts: rapid disparity of phylogenetic niche conservatism and divergence in mountain vipers.

BACKGROUND: The orogeny of the eastern Mediterranean region has substantially affected ecological speciation patterns, particularly of mountain-dwelling species. Mountain vipers of the genus Montivipera are among the paramount examples of Mediterranean neo-endemism, with restricted ranges in the mountains of Anatolia, the Levant, Caucasus, Alborz, and Zagros. Here we explore the phylogenetic and ecological diversification of Montivipera to reconstruct its ecological niche evolution and biogeographic history. Using 177 sequences of three mitochondrial genes, a dated molecular phylogeny of mountain vipers was reconstructed. Based on 320 occurrence points within the entire range of the genus and six climatic variables, ecological niches were modelled and used to infer ancestral niche occupancy. In addition, the biogeographic history and ancestral states of the species were reconstructed across climate gradients. RESULTS: Dated phylogenetic reconstruction revealed that the ancestor of mountain vipers split into two major clades at around 12.18 Mya followed by multiple vicariance events due to rapid orogeny. Montivipera colonised coastal regions from a mountain-dwelling ancestor. We detected a highly complex ecological niche evolution of mountain vipers to temperature seasonality, a variable that also showed a strong phylogenetic signal and high contribution in niche occupation. CONCLUSION: Raising mountain belts in the Eastern Mediterranean region and subsequent remarkable changes in temperature seasonality have led to the formation of important centres of diversification and endemism in this biodiversity hotspot. High rates of niche conservatism, low genetic diversity, and segregation of ranges into the endemic distribution negatively influenced the adaptive capacity of mountain vipers. We suggest that these species should be considered as evolutionary significant units and priority species for conservation in Mediterranean mountain ecosystems.

Morphological diversity and ecological niche divergence in goitered and sand gazelles.

The phylogeny and species boundaries of Gazella subgutturosa and G. marica have been long debated. The achievements of past conservation efforts have been compromised by a lack of knowledge about the phylogeny and taxonomic status of different populations. We integrated the recent genetic findings by previous studies with morphometric analyses and ecological niche modeling (ENM) to assess discreteness among populations of these gazelle species in Asia. Taxonomic diversity of gazelles was investigated by using principal components analysis (PCA) based on 14 cranial measures of male skulls. Ecological niche divergence was examined based on a PCA on climatic factors and a species distribution modeling (SDM) with environmental variables. Morphometric results indicated substantial differentiation in size between skulls of the western Zagros Mountains including west and south-western Iran and Arabian Peninsula from all other samples east of the Zagros Mountains from Iran to China. ENM also revealed that gazelles in the east and west of Zagros Mountains occupy distinct niches and that there are apparent areas of disconnection across the goitered gazelle suitable range. A complete divergent niche occupation was also observed between goitered gazelles of northern Mongolia and other populations of the species, except those in China. Taking the inferences from ENM and morphology together with previous genetics results, we conclude that gazelles in the west and south-west of Iran may represent G. marica. Also, our combined analyses revealed divergence among gazelles of Iran, Central Asia, and Mongolia/China. These results may pave the way for future studies and have conservation implications particularly for reintroduction/supplementation programs.

Human dimensions of wildlife conservation in Iran: Assessment of human-wildlife conflict in restoring a wide-ranging endangered species.

Human-wildlife conflicts restrict conservation efforts, especially for wide-ranging animals whose home ranges overlap with human activities. We conducted a study to understand conflicts with, and factors influencing the perceived value of an expanding population of onagers (Equus hemionus onager) in local communities in southern Iran. We asked about locals' perceptions of six potential management strategies intended to lessen human-onager conflict. We found that human-onager conflict was restricted to 45% of respondents within the Bahram-e-Goor Protected Area, all of whom were involved in farming or herding activities. Locals within the protected area were more knowledgeable about onagers and valued onagers more than those living outside the protected area. The perceived value of onagers increased with level of education, total annual income, and perceptions of onager population trends; the perceived value of onagers decreased with the magnitude of conflict between onagers and locals. To tolerate or avoid conflicts with onagers, locals were supportive of monetary compensation and changing from a traditional lifestyle to industrialized farming (for farmers) or livestock production (for herders) with the help of government; locals did not support selling land to the government. Our study is among the first in human-wildlife conflict and local attitudes towards an endangered species and its recovery in Iran. We conclude that current levels of human-onager conflict are relatively low and perceived value of onagers is still relatively high. Therefore, wildlife authorities should consider the development of mitigation strategies with local communities before conflicts intensify.

Extinction risks of a Mediterranean neo-endemism complex of mountain vipers triggered by climate change.

Climate change is among the most important drivers of biodiversity decline through shift or shrinkage in suitable habitat of species. Mountain vipers of the genus Montivipera are under extreme risk from climate changes given their evolutionary history and geographic distribution. In this study, we divided all Montivipera species into three phylogenetic-geographic Montivipera clades (PGMC; Bornmuelleri, Raddei and Xanthina) and applied an ensemble ecological niche modelling (ENM) approach under different climatic scenarios to assess changes in projected suitable habitats of these species. Based on the predicted range losses, we assessed the projected extinction risk of the species relative to IUCN Red List Criteria. Our result revealed a strong decline in suitable habitats for all PGMCs (63.8%, 79.3% and 96.8% for Xanthina, Raddei and Bornmuelleri, respectively, by 2070 and under 8.5 RCP scenario) with patterns of altitudinal range shifts in response to projected climate change. We found that the mountains close to the Mediterranean Sea are exposed to the highest threats in the future (84.6 ± 9.1 percent range loss). We also revealed that disjunct populations of Montivipera will be additionally highly isolated and fragmented in the future. We argue that leveraging climate niche projections into the risk assessment provides the opportunity to implement IUCN criteria and better assess forthcoming extinction risks of species.

Geographic separation and genetic differentiation of populations are not coupled with niche differentiation in threatened Kaiser's spotted newt (Neurergus kaiseri).

The combination of niche modelling and landscape genetics (genomics) helps to disentangle processes that have shaped population structure in the evolutionary past and presence of species. Herein, we integrate a comprehensive genomic dataset with ecological parameters and niche modelling for the threatened Kaiser's newt, a newt species adapted to mountain spring-ponds in Iran. Genomic analysis suggests the existence of two highly differentiated clades North and South of the Dez River. Genetic variation between the two clades (76.62%) was much greater than within clades (16.25%), suggesting that the Dez River prevented gene flow. River disconnectivity, followed by geographic distance, contributed mostly to genetic differentiation between populations. Environmental niche and landscape resistance had no significant influence. Though a significant difference between climatic niches occupied by each clade at the landscape-scale, habitat niches at the local-scale were equivalent. 'Niche similarity analysis' supported niche conservatism between the two clades despite the southward shift in the climatic niche of the Southern clade. Accordingly, populations of different clades may occupy different climatic niches within their ancestral niche. Our results indicate that the change of climatic conditions of geographically and genetically separated populations does not necessarily result in the shift of an ecological niche.

Taming the late Quaternary phylogeography of the Eurasiatic wild ass through ancient and modern DNA.

Taxonomic over-splitting of extinct or endangered taxa, due to an incomplete knowledge of both skeletal morphological variability and the geographical ranges of past populations, continues to confuse the link between isolated extant populations and their ancestors. This is particularly problematic with the genus Equus. To more reliably determine the evolution and phylogeographic history of the endangered Asiatic wild ass, we studied the genetic diversity and inter-relationships of both extinct and extant populations over the last 100,000 years, including samples throughout its previous range from Western Europe to Southwest and East Asia. Using 229 bp of the mitochondrial hypervariable region, an approach which allowed the inclusion of information from extremely poorly preserved ancient samples, we classify all non-African wild asses into eleven clades that show a clear phylogeographic structure revealing their phylogenetic history. This study places the extinct European wild ass, E. hydruntinus, the phylogeny of which has been debated since the end of the 19th century, into its phylogenetic context within the Asiatic wild asses and reveals recent mitochondrial introgression between populations currently regarded as separate species. The phylogeographic organization of clades resulting from these efforts can be used not only to improve future taxonomic determination of a poorly characterized group of equids, but also to identify historic ranges, interbreeding events between various populations, and the impact of ancient climatic changes. In addition, appropriately placing extant relict populations into a broader phylogeographic and genetic context can better inform ongoing conservation strategies for this highly-endangered species.