Predictive modelling in paleoenvironmental reconstruction: The micromammals of Manot Cave, Israel.

This paper describes the micromammalian remains and paleoenvironment of the Upper Paleolithic sequence of Manot Cave (46-34 ka), southern Levant. Micromammal remains were identified from Ahmarian (46-42 ka), Levantine Aurignacian (38-34 ka) and post-Levantine Aurignacian (34-33 ka) layers. To identify taphonomic agents, molar digestion was modelled for seven local raptor species, and model predictions were compared with observed digestion scores in the Manot Cave material. Raptor species differed significantly in molar digestion patterns, allowing us to identify Tyto alba as the bone accumulator in the Ahmarian and Post-Levantine Aurignacian units. Data were insufficient for species-level identification of the taphonomic agent in the Levantine Aurignacian. Günther's voles (Microtus guentheri) were dominant, though woodland species (Apodemus spp., Sciurus anomalus and Dryomys nitedula) also occurred through the entire sequence. Manot Cave furnished the first fossil record of the Eurasian snow vole (Chionomys nivalis) in the southern Levant during the Late Pleistocene and its first secure record of the water vole (Arvicola terrestris) outside the Hula Valley. Records of other taxa (Acomys dimidiatus and D. nitedula) provide the earliest occurrence datum in the region. Using counts of both identified specimens and distinct elements (lower first molars) we modelled paleoenvironmental conditions through both Weighted Averaging Partial Least Squares regression and a qualitative analysis considering niche preferences of species. Model predictions indicate the unexpectedly pronounced dominance of open habitats compared to present conditions near the cave, though the occurrence and abundance of woodland species also indicate some woodland expansion relative to the preceding Middle Paleolithic period. Combining statistical models and species niche considerations we show that the time span between the Ahmarian, Levantine Aurignacian and post-Levantine Aurignacian was marked by a clear-cut climatic oscillation to cooler and likely wetter conditions.

Increased songbird nest depredation due to Aleppo pine (Pinus halepensis) encroachment in Mediterranean shrubland.

BACKGROUND: In recent decades, a decrease of passerine densities was documented in Mediterranean shrublands. At the same time, a widespread encroachment of Aleppo pines (Pinus halepensis) to Mediterranean shrubland occurred. Such changes in vegetation structure may affect passerine predator assemblage and densities, and in turn impact passerine densities. Depredation during the nesting season is an important factor to influence passerine population size. Understanding the effects of changes in vegetation structure (pine encroachment) on passerine nesting success is the main objective of this study. We do so by assessing the effects of Aleppo pine encroachment on Sardinian warbler (Sylvia melanocephala) nest depredation in Mediterranean shrublands. We examined direct and indirect predation pressures through a gradients of pine density, using four methods: (1) placing dummy nests; (2) acoustic monitoring of mobbing events; (3) direct observations on nest predation using cameras; and (4) observation of Eurasian jay (Garrulus glandarius) behaviour as indirect evidence of predation risk. RESULTS: We found that Aleppo pine encroachment to Mediterranean shrublands increased nest predation by Eurasian jays. Nest predation was highest in mixed shrubland and pines. These areas are suitable for warblers but had high occurrence rate of Eurasian jays. CONCLUSIONS: Encroaching pines directly increase activity of Eurasian jays in shrubland habitats, which reduced the nesting success of Sardinian warblers. These findings are supported by multiple methodologies, illustrating different predation pressures along a gradient of pine densities in natural shrublands. Management of Aleppo pine seedlings and removal of unwanted trees in natural shrubland might mitigate arrival and expansion of predators and decrease the predation pressure on passerine nests.

Interbasin water transfer for the rehabilitation of a transboundary Mediterranean stream: An economic analysis.

Global demand for freshwater is increasing as human population grows, climate changes and water resources are being overexploited. Consequently, many freshwater ecosystems, particularly in water-stressed regions, are severely degraded. Here we present a unique case of an Interbasin Water Transfer (IWT) project aiming at ecosystem rehabilitation and recreation enhancement of an intermittent transboundary stream (Ayun, Israel). For the past century, water diversion at the Lebanese side had led to flow secession in the Israeli Ayun Nature Reserve during the dry season (May-November). To restore flow continuum, a sum of 0.5 million cubic meters of high quality freshwaters have been allocated annually during the dry months. The aim of this study is to evaluate the IWT project by: (1) examining the correlation between water flow in the Ayun and recreational visitation, and (2) performing a cost-benefit analysis of the IWT scheme, including non-market benefits. A time-series regression (Adj. R2 = 0.688, n = 125) shows that a 10% increase in water flow corresponds to a 2.1% increase in monthly visitors. An estimated 18.8% of the visitation rate between 2009 and 2015 can be attributed to the water reallocation project. Through a single-site travel cost model, we estimate the visitor's willingness to pay in US$ 37.8 per person per trip. When non-market benefits for recreation are included, the total benefits of the IWT project substantially exceed its costs. Our results suggest that IWT can be applied to restore water flow and enhance ecosystem services also in water-stressed regions.

Foraging Activity Pattern Is Shaped by Water Loss Rates in a Diurnal Desert Rodent.

Although animals fine-tune their activity to avoid excess heat, we still lack a mechanistic understanding of such behaviors. As the global climate changes, such understanding is particularly important for projecting shifts in the activity patterns of populations and communities. We studied how foraging decisions vary with biotic and abiotic pressures. By tracking the foraging behavior of diurnal desert spiny mice in their natural habitat and estimating the energy and water costs and benefits of foraging, we asked how risk management and thermoregulatory requirements affect foraging decisions. We found that water requirements had the strongest effect on the observed foraging decisions. In their arid environment, mice often lose water while foraging for seeds and cease foraging even at high energetic returns when water loss is high. Mice also foraged more often when energy expenditure was high and for longer times under high seed densities and low predation risks. Gaining insight into both energy and water balance will be crucial to understanding the forces exerted by changing climatic conditions on animal energetics, behavior, and ecology.

Chronobiology by moonlight.

Most studies in chronobiology focus on solar cycles (daily and annual). Moonlight and the lunar cycle received considerably less attention by chronobiologists. An exception are rhythms in intertidal species. Terrestrial ecologists long ago acknowledged the effects of moonlight on predation success, and consequently on predation risk, foraging behaviour and habitat use, while marine biologists have focused more on the behaviour and mainly on reproduction synchronization with relation to the Moon phase. Lately, several studies in different animal taxa addressed the role of moonlight in determining activity and studied the underlying mechanisms. In this paper, we review the ecological and behavioural evidence showing the effect of moonlight on activity, discuss the adaptive value of these changes, and describe possible mechanisms underlying this effect. We will also refer to other sources of night-time light ('light pollution') and highlight open questions that demand further studies.

Seasonal fluctuations attenuate stimulatory or inhibitory impacts of colonial birds on abundance, structure and diversity of soil biota.

Soil microorganisms and free-living nematodes were investigated in association with the nesting and roosting habitats of the following piscivorous and omnivorous colonial birds: black kite (Milvus migrans), great cormorant (Phalacrocorax carbo), black-crowned night heron (Nycticorax nycticorax) and little egret (Egretta garzetta), in Israel's Mediterranean region. Abiotic variables, abundance, trophic structure, sex ratio and genus diversity of soil free-living nematodes, and total abundance of bacteria and fungi, were measured during the wet season, following our previous study conducted during the dry season. The observed soil properties were important drivers of soil biota structure. Presence of the most efficient elements for soil organisms, such as phosphorus and nitrogen, was strongly dependent on the diet of the compared piscivorous and omnivorous bird colonies; levels of these nutrients were notably higher in the bird habitats than in their respective control sites during the study period. Ecological indices showed that the different species of colonial birds can have different (stimulatory or inhibitory) impacts on abundance and diversity of the soil biota, affecting the structure of the soil free-living nematode population at the generic, trophic and sexual levels during the wet season. A comparison with results from the dry season illustrated that seasonal fluctuations can change, and even attenuate the effect of bird activity on the abundance, structure and diversity of the soil communities.

Exploring scenarios for the food system-zoonotic risk interface.

The unprecedented economic and health impacts of the COVID-19 pandemic have shown the global necessity of mitigating the underlying drivers of zoonotic spillover events, which occur at the human-wildlife and domesticated animal interface. Spillover events are associated to varying degrees with high habitat fragmentation, biodiversity loss through land use change, high livestock densities, agricultural inputs, and wildlife hunting-all facets of food systems. As such, the structure and characteristics of food systems can be considered key determinants of modern pandemic risks. This means that emerging infectious diseases should be more explicitly addressed in the discourse of food systems to mitigate the likelihood and impacts of spillover events. Here, we adopt a scenario framework to highlight the many connections among food systems, zoonotic diseases, and sustainability. We identify two overarching dimensions: the extent of land use for food production and the agricultural practices employed that shape four archetypal food systems, each with a distinct risk profile with respect to zoonotic spillovers and differing dimensions of sustainability. Prophylactic measures to curb the emergence of zoonotic diseases are therefore closely linked to diets and food policies. Future research directions should explore more closely how they impact the risk of spillover events.

Possible origins and implications of atypical morphologies and domestication-like traits in wild golden jackals (Canis aureus).

Deciphering the origins of phenotypic variations in natural animal populations is a challenging topic for evolutionary and conservation biologists. Atypical morphologies in mammals are usually attributed to interspecific hybridisation or de-novo mutations. Here we report the case of four golden jackals (Canis aureus), that were observed during a camera-trapping wildlife survey in Northern Israel, displaying anomalous morphological traits, such as white patches, an upturned tail, and long thick fur which resemble features of domesticated mammals. Another individual was culled under permit and was genetically and morphologically examined. Paternal and nuclear genetic profiles, as well as geometric morphometric data, identified this individual as a golden jackal rather than a recent dog/wolf-jackal hybrid. Its maternal haplotype suggested past introgression of African wolf (Canis lupaster) mitochondrial DNA, as previously documented in other jackals from Israel. When viewed in the context of the jackal as an overabundant species in Israel, the rural nature of the surveyed area, the abundance of anthropogenic waste, and molecular and morphological findings, the possibility of an individual presenting incipient stages of domestication should also be considered.

Foraging sequence, energy intake and torpor: an individual-based field study of energy balancing in desert golden spiny mice.

We studied the relationship between sequence of foraging, energy acquired and use of torpor as an energy-balancing strategy in diurnally active desert golden spiny mice. We hypothesised that individuals that arrive earlier to forage will get higher returns and consequently spend less time torpid. If that is the case, then early foragers can be viewed as more successful; if the same individuals arrive repeatedly early, they are likely to have higher fitness under conditions of resource limitation. For the first time, we show a relationship between foraging sequence and amount of resources removed, with individuals that arrive later to a foraging patch tending to receive lower energetic returns and to spend more time torpid. Torpor bears not only benefits but also significant costs, so these individuals pay a price both in lower energy intake and in extended periods of torpor, in what may well be a positive feedback loop.

Biophysical modeling of the temporal niche: from first principles to the evolution of activity patterns.

Most mammals can be characterized as nocturnal or diurnal. However infrequently, species may overcome evolutionary constraints and alter their activity patterns. We modeled the fundamental temporal niche of a diurnal desert rodent, the golden spiny mouse, Acomys russatus. This species can shift into nocturnal activity in the absence of its congener, the common spiny mouse, Acomys cahirinus, suggesting that it was competitively driven into diurnality and that this shift in a small desert rodent may involve physiological costs. Therefore, we compared metabolic costs of diurnal versus nocturnal activity using a biophysical model to evaluate the preferred temporal niche of this species. The model predicted that energy expenditure during foraging is almost always lower during the day except during midday in summer at the less sheltered microhabitat. We also found that a shift in summer to foraging in less sheltered microhabitats in response to predation pressure and food availability involves a significant physiological cost moderated by midday reduction in activity. Thus, adaptation to diurnality may reflect the "ghost of competition past"; climate-driven diurnality is an alternative but less likely hypothesis. While climate is considered to play a major role in the physiology and evolution of mammals, this is the first study to model its potential to affect the evolution of activity patterns of mammals.

In Its Southern Edge of Distribution, the Tawny Owl (Strix aluco) Is More Sensitive to Extreme Temperatures Than to Rural Development.

Populations at the warm edge of distribution are more genetically diverse, and at the same time are more susceptible to climate change. Between 1987-1996, we studied Tawny Owls in Israel, the species' global southern edge of distribution and a country undergoing a rapid land cover transformation for over a century. To assess the potential impacts of land cover transformation, we modelled the species' most suitable habitat and climate and analyzed how climate and habitat affected the nesting success and prey selection. Moreover, we monitored Tawny Owl juveniles' survival and ontogeny from eggs to dietary independent young, to find out whether the Israeli population is a sink. While the species distribution model correctly predicted the Tawny Owl's densest areas of occurrence, it failed to predict its occurrence in adjacent regions. The model also predicted that areas included in the species' historical range remained suitable habitats. The number of fledglings increased with precipitation and in rural settings but was adversely affected by extreme temperatures. While voles dominated the diet in all habitats, the Tawny Owl's diet is considerably more variable than other Israeli owls. Our results suggest that the Tawny Owl can adapt to rural-agricultural environments, but is susceptible to climate change.

Community-wide character displacement in the presence of clines: a test of Holarctic weasel guilds.

1. Competition is thought to be a major influence on community assembly, ecology and evolution; presence of competitors may cause divergence in traits related to resource use (character displacement). 2. Such traits, however, often vary clinally, and this phenomenon may be independent of the presence or absence of competing species. 3. The presence of such clines can either obscure the effects of competition, or create an impression that competition is operating when, in fact, it is not. 4. We corrected for clinal variation while testing for character displacement in two well-studied weasel (Mustela) guilds, in the Nearctic and the west Palaearctic. 5. Without accounting for clines, our results agreed with previous studies suggesting character displacement in these guilds. 6. However, when we corrected for clines, predictions of competition theory were not met - and often we obtained evidence for character convergence in sympatry. 7. This may suggest that the nature of the resource base may be more important than interspecific competition in shaping morphology and size in these carnivores. 8. Our results highlight the need to account for geographic variation when studying character displacement and cast some doubt on prevailing ideas regarding the effect of competition on morphological evolution.

A model of digestive tooth corrosion in lizards: experimental tests and taphonomic implications.

Corrosion patterns induced by gastric fluids on the skeleton of prey animals may depend on the nature of the corrosive agents (acid, enzymes) as well as on the composition of the hard parts and the soft tissues that surround them. We propose a framework for predicting and interpreting corrosion patterns on lizard teeth, our model system, drawing on the different digestive pathways of avian and non-avian vertebrate predators. We propose that high-acid, low-enzyme systems (embodied by mammalian carnivores) will lead to corrosion of the tooth crowns, whereas low-acid, high-enzyme systems (embodied by owls) will lead to corrosion of the tooth shafts. We test our model experimentally using artificial gastric fluids (with HCl and pepsin) and feeding experiments, and phenomenologically using wild-collected owl pellets with lizard remains. Finding an association between the predictions and the experimental results, we then examine corrosion patterns on nearly 900 fossil lizard jaws. Given an appropriate phylogenetic background, our focus on physiological rather than taxonomic classes of predators allows the extension of the approach into Deep Time.

Life on the edge: carnivore body size variation is all over the place.

Evolutionary biologists have long been fascinated by both the ways in which species respond to ecological conditions at the edges of their geographic ranges and the way that species' body sizes evolve across their ranges. Surprisingly, though, the relationship between these two phenomena is rarely studied. Here, we examine whether carnivore body size changes from the interior of their geographic range towards the range edges. We find that within species, body size often varies strongly with distance from the range edge. However, there is no general tendency across species for size to be either larger or smaller towards the edge. There is some evidence that the smallest guild members increase in size towards their range edges, but results for the largest guild members are equivocal. Whether individuals vary in relation to the distance from the range edges often depends on the way edge and interior are defined. Neither geographic range size nor absolute body size influences the tendency of size to vary with distance from the range edge. Therefore, we suggest that the frequent significant association between body size and the position of individuals along the edge-core continuum reflects the prevalence of geographic size variation and that the distance to range edge per se does not influence size evolution in a consistent way.

What determines prey selection in owls? Roles of prey traits, prey class, environmental variables, and taxonomic specialization.

Ecological theory suggests that prey size should increase with predator size, but this trend may be masked by other factors affecting prey selection, such as environmental constraints or specific prey preferences of predator species. Owls are an ideal case study for exploring how predator body size affects prey selection in the presence of other factors due to the ease of analyzing their diets from owl pellets and their widespread distributions, allowing interspecific comparisons between variable habitats. Here, we analyze various dimensions of prey resource selection among owls, including prey size, taxonomy (i.e., whether or not particular taxa are favored regardless of their size), and prey traits (movement type, social structure, activity pattern, and diet). We collected pellets of five sympatric owl species (Athene noctua, Tyto alba, Asio otus, Strix aluco, and Bubo bubo) from 78 sites across the Mediterranean Levant. Prey intake was compared between sites, with various environmental variables and owl species as predictors of abundance. Despite significant environmental impacts on prey intake, some key patterns emerge among owl species studied. Owls select prey by predator body size: Larger owls tend to feed on wider ranges of prey sizes, leading to higher means. In addition, guild members show both specialization and generalism in terms of prey taxa, sometimes in contrast with the expectations of the predator-prey body size hypothesis. Our results suggest that while predator body size is an important factor in prey selection, taxon specialization by predator species also has considerable impact.

Fitness effects of interspecific competition between two species of desert rodents.

Many factors affect individual fitness, but while some factors, such as resource availability, have received strong experimental support, others including interspecific competition have rarely been quantified. Nevertheless, interspecific competition is commonly mentioned in the context of reproductive success and fitness. In general, when reproduction is likely to fail, reproductive suppression may occur. We studied the golden spiny mouse (Acomys russatus) and the common spiny mouse (A. cahirinus; however, recent molecular analysis in spiny mice from Jordan and Sinai suggests this species is A. dimidiatus (Frynta et al., 2010), as a model for the effect of competition on reproduction in four field enclosures: two populated only by A. russatus individuals, and two populated by individuals of both species. In presence of A. cahirinus, fitness of A. russatus was lower: the number of A. rusatus offspring was significantly lower; more males had regressed testes (indicating reproductive depression); more A. russatus young had damaged tails. However, no clear effect was evident in A. russatus female vaginal smear cytology. We conclude that the presence of A. cahirinus impairs fitness and reproductive success of A. russatus. While various direct and/or indirect mechanisms may be responsible for the effect of competition on reproduction, a plausible mechanism is increased use of torpor induced by the presence of A. cahirinus previously documented in A. russatus.

Opportunism or aquatic specialization? Evidence of freshwater fish exploitation at Ohalo II- A waterlogged Upper Paleolithic site.

Analysis of ca. 17,000 fish remains recovered from the late Upper Paleolithic/early Epi-Paleolithic (LGM; 23,000 BP) waterlogged site of Ohalo II (Rift Valley, Israel) provides new insights into the role of wetland habitats and the fish inhabiting them during the evolution of economic strategies prior to the agricultural evolution. Of the current 19 native fish species in Lake Kinneret (Sea of Galilee), eight species were identified at Ohalo II, belonging to two freshwater families: Cyprinidae (carps) and Cichlidae (St. Peter fish). Employing a large set of quantitative and qualitative criteria (NISP, species richness, diversity, skeletal element representation, fragmentation, color, spatial distribution, etc.), we demonstrate that the inhabitants of Ohalo II used their knowledge of the breeding behavior of different species of fish, for year-round intensive exploitation.

Reliability of a higher-taxon approach to richness, rarity, and composition assessments at the local scale.

A promising shortcut for quantifying species patterns is to use genera and families as surrogates of species. At large spatial scales, concurrence between patterns of richness, rarity, and composition of species and higher taxa is generally high. Only a few researchers, however, have examined this relationship at the local scale, which is frequently the relevant scale in land-use conflicts. We investigated the reliability of the higher-taxon approach in assessing patterns of species richness, rarity, and composition at the local scale. We studied diversity patterns of three commonly used surrogate taxa: vascular plants, ground-dwelling beetles, and moths. We conducted year-round field surveys for these taxa in the Jerusalem Mountains and the Judean foothills, Israel. Richness and composition of species were highly correlated with richness and composition of genera for all taxa. At the family level, correlations with richness and composition of species were much lower. Excluding monotypic genera and families did not affect these relations. Rarity representation based on higher taxa varied considerably depending on the taxon, and rarity scale and was weaker compared with richness and composition representation. Cumulative richness curves of species and genera showed similar patterns, leveling off at equivalent sampling efforts. Genus-level assessments were a reliable surrogate for local patterns of species richness, rarity, and composition, but family-level assessments performed poorly. The advantage of using coarse taxonomic scales in local diversity surveys is that it may decrease identification time and the need for experts, but it will not reduce sampling effort.

The relationship between the golden spiny mouse circadian system and its diurnal activity: an experimental field enclosures and laboratory study.

Examples of animals that switch activity times between nocturnality and diurnality in nature are relatively infrequent. Furthermore, the mechanism for switching activity time is not clear: does a complete inversion of the circadian system occur in conjunction with activity pattern? Are there switching centers downstream from the internal clock that interpret the clock differently? Or does the switch reflect a masking effect? Answering these key questions may shed light on the mechanisms regulating activity patterns and their evolution. The golden spiny mouse (Acomys russatus) can switch between nocturnal and diurnal activity. This study investigated the relationship between its internal circadian clock and its diurnal activity pattern observed in the field. The goal is to understand the mechanisms underlying species rhythm shifts in order to gain insight into the evolution of activity patterns. All golden spiny mice had opposite activity patterns in the field than those under controlled continuous dark conditions in the laboratory. Activity and body temperature patterns in the field were diurnal, while in the laboratory all individuals immediately showed a free-running rhythm starting with a nocturnal pattern. No phase transients were found toward the preferred nocturnal activity pattern, as would be expected in the case of true entrainment. Moreover, the fact that the free-running activity patterns began from the individuals' subjective night suggests that golden spiny mice are nocturnal and that their diurnality in their natural habitat in the field results from a change that is downstream to the internal clock or reflects a masking effect.

Temporal niche expansion in mammals from a nocturnal ancestor after dinosaur extinction.

Most modern mammals, including strictly diurnal species, exhibit sensory adaptations to nocturnal activity that are thought to be the result of a prolonged nocturnal phase or 'bottleneck' during early mammalian evolution. Nocturnality may have allowed mammals to avoid antagonistic interactions with diurnal dinosaurs during the Mesozoic. However, understanding the evolution of mammalian activity patterns is hindered by scant and ambiguous fossil evidence. While ancestral reconstructions of behavioural traits from extant species have the potential to elucidate these patterns, existing studies have been limited in taxonomic scope. Here, we use an extensive behavioural dataset for 2,415 species from all extant orders to reconstruct ancestral activity patterns across Mammalia. We find strong support for the nocturnal origin of mammals and the Cenozoic appearance of diurnality, although cathemerality (mixed diel periodicity) may have appeared in the late Cretaceous. Simian primates are among the earliest mammals to exhibit strict diurnal activity, some 52-33 million years ago. Our study is consistent with the hypothesis that temporal partitioning between early mammals and dinosaurs during the Mesozoic led to a mammalian nocturnal bottleneck, but also demonstrates the need for improved phylogenetic estimates for Mammalia.