Decline and recovery in otter Lutra lutra populations in Italy

• 1 Changes in otter Lutra lutra distribution in Italy were examined by analysing geographical, historical and survey data.
• 2 As in other European countries, otters declined sharply in Italy during the late 1970s and throughout the 1980s. Between 1985 and 2004 the species became extinct, except for some reintroductions, in northern and most of central Italy while the species appears to have substantially recovered in its southern range. This recovery of the otter population is apparently not due to increased research effort, but reflects a real expansion of range.
• 3 Differences in the degree of range reduction during the second half of the 20th century and in the availability of suitable habitats are probably the major factors that have determined the two opposite trends shown by otter populations in the last 20 years.
• 4 Better knowledge of otter status and distribution is needed for effective conservation management. We suggest that a national survey of the Italian otter population that employs genetic approaches should represent a first step of the national action plan for otter conservation.

     

Genetic signature of the colonisation dynamics along a coastal expansion front in the damselfly Coenagrion scitulum

1. Many insects are expanding their distribution range polewards as a result of climate change, which has been shown to be associated with founder effects leading to a reduction in genetic diversity and an increase in genetic differentiation. These spatial genetic patterns may arise from colonisation from a broad expansion front or a limited neighbourhood after a stepping stone model of dispersal. The temporal persistence of such founder effects are poorly understood, mainly because studies looking at the fine‐scale initial temporal dynamics of the genetic signature of a range expansion are rare. 2. Using microsatellite markers, we performed a detailed spatiotemporal genetic analysis of the range expanding damselfly Coenagrion scitulum (Rambur) along a coastal axis during the first years after colonisation. 3. A decrease was in (private) allelic richness when going northwards along the coastline, which is consistent with a scenario of cumulative founder events. In spite of the spatiotemporal dynamics in the observation records of the species along the coastline, the spatial genetic data indicated a major contribution from the broad expansion front during the colonisation of the coastline rather than a stepping‐stone colonisation process. 4. The fine‐scale temporal dynamics of the range expansion indicated the absence of persistent founder effects and instead showed considerable temporal instability in genetic indices at the more northern edge populations. This may be explained by genetic immigration and admixture from the broad expansion front in this active disperser.     

Reid's paradox revisited: the evolution of dispersal kernels during range expansion

Current approaches to modeling range advance assume that the distribution describing dispersal distances in the population (the "dispersal kernel") is a static entity. We argue here that dispersal kernels are in fact highly dynamic during periods of range advance because density effects and spatial assortment by dispersal ability ("spatial selection") drive the evolution of increased dispersal on the expanding front. Using a spatially explicit individual-based model, we demonstrate this effect under a wide variety of population growth rates and dispersal costs. We then test the possibility of an evolved shift in dispersal kernels by measuring dispersal rates in individual cane toads (Bufo marinus) from invasive populations in Australia (historically, toads advanced their range at 10 km/year, but now they achieve >55 km/year in the northern part of their range). Under a common-garden design, we found a steady increase in dispersal tendency with distance from the invasion origin. Dispersal kernels on the invading front were less kurtotic and less skewed than those from origin populations. Thus, toads have increased their rate of range expansion partly through increased dispersal on the expanding front. For accurate long-range forecasts of range advance, we need to take into account the potential for dispersal kernels to be evolutionarily dynamic.     

Land use drivers of species re‐expansion: inferring colonization dynamics in Eurasian otters

Aim Land use intensity has been recognized as one of the major determinants of native species declines. The re‐expansion of species previously constrained by habitat degradation has been rarely investigated. Here, we use site occupancy models incorporating imperfect detection to identify the land use drivers of the re‐expansion of the Eurasian otter (Lutra lutra). Location Czech Republic. Methods We applied multi‐season occupancy models to otter presence–non‐detection data collected in three national surveys (1992, 2000, 2006) at 552 sites (11.2 × 12 km grid cells). Model parameters included site occupancy, colonization and extinction probabilities, and detection probability at a sub‐site level. We modelled changes in occupancy over time as a function of agricultural, urban and industrial land use and change in the extent of agricultural land use. Results Under the best fitting model, occupancy was estimated to be 34.6% in 1992, 51.3% in 2000 and 83.7% in 2006. Detection probability was neither perfect nor constant. Occupancy probability in 1992 was negatively related to land use gradients. Colonization was more likely to occur where a reduction in agricultural land was larger. Variation in extinction and colonization rates along land use gradients resulted in increased occupancy in industrial and especially urban landscapes. Conversely, occupancy remained almost unchanged along agricultural gradients. Main conclusions Dynamics of otter expansion were strongly associated with the two main patterns of the rapid environmental transition that has taken place in the Czech Republic since the early 1990s. Results show that a reduction in intensive agricultural land use led to an increase in otter distribution, providing evidence of the impact of agricultural land use on stream ecosystems. Moreover, otters recolonized urban and industrial landscapes, probably as a result of extensive reduction in water pollution from point sources. Our results suggest that active conservation of otter populations should focus on restoration of freshwater habitat at large scales, especially in agricultural landscapes.     

Forecasting range expansion into ecological traps: climate‐mediated shifts in sea turtle nesting beaches and human development

Some species are adapting to changing environments by expanding their geographic ranges. Understanding whether range shifts will be accompanied by increased exposure to other threats is crucial to predicting when and where new populations could successfully establish. If species overlap to a greater extent with human development under climate change, this could form ecological traps which are attractive to dispersing individuals, but the use of which substantially reduces fitness. Until recently, the core nesting range for the Critically Endangered Kemp's ridley sea turtle (Lepidochelys kempii) was ca. 1000 km of sparsely populated coastline in Tamaulipas, Mexico. Over the past twenty‐five years, this species has expanded its range into populated areas of coastal Florida (>1500 km outside the historical range), where nesting now occurs annually. Suitable Kemp's ridley nesting habitat has persisted for at least 140 000 years in the western Gulf of Mexico, and climate change models predict further nesting range expansion into the eastern Gulf of Mexico and northern Atlantic Ocean. Range expansion is 6–12% more likely to occur along uninhabited stretches of coastline than are current nesting beaches, suggesting that novel nesting areas will not be associated with high levels of anthropogenic disturbance. Although the high breeding‐site fidelity of some migratory species could limit adaptation to climate change, rapid population recovery following effective conservation measures may enhance opportunities for range expansion. Anticipating the interactive effects of past or contemporary conservation measures, climate change, and future human activities will help focus long‐term conservation strategies.     

Exploring the range expansion of the yellow-spotted longhorn beetle Psacothea hilaris hilaris in northern Italy

1. Psacothea hilaris hilaris is an exotic cerambycid detected in Lombardy (northern Italy) in 2005 and now established and undergoing an expansion phase. The species constitutes a serious pest for Ficus spp. and Morus spp.
2. The aim of this work is to update the distribution area of the species in Italy, investigating its expansion over 16 years. Data were acquired through the authors' direct field records and community science, by data mining on national and international web platforms and social networks. Reports were analysed and GPS points plotted in ArcMap 10.8.1 to obtain distribution maps.
3. Of the records collected, 34% were acquired during field surveys, 22% via e-mail and 44% through on-line forums and social networks.
4. Adults of P. h. hilaris showed a long period of activity, from April to the beginning of December, with a peak of presence from June to September. The presence of the pest has been ascertained over six provinces in Lombardy covering an area estimated at more than 1750 km² in 2021. New detections occurred within 2 km of a known infestation from the previous year in 41.6% of cases, and between 2 and 4 km away in 37.7% of cases. The mean rate of range expansion estimated from reported records was 3.17 ± 0.33 km/year from 2010 to 2021. After an initial phase of settlement, the insect is now spreading southward.
5. Given the relevant phytosanitary interest that the species may have for fig plants in Italy and in the Mediterranean Basin, it is fundamental to deepen the knowledge about its flight biology, dynamics and active dispersal capabilities in order to estimate the future range expansion.

     

Birds on the move in the face of climate change: High species turnover in northern Europe

Species richness is predicted to increase in the northern latitudes in the warming climate due to ranges of many southern species expanding northwards. We studied changes in the composition of the whole avifauna and in bird species richness in a period of already warming climate in Finland (in northern Europe) covering 1,100 km in south–north gradient across the boreal zone (over 300,000 km²). We compared bird species richness and species‐specific changes (for all 235 bird species that occur in Finland) in range size (number of squares occupied) and range shifts (measured as median of area of occupancy) based on bird atlas studies between 1974–1989 and 2006–2010. In addition, we tested how the habitat preference and migration strategy of species explain species‐specific variation in the change of the range size. The study was carried out in 10 km squares with similar research intensity in both time periods. The species richness did not change significantly between the two time periods. The composition of the bird fauna, however, changed considerably with 37.0% of species showing an increase and 34.9% a decrease in the numbers of occupied squares, that is, about equal number of species gained and lost their range. Altogether 95.7% of all species (225/235) showed changes either in the numbers of occupied squares or they experienced a range shift (or both). The range size of archipelago birds increased and long‐distance migrants declined significantly. Range loss observed in long‐distance migrants is in line with the observed population declines of long‐distance migrants in the whole Europe. The results show that there is an ongoing considerable species turnover due to climate change and due to land use and other direct human influence. High bird species turnover observed in northern Europe may also affect the functional diversity of species communities.     

Evolutionarily accelerated invasions: the rate of dispersal evolves upwards during the range advance of cane toads

Human activities are changing habitats and climates and causing species' ranges to shift. Range expansion brings into play a set of powerful evolutionary forces at the expanding range edge that act to increase dispersal rates. One likely consequence of these forces is accelerating rates of range advance because of evolved increases in dispersal on the range edge. In northern Australia, cane toads have increased their rate of spread fivefold in the last 70 years. Our breeding trials with toads from populations spanning the species' invasion history in Australia suggest a genetic basis to dispersal rates and interpopulation genetic variation in such rates. Toads whose parents were from the expanding range front dispersed faster than toads whose parents were from the core of the range. This difference reflects patterns found in their field-collected mothers and fathers and points to heritable variance in the traits that have accelerated the toads' rate of invasion across tropical Australia over recent decades. Taken together with demonstrated spatial assortment by dispersal ability occurring on the expanding front, these results point firmly to ongoing evolution as a driving force in the accelerated expansion of toads across northern Australia.     

Deleterious mutations can surf to high densities on the wave front of an expanding population

There is an increasing recognition that evolutionary processes play a key role in determining the dynamics of range expansion. Recent work demonstrates that neutral mutations arising near the edge of a range expansion sometimes surf on the expanding front leading them rather than that leads to reach much greater spatial distribution and frequency than expected in stationary populations. Here, we extend this work and examine the surfing behavior of nonneutral mutations. Using an individual-based coupled-map lattice model, we confirm that, regardless of its fitness effects, the probability of survival of a new mutation depends strongly upon where it arises in relation to the expanding wave front. We demonstrate that the surfing effect can lead to deleterious mutations reaching high densities at an expanding front, even when they have substantial negative effects on fitness. Additionally, we highlight that this surfing phenomenon can occur for mutations that impact reproductive rate (i.e., number of offspring produced) as well as mutations that modify juvenile competitive ability. We suggest that these effects are likely to have important consequences for rates of spread and the evolution of spatially expanding populations.     

Strong genetic differentiation but not local adaptation toward the range limit of a coastal dune plant

All species have limited geographic distributions; but the ecological and evolutionary mechanisms causing range limits are largely unknown. That many species’ geographic range limits are coincident with niche limits suggests limited evolutionary potential of marginal populations to adapt to conditions experienced beyond the range. We provide a test of range limit theory by combining population genetic analysis of microsatellite polymorphisms with a transplant experiment within, at the edge of, and 60 km beyond the northern range of a coastal dune plant. Contrary to expectations, lifetime fitness increased toward the range limit with highest fitness achieved by most populations at and beyond the range edge. Genetic differentiation among populations was strong, with very low, nondirectional gene flow suggesting range limitation via constraints to dispersal. In contrast, however, local adaptation was negligible, and a distance‐dependent decline in fitness only occurred for those populations furthest from home when planted beyond the range limit. These results challenge a commonly held assumption that stable range limits match niche limits, but also raise questions about the unique value of peripheral populations in expanding species’ geographical ranges.     

Evaluating range‐expansion models for calculating nonnative species' expansion rate

Species range shifts associated with environmental change or biological invasions are increasingly important study areas. However, quantifying range expansion rates may be heavily influenced by methodology and/or sampling bias. We compared expansion rate estimates of Roesel''s bush-cricket (Metrioptera roeselii, Hagenbach 1822), a nonnative species currently expanding its range in south-central Sweden, from range statistic models based on distance measures (mean, median, 95^th gamma quantile, marginal mean, maximum, and conditional maximum) and an area-based method (grid occupancy). We used sampling simulations to determine the sensitivity of the different methods to incomplete sampling across the species'' range. For periods when we had comprehensive survey data, range expansion estimates clustered into two groups: (1) those calculated from range margin statistics (gamma, marginal mean, maximum, and conditional maximum: ˜3 km/year), and (2) those calculated from the central tendency (mean and median) and the area-based method of grid occupancy (˜1.5 km/year). Range statistic measures differed greatly in their sensitivity to sampling effort; the proportion of sampling required to achieve an estimate within 10% of the true value ranged from 0.17 to 0.9. Grid occupancy and median were most sensitive to sampling effort, and the maximum and gamma quantile the least. If periods with incomplete sampling were included in the range expansion calculations, this generally lowered the estimates (range 16–72%), with exception of the gamma quantile that was slightly higher (6%). Care should be taken when interpreting rate expansion estimates from data sampled from only a fraction of the full distribution. Methods based on the central tendency will give rates approximately half that of methods based on the range margin. The gamma quantile method appears to be the most robust to incomplete sampling bias and should be considered as the method of choice when sampling the entire distribution is not possible.     

Mutation surfing and the evolution of dispersal during range expansions

A growing body of empirical evidence demonstrates that at an expanding front, there can be strong selection for greater dispersal propensity, whereas recent theory indicates that mutations occurring towards the front of a spatially expanding population can sometimes ‘surf’ to high frequency and spatial extent. Here, we consider the potential interplay between these two processes: what role may mutation surfing play in determining the course of dispersal evolution and how might dispersal evolution itself influence mutation surfing? Using an individual‐based coupled‐map lattice model, we first run simulations to determine the fate of dispersal mutants that occur at an expanding front. Our results highlight that mutants that have a slightly higher dispersal propensity than the wild type always have a higher survival probability than those mutants with a dispersal propensity lower than, or very similar to, the wild type. However, it is not always the case that mutants with very high dispersal propensity have the greatest survival probability. When dispersal mortality is high, mutants of intermediate dispersal survive most often. Interestingly, the rate of dispersal that ultimately evolves at an expanding front is often substantially higher than that which confers a novel mutant with the greatest probability of survival. Second, we run a model in which we allow dispersal to evolve over the course of a range expansion and ask how the fate of a neutral or nonneutral mutant depends upon when and where during the expansion it arises. These simulations highlight that the success of a neutral mutant depends upon the dispersal genotypes that it is associated with. An important consequence of this is that novel mutants that arise at the front of an expansion, and survive, typically end up being associated with more dispersive genotypes than the wild type. These results offer some new insights into causes and the consequences of dispersal evolution during range expansions, and the methodology we have employed can be readily extended to explore the evolutionary dynamics of other life history characteristics.     

Rapid adaptive evolution of northeastern coyotes via hybridization with wolves

The dramatic expansion of the geographical range of coyotes over the last 90 years is partly explained by changes to the landscape and local extinctions of wolves, but hybridization may also have facilitated their movement. We present mtDNA sequence data from 686 eastern coyotes and measurements of 196 skulls related to their two-front colonization pattern. We find evidence for hybridization with Great Lakes wolves only along the northern front, which is correlated with larger skull size, increased sexual dimorphism and a five times faster colonization rate than the southern front. Northeastern haplotype diversity is low, suggesting that this population was founded by very few females moving across the Saint Lawrence River. This northern front then spread south and west, eventually coming in contact with an expanding front of non-hybrid coyotes in western New York and Pennsylvania. We suggest that hybridization with wolves in Canada introduced adaptive variation that contributed to larger size, which in turn allowed eastern coyotes to better hunt deer, allowing a more rapid colonization of new areas than coyotes without introgressed wolf genes. Thus, hybridization is a conduit by which genetic variation from an extirpated species has been reintroduced into northeastern USA, enabling northeastern coyotes to occupy a portion of the niche left vacant by wolves.     

Rapid range expansion increases genetic differentiation while causing limited reduction in genetic diversity in a damselfly

Many ectothermic species are currently expanding their geographic range due to global warming. This can modify the population genetic diversity and structure of these species because of genetic drift during the colonization of new areas. Although the genetic signatures of historical range expansions have been investigated in an array of species, the genetic consequences of natural, contemporary range expansions have received little attention, with the only studies available focusing on range expansions along a narrow front. We investigate the genetic consequences of a natural range expansion in the Mediterranean damselfly Coenagrion scitulum, which is currently rapidly expanding along a broad front in different directions. We assessed genetic diversity and genetic structure using 12 microsatellite markers in five centrally located populations and five recently established populations at the edge of the geographic distribution. Our results suggest that, although a marginal significant decrease in the allelic richness was found in the edge populations, genetic diversity has been preserved during the range expansion of this species. Nevertheless, edge populations were genetically more differentiated compared with core populations, suggesting genetic drift during the range expansion. The smaller effective population sizes of the edge populations compared with central populations also suggest a contribution of genetic drift after colonization. We argue and document that range expansion along multiple axes of a broad expansion front generates little reduction in genetic diversity, yet stronger differentiation of the edge populations.     

Host‐targeted RAD‐Seq reveals genetic changes in the coral Oculina patagonica associated with range expansion along the Spanish Mediterranean coast

Many organisms are expanding their ranges in response to changing environmental conditions. Understanding the patterns of genetic diversity and adaptation along an expansion front is crucial to assessing a species’ long‐term success. While next‐generation sequencing techniques can reveal these changes in fine detail, ascribing them to a particular species can be difficult for organisms that live in close association with symbionts. Using a novel modified restriction site‐associated DNA sequencing (RAD‐Seq) protocol to target coral DNA, we collected 595 coral‐specific single nucleotide polymorphisms from 189 colonies of the invasive coral Oculina patagonica from the Spanish Mediterranean coast, including established core populations and two expansion fronts. Surprisingly, populations from the recent northern expansion are genetically distinct from the westward expansion and core populations and also harbour greater genetic diversity. We found that temperature may have driven adaptation along the northern expansion, as genome scans for selection found three candidate loci associated with temperature in the north but none in the west. We found no genomic signature of selection associated with artificial substrate, which has been proposed for explaining the rapid spread of O. patagonica. This suggests that this coral is simply an opportunistic colonizer of free space made available by coastal habitat modifications. Our results suggest that unique genetic variation, possibly due to limited dispersal across the Ibiza Channel, an influx of individuals from different depths and/or adaptation to cooler temperatures along the northern expansion front may have facilitated the northward range expansion of O. patagonica in the western Mediterranean.     

Survey of an otter<Emphasis Type="Italic">Lutra lutra</Emphasis> population in Southern Italy: site occupancy and influence of sampling season on species detection

In order to assess the current status of the otterLutra lutra Linnaeus, 1758 in Southern Campania (Italy), we surveyed 141 sites for spraints (faeces) in 2001. Fifty-eight sites were surveyed during the winter and in the following summer in order to test, through an estimation-based approach, the influence of sampling season on species detection. Site occupation in the study area was high (69.5%) and possibly underestimated, because the survey was affected by non-detection errors. Our analyses showed that winter surveying markedly underestimated true otter occupancy at the 58 sites (51.7% vs 97.1%), whereas summer surveying was very reliable (91.4%). Rains and floods may have removed spraints during winter, thereby reducing the detection probability to 0.534. These results suggest that otter standard surveys in areas with Mediterranean-type climates should be conducted during summer or periods without prolonged precipitation. Comparing our results with those of the 1985 National survey, we found an occupancy increase from 65.8 to 100%. We could not establish whether this change reflected a population increase or was due to possible non-detection errors that occurred in the 1985 survey. However, the present occupancy substantiates the strategic relevance of the study area for planning the conservation of Italian otters.     

Natural,human and spatial constraints to expanding populations of otters in the Iberian Peninsula

Aim To determine the relationships between otter (Lutra lutra) distribution dynamics and environmental and spatial constraints over a 20‐year period. Location Andalusia, southern Iberian Peninsula. Methods We synthesized otter distribution data from three otter surveys (1985, 1995 and 2005) using subcatchment areas defined by hydrological barriers. Subcatchments were characterized by two ‘natural’ (climatic and orographic variables) and two ‘human’ (land use and population density) gradients. In addition, we calculated two contagion variables (the distance to previously occupied subcatchments and the percentage of occupied subcatchments within a 50 km buffer) for consecutively surveyed subcatchments. Results Between 1985 and 2005 the percentage of subcatchments with otters present increased from 42% to 72%. Otters tended to be rare or absent from human‐dominated areas. Anthropogenic gradients were better predictors of otter distribution than natural ones. Human and natural gradients showed strong covariation, but for any value of the natural gradients otters tended to be present in subcatchments with lower human impacts. Colonization of new subcatchments was found to be strongly related to contagion variables and expansion rates were slower than those estimated in other studies. Newly colonized areas tended to be located in areas with intermediate human influence, while repeated absences occurred mainly in areas where human impact was most severe. Main conclusions Our results suggest that recent otter expansion across Andalusia is a reflection of large‐scale improvement in environmental conditions. Otter populations that survived the period of strong and generalized declines appear to be acting as sources from which neighbouring areas are colonized, probably aided by improved water quality and increases in food availability. However, the further expansion of otters into their full original range is likely to be constrained by human‐impacted landscapes.     

Ecophysiological constraints shape autumn migratory response to climate change in the North American field sparrow

Our ability to accurately forecast species' geographical responses to climate change requires knowledge of the proximate and ultimate drivers of their distribution. Here, we consider the ecophysiological and demographic determinants of the distribution of a partial migrant, the North American field sparrow, Spizella pusilla. From 1940 to 1963, the field sparrow extended its winter northern range margin 222km polewards. Such expansion was coincident with not only a geographical expansion into suitable breeding habitats, but also a decrease in mean abundance across sites occupied during the winter surveys. Combined, these trends suggest that declining populations along the expansion front either stopped migrating or altered their autumn migration. The poleward expansion was not coincident with climatically induced decreases in peak metabolic energy demand, but it did track increases in ecosystem net primary productivity. After 1963, the species' lower lethal temperature prevented further poleward movement. These findings show how different ecophysiological constraints can interact to change migration and distribution in a demographically declining species.     

Long‐distance dispersal maximizes evolutionary potential during rapid geographic range expansion

Conventional wisdom predicts that sequential founder events will cause genetic diversity to erode in species with expanding geographic ranges, limiting evolutionary potential at the range margin. Here, we show that invasive European starlings (Sturnus vulgaris) in South Africa preserve genetic diversity during range expansion, possibly as a result of frequent long‐distance dispersal events. We further show that unfavourable environmental conditions trigger enhanced dispersal, as indicated by signatures of selection detected across the expanding range. This brings genetic variation to the expansion front, counterbalancing the cumulative effects of sequential founding events and optimizing standing genetic diversity and thus evolutionary potential at range margins during spread. Therefore, dispersal strategies should be highlighted as key determinants of the ecological and evolutionary performances of species in novel environments and in response to global environmental change.