Range reexpansion after long stasis: Italian otters (Lutra lutra) at their northern edge

Species range shifts and expansion are subjects of primary research interest in the context of climate warming and biological invasions. Few studies have focused on reexpansion of species that suffered severe declines. Here, we focused on population recovery of Eurasian otters (Lutra lutra) in Italy, first detected in 2003 after a southward range contraction. We modeled the rate of range expansion and occupancy at the northern expanding front (central Italy), to gain insights into the progress of recovery and mechanisms of reexpansion. We performed a field survey in 2021, which redefined the northern limit of distribution further north, in close proximity to the Gran Sasso National Park. Then we analyzed a time series (1985–2021) of distances of northernmost occurrences from the center of the 1985 range. Using segmented regression, we were able to identify a prolonged stasis of the northern range edge and a simultaneous increase in occupancy from 0.151 to 0.4. A breakpoint was estimated in 2006, after which the range expanded northwards at an average rate of 5.48 km/year. From 2006 to 2021, the overall northward shift was about 80 km. Occupancy continued to increase until 2019 and abruptly declined in 2021. These patterns suggest that the reexpansion of the range can be limited by low occupancy at the expanding front. As occupancy increases, long-distance dispersal increases and then range expands. The low occupancy at the current distribution limit of otters may reflect a higher anthropogenic pressure on northern habitats, which could slow down the reexpansion process.     

A critical evaluation of host ranges of parasitoids of the subtribe Diabroticina (Coleoptera: Chrysomelidae: Galerucinae: Luperini) using field and laboratory host records

The subtribe Diabroticina is a large group of New World Chrysomelidae that includes corn rootworms, cucumber beetles and other pests. Recent introductions of Diabrotica virgifera virgifera LeConte into Europe, and the development of resistances to current management practices of rootworms in the USA have increased interest in new sustainable options for managing those pests. The only parasitoids that have been shown to consistently target and develop inside the beetle adults are Centistes gasseni Shaw, Centistes diabroticae Gahan (both Hym.: Braconidae), and Celatoria diabroticae Shimer, Celatoria compressa (Wulp), Celatoria bosqi Blanchard, and Celatoria setosa Coquillett (all Diptera: Tachinidae). This review improves our understanding of the realised and potential host range of these known parasitoids by rectifying erroneous references in light of new host records and from laboratory host range tests. Based on this critical review, all tachinid and braconid species studied are considered to be specific at least to the level of subtribe, i.e. Diabroticina. Celatoria setosa, Celatoria diabroticae and C. bosqi, have a narrow realized and potential host range; the former is restricted to the genus Acalymma and the last two to the fucata and virgifera groups of the genus Diabrotica. The braconids Centistes gasseni and C. diabroticae are also specific. The realized host range of C. gasseni includes species in the Diabrotica fucata and virgifera groups; while its potential host range also includes Acalymma species. The realized and potential host range of Centistes diabroticae includes Acalymma species as well as species in the fucata and virgifera groups of Diabrotica. Celatoria compressa has the broadest realised range compared to the other species studied, since it was obtained from species in several genera of Diabroticina; and its potential host range may also include Old World Aulacophora species.     

Determination of gasoline range,diesel range,and mineral oil range organics in soils and water by flame lonization gas chromatography

This article describes a method for the determination of gasoline range, diesel range, and mineral oil range organics in soils and water. It represents the culmination of a series of efforts to go beyond typical GRO and DRO methodology currently available in the literature to include a quantitative determination of mineral oil organics having a boiling range up to C₄₄. It also is the result of an attempt to develop a cost‐effective method that enables the analyst to quantify three different types of hydrocarbon components in one GC run under conditions without a concentration step. Method performance is comparable to that of current protocols for GRO and DRO determinations and validated further by comparisons to certified standards and in‐house standards. Accuracy as percentage recovery for GRO in water is 82 to 84 and 91 to 92% for soils. Accuracy as percentage recovery for MRO in water is 84 to 102 and 75 to 80% for soils. Accuracy as percentage recovery for DRO in water is 78 to 100 and 71 to 90% for soils. Results from the analysis of in‐house standards and certified standards for DRO and MRO gave higher recoveries than was demonstrated in the MDL studies. The MDL for DRO in water is 31 mg/l and 14 mg/kg for soils. The MDL for GRO in water is 8 mg/l and 4 mg/kg for soils. For MRO in water, the MDL is 7 mg/l and for soils 10 mg/kg. Future proposed improvements to this method will involve updated software that will allow automatic blank subtraction, automatic calculation of surrogate recoveries and the automatic incorporation of dry weight factors in the final calculations for soils. In addition, a GRO method with a run time of only 24 min will be used routinely when only GRO analyses have to be performed.     

Demographic and population‐genetic tests provide mixed support for the abundant centre hypothesis in the endemic plant Leavenworthia stylosa

The abundant centre hypothesis (ACH) assumes that population abundance, population size, density and per‐capita reproductive output should peak at the centre of a species' geographic range and decline towards the periphery. Increased isolation among and decreased reproductive output within edge populations should reduce within‐population genetic diversity and increase genetic differentiation among edge relative to central populations. The ACH also predicts asymmetrical gene flow, with net movement of migrants from the centre to edges. We evaluated these ecological assumptions and population‐genetic predictions in the endemic flowering plant Leavenworthia stylosa. Although populations were more spatially isolated near range edges, the geographic centre was surrounded by and not coincident with areas of peak population abundance, and plant density increased towards range edges. Per‐capita seed number was not associated with distance to the range centre, but seed number/m² increased near range edges. In support of ACH predictions, allelic diversity at 12 microsatellite loci declined with distance from the range centre, and pairwise F_ST values were higher between edge populations than between central populations. Coalescent analyses confirmed that gene flow was most infrequent between edge populations, but there was not an asymmetric pattern of gene flow predicted by the ACH. This study shows that among‐population demographic variability largely did not support the ACH, while patterns of genetic diversity, differentiation and gene flow were generally consistent with its predictions. Such mixed support has frequently been observed in tests of the ACH and raises concerns regarding the generality of this hypothesis for species range limits.     

Do local abundances of British birds change with proximity to range edge?

Aim Species generally vary in the density they attain at different sites, prompting the question of whether this variation is systematic across their range. We investigate this question using data on the abundance and distribution of thirty-two species of passerine birds across Britain derived from censuses organized by the British Trust for Ornithology. Methods Analysis is complicated by the issue of quantifying the distance of any particular census location from the edge of the range of a species when the study area encompasses only part of its entire distribution. No measure of this quantity is a priori the correct one, and so we use a variety of different measures which make differing assumptions about how abundances might be structured across species ranges. Results None of the measures used reveal any consistent relationships between the density attained by species at census sites and the spatial positions of those sites. Only thirteen species show significant relationships with any of the measures, and no more than seven species with any single measure. Main conclusion In summary, there is no convincing evidence that passerine bird densities are usually lower towards range edges in Britain. We discuss possible reasons for these findings.     

Comparison of overwintering survival and fertility of Zaprionus indianus (Diptera: Drosophilidae) flies from native and invaded ranges

Zaprionus indianus is a fly species native to the Afrotropical biogeographic region that invaded the South American continent 20 years ago. Its southernmost record is 34°S in areas with temperate climates with cold winters. To better understand its invasion biology, we investigated physiological responses to winter-like abiotic conditions that may be relevant in Z. indianus geographic expansion. We characterized Z. indianus females reproductive traits (ovarian maturation and fertility) and survival in response to cold treatments with summer-like and winter-like photoperiods. We also compared these traits between native (Yokadouma, Africa) and invasive (Yuto, South America) range wild-derived flies. We showed that Z. indianus females have the ability to arrest ovarian maturation and maintain fertility following recovery from cold stress. The critical temperature for ovarian maturation of this species was estimated at c. 13 °C, an intermediate value between those of tropical and temperate drosophilid species. Wild-derived females from Yuto responded to winter-like photoperiod by slowing down ovarian maturation at low but permissive temperatures of 14 °C and 16 °C and also delayed the start of oviposition after cold treatment. Yuto flies also survived better and recovered 20% faster from chill coma than flies from Yokadouma. These results are consistent with a scenario of local adaptations or phenotypic plasticity in the invaded range, and suggest that photoperiod could act as modulator of ovarian arrest. Conversely, the fact that native range flies showed higher fertility after cold recovery than females from invaded range is not indicative of local adaptation. All in all, our findings report a set of physiological responses that would enable Z. indianus expansion to temperate and cold areas, but also results that are compatible with a limitation to the invasion process.     

Marginal range expansion in a host-limited butterfly species Gonepteryx rhamni

1. The British distribution of the butterfly Gonepteryx rhamni (L.) follows closely the range of its natural host plants, Rhamnus catharticus L. and Frangula alnus Miller, suggesting that it is one of the few British butterflies that has a host‐limited distribution. In North Wales, this species has its range margin, and it was recorded only occasionally in a 35‐km² area prior to the 1980s. Frangula alnus bushes were planted in the area in about 1986, allowing the hypothesis that G. rhamni would expand its range following increased host plant availability to be tested. 2. From 1996 to 1998, the distribution of the butterfly and its host plants, R. catharticus (native), Rhamnus alaternus L. (introduced), and F. alnus (introduced to the area but native to Britain), was mapped in the study area. It was found that the butterfly was more widespread than any of its host plants. Frangula alnus was the most widespread of the host plants, and received most eggs, suggesting that the carrying capacity of the habitat would have increased substantially following the planting of this species. Gonepteryx rhamni was able to complete its lifecycle on both introduced species in the study area. 3. A mark–release–recapture study showed that adult G. rhamni moved an average of 512 m, and 50% of movements were further than 400 m; these values are underestimates. The relatively high mobility of this species suggests that it probably perceives host plants and nectar sources as resource patches (patchy population) in this fragmented landscape, and this population now represents a satellite population of the butterfly's main distribution in Britain. 4. The results presented here confirm empirically the host‐limited distribution of G. rhamni, which expanded following the planting of extra host plants.     

Does daily climate variation have an effect on species’ elevational range size?

In their recent paper published in Science (2016, 351 , 1437–1439), Chan et al. analysed 137 montane gradients, concluding that they found a novel pattern—a negative relationship between mean elevational range size of species and daily temperature variation, which was claimed as empirical evidence for a novel macrophysiological principle (Gilchrist's hypothesis). This intriguing possibility was their key conceptual contribution. Unfortunately, as we show, the empirical evidence was flawed because of errors in the analyses and substantial sampling bias in the data. First, we re‐ran their analyses using their data, finding that their model should have been rejected. Second, we performed two additional re‐analyses of their data, addressing biases and pseudoreplication in different ways, both times again rejecting the evidence claimed to support Gilchrist's hypothesis. These results overturn the key empirical findings of Chan et al.'s study. Therefore, the “macrophysiological principle” should be regarded as currently remaining unsupported by empirical evidence.     

Experimental studies of adaptation in Clarkia xantiana. II. Fitness variation across a subspecies border

Because the range boundary is the locale beyond which a taxon fails to persist, it provides a unique opportunity for studying the limits on adaptive evolution. Adaptive constraints on range expansion are perplexing in view of widespread ecotypic differentiation by habitat and region within a species' range (regional adaptation) and rapid evolutionary response to novel environments. In this study of two parapatric subspecies, Clarkia xantiana ssp. xantiana and C. x. ssp. parviflora, we compared the fitness of population transplants within their native region, in a non-native region within the native range, and in the non-native range to assess whether range expansion might be limited by a greater intensity of selection on colonists of a new range versus a new region within the range. The combined range of the two subspecies spans a west-to-east gradient of declining precipitation in the Sierra Nevada of California, with ssp. xantiana in the west being replaced by ssp. parviflora in the east. Both subspecies had significantly higher fitness in the native range (range adaptation), whereas regional adaptation was weak and was found only in the predominantly outcrossing ssp. xantiana but was absent in the inbreeding ssp. parvifilora. Because selection intensity on transplants was much stronger in the non-native range relative to non-native regions, there is a larger adaptive barrier to range versus regional expansion. Three of five sequential fitness components accounted for regional and range adaptation, but only one of them, survivorship from germination to flowering, contributed to both. Flower number contributed to regional adaptation in ssp. xantiana and fruit set (number of fruits per flower) to range adaptation. Differential survivorship of the two taxa or regional populations of ssp. xantiana in non-native environments was attributable, in part, to biotic interactions, including competition, herbivory, and pollination. For example, low fruit set in ssp. xantiana in the east was likely due to the absence of its principal specialist bee pollinators in ssp. parviflora's range. Thus, convergence on self-fertilization may be necessary for ssp. xantiana to invade ssp. parviflora's range, but the evolution of outcrossing would not be required for ssp. parviflora to invade ssp. xantiana's range.     

Widespread latitudinal asymmetry in the performance of marginal populations: A meta-analysis

Aim

Range shifts are expected to occur when populations at one range margin perform better than those at the other margin, yet no global trend in population performances at range margins has been demonstrated empirically across a wide range of taxa and biomes. Here we test the prediction that, if impacts of ongoing climate change on performance in marginal populations are widespread, then populations from the high-latitude margin (HLM) should perform as well as or better than central populations, whereas low-latitude margin (LLM) populations should perform worse.

Location

Global.

Time period

1995–2019.

Major taxa studied

Plants and animals.

Methods

To test our prediction, we used a meta-analysis to quantify empirical support for asymmetry in the performance of high- and low-latitude margin populations compared to central populations. Performance estimates (survival, reproduction, or lifetime fitness) for populations occurring in their natural environment were derived from 51 papers involving 113 margin-centre comparisons from 54 species and 705 populations from the Americas, Europe, Africa and Australia. We then related these performance differences to climatic differences among populations. We also tested whether patterns are consistent across taxonomic kingdoms (plants vs animals) and across realms (marine vs terrestrial).

Results

Populations at margins performed significantly worse than central populations, and this trend was primarily driven by the low-latitude margin. Although the difference was of small magnitude, it was largely consistent across biological kingdoms and realms. Differences in performance were weakly (p = .08) related to the difference in average temperatures between central and marginal populations.

Main conclusions

The observed asymmetry in performance in marginal populations is consistent with predictions about the effects of global climate change, though further research is needed to confirm the effect of climate. It indicates that changes in demographic rates in marginal populations can serve as early-warning signals of impending range shifts.     

What limits male range sizes at different population densities? Evidence from three populations of water voles

In many mammalian species, females compete with each other for food and space to raise offspring, while males compete with each other for access to females. Few studies have examined the factors which limit male range sizes or the degree of overlap between male ranges. We deduced four possible responses of the range -sizes of non-territorial male small mammals to increasing population density and/or levels of forage abundance. These were: (1) male range sizes might remain the same irrespective of population density; (2) at high population densities males may become territorial, and their ranges small and non-overlapping; (3) at high forage abundance/population densities, males' ranges may become smaller but remain intra-sexually overlapping; (4) at high forage abundance, male range sizes may increase. We examined the relationship between population density, range lengths and range overlaps and body weights of both sexes in a comparative study of three populations of water voles Arvicola terrestris . Male range sizes were smaller at higher population densities, but their ranges remained both inter- and intra-sexually overlapping. Heavier males had larger ranges than did lighter males at all sites. These results comply with what would be expected if male range sizes were at least partially restricted by the number of ranges of other individuals with which they overlapped. Although we could not discount the hypothesis that forage abundance may also have had a direct effect on male range sizes, our results implied that male range sizes were at least partially determined by social factors.     

Techniques for evaluating the spatial behaviour of river fish

Radio-tagging is widely used for studies of movements, resource use and demography of land vertebrates, with potential to combine such data for predictive modelling of populations from individuals. Such modelling requires standard measures of individual space use, for combination with data on resources, survival, dispersal and breeding. This paper describes how protocols for efficient collection of space-use data can be developed during a pilot study, and reviews the ways in which such data can be used for space-use indices that help answer biological questions, with examples from a study of riverine pike (Esox lucius). Analyses of diurnal activity and spatio-temporal correlation were used to assess when to record locations, and analyses of home range increments were used to define the number of location records necessary to assess seasonal ranges. We stress the importance of developing protocols that use minimal numbers of locations from each individual, so that analyses can be based on samples of many individuals. The efficacy of link-distance (e.g. cluster analysis) and location density (e.g. contouring) techniques for spatial analysis for river fish were compared, and the utility of clipping off areas to river banks was assessed. In addition, a new automated analysis was used to estimate distances along river mid-lines. These techniques made it possible to quantify interactions between individuals and their habitat: including a significant increase in core range size during floods, significant preference for deep pools, and a lack of exclusive territories.     

Racing against change: understanding dispersal and persistence to improve species' conservation prospects

Climate change is contributing to the widespread redistribution, and increasingly the loss, of species. Geographical range shifts among many species were detected rapidly after predictions of the potential importance of climate change were specified 35 years ago: species are shifting their ranges towards the poles and often to higher elevations in mountainous areas. Early tests of these predictions were largely qualitative, though extraordinarily rapid and broadly based, and statistical tests distinguishing between climate change and other global change drivers provided quantitative evidence that climate change had already begun to cause species’ geographical ranges to shift. I review two mechanisms enabling this process, namely development of approaches for accounting for dispersal that contributes to range expansion, and identification of factors that alter persistence and lead to range loss. Dispersal in the context of range expansion depends on an array of processes, like population growth rates in novel environments, rates of individual species movements to new locations, and how quickly areas of climatically tolerable habitat shift. These factors can be tied together in well-understood mathematical frameworks or modelled statistically, leading to better prediction of extinction risk as climate changes. Yet, species'' increasing exposures to novel climate conditions can exceed their tolerances and raise the likelihood of local extinction and consequent range losses. Such losses are the consequence of processes acting on individuals, driven by factors, such as the growing frequency and severity of extreme weather, that contribute local extinction risks for populations and species. Many mechanisms can govern how species respond to climate change, and rapid progress in global change research creates many opportunities to inform policy and improve conservation outcomes in the early stages of the sixth mass extinction.     

Measuring range sizes of South-East Asian hawkmoths (Lepidoptera: Sphingidae): effects of scale, resolution and phylogeny

Aim  The effects of resolution and spatial extent on range measures were explored in estimates of the geographic distribution of tropical hawkmoths. Furthermore, data were tested for phylogenetic autocorrelation.
Location  South-East Asia.
Methods  Various range measures, such as geographic information system (GIS)-supported range estimates, minimum convex polygons, latitudinal and longitudinal extents, and their products, were derived from original distribution records and compared to each other. A taxonomic classification of the species was used to analyse phylogenetic effects on range sizes.
Results  Range size measures exhibit a strongly right-skewed frequency distribution with many geographically restricted species and few widespread taxa. Rankings from GIS-supported, comprehensive range size estimates do not deviate greatly from more crude measurements of lower resolution. Comprehensive ranges and ranges within South-East Asia are correlated strongly, but already at this rather large scale the ranking of species changes considerably. Other measures of occupancy with an increasingly more localized consideration of 'range' show decreasing strengths of correlation. We found a weak, but significant, autocorrelation in range area data: related groups of species have ranges of similar size.
Main conclusions  Spatial resolution did not affect range ranking greatly in our data. However, macroecological studies based only on parts of species' ranges must be viewed critically, particularly if their extent is small compared to comprehensive ranges. Phylogenetic non-independence of range size data must be considered in comparative analyses.     

T-even type phages can change their host range by recombination with gene 34 (tail fibre) or gene 23 (head)

Summary T-even type phages recognize their cellular receptors with the tip of their long tail fibres. The gene products involved in receptor recognition are proteins 37 and 38. While screening libraries of phage K3 with a probe of gene38 from phage T2, a class of weakly hybridizing clones was found in addition to the expected clones of gene38 of K3. One of these clones was identified as being from gene23 of the phage which codes for the major head subunit; another clone originated from gene34, which codes for the proximal half of the long tail fibres. Neither gene product 23 nor 34 is involved in receptor recognition. Phages can recombine with the DNA of the gene23 and gene34 clones and change the host range. This paper is dedicated to Prof. Dr. Wolfgang Beermann, Tübingen, on the occation of his 65^th birthday