The effect of the spatial configuration of habitat fragmentation on invasive spread

To address how the spatial configuration of habitat fragmentation influences the persistence and the rate of spread of an invasive species, we consider three simple periodically fragmented environments, a lattice-like corridor environment, an island-like environment and a striped environment. By numerically analyzing Fisher’s equation with a spatially varying diffusion coefficient and the intrinsic growth rate, we find the following. (1) When the scale of fragmentation is sufficiently large, the minimum favorable area needed for successful invasion reduces in the following order: lattice-like corridor, striped and island-like environments. (2) When the scale of fragmentation and the fraction of favorable area are sufficiently large, the spreading speeds along contiguous favorable habitats in the lattice-like corridor and striped environments are faster than the speeds across isolated favorable habitats in the island-like environment and the striped environment. (3) When the periodicity of fragmentation is relaxed by stochastically shifting the boundaries between favorable and unfavorable habitats, the average speed increases with increases in the irregularity of fragmentation.     

Impact of Directed Movement on Invasive Spread in Periodic Patchy Environments

To address the effect of taxis of invasive animals on their spreading speed in heterogeneous environments, we deal with an advection-diffusion-reaction equation (ADR) in a periodic patchy environment. Two-types of advection that spatially vary depending on environmental heterogeneity are taken into consideration: a stepwise taxis function and a saw-like taxis function. We first analyze the ADR with the stepwise taxis advection, and derive an invasion criterion. When the invasion criterion holds, an initially localized population evolves to a traveling periodic wave (TPW). The asymptotic speed of the TPW is found to be equal to the minimal speed of the TPW analytically derived. Thus, we examine how the minimal speed is influenced by the taxis. The major results are: (1)?As the magnitude of the taxis toward favorable patches increases, invasion becomes more feasible. However, the spreading speed increases at first, and then decreases to show a one-humped curve against the magnitude of the taxis; (2)?As the scale of fragmentation in the patchy environment is increased, the spreading speed increases when the magnitude of the taxis is small, while it decreases when the magnitude of the taxis becomes sufficiently large. These characteristic features qualitatively apply to the ADR model with the saw-like taxis function.     

Genetic diversity in butterflies: Interactive effects of habitat fragmentation and climate-driven range expansion

Some species are expanding their ranges polewards during current climate warming. However, anthropogenic fragmentation of suitable habitat is affecting expansion rates and here we investigate interactions between range expansion, habitat fragmentation and genetic diversity. We examined three closely related Satyrinae butterflies, which differ in their habitat associations, from six sites along a transect in England from distribution core to expanding range margin. There was a significant decline in allozyme variation towards an expanding range margin in Pararge aegeria, which has the most restricted habitat availability, but not in Pyronia tithonus whose habitat is more widely available, or in a non-expanding 'control species' (Maniola jurtina). Moreover, data from another transect in Scotland indicated that declines in genetic diversity in P. aegeria were evident only on the transect in England, which had greater habitat fragmentation. Our results indicate that fragmentation of breeding habitats leads to more severe founder events during colonization, resulting in reduced diversity in marginal populations in more specialist species. The continued widespread loss of suitable habitats in the future may increase the likelihood of loss of genetic diversity in expanding species, which may affect whether or not species can adapt to future environmental change.     

Modelling and analysing evolution of dispersal in populations at expanding range boundaries

Abstract 1. Species would be expected to shift northwards in response to current climate warming, but many are failing to do so because of fragmentation of breeding habitats. Dispersal is important for colonisation and an individual‐based spatially explicit model was developed to investigate impacts of habitat availability on the evolution of dispersal in expanding populations. Model output was compared with field data from the speckled wood butterfly Pararge aegeria, which currently is expanding its range in Britain. 2. During range expansion, models simulated positive linear relationships between dispersal and distance from the seed location. This pattern was observed regardless of quantity (100% to 10% habitat availability) or distribution (random vs. gradient distribution) of habitat, although higher dispersal evolved at expanding range margins in landscapes with greater quantity of habitat and in gradient landscapes. Increased dispersal was no longer evident in any landscape once populations had reached equilibrium; dispersal values returned to those of seed populations. However, in landscapes with the least quantity of habitat, reduced dispersal (below that of seed populations) was observed at equilibrium. 3. Evolutionary changes in adult flight morphology were examined in six populations of P. aegeria along a transect from the distribution core to an expanding range margin in England (spanning a latitudinal distance of >200 km). Empirical data were in agreement with model output and showed increased dispersal ability (larger and broader thoraxes, smaller abdomens, higher wing aspect ratios) with increasing distance from the distribution core. Increased dispersal ability was evident in populations from areas colonised >30 years previously, although dispersal changes were generally evident only in females. 4. Evolutionary increases in dispersal ability in expanding populations may help species track future climate changes and counteract impacts of habitat fragmentation by promoting colonisation. However, at the highest levels of habitat loss, increased dispersal was less evident during expansion and reduced dispersal was observed at equilibrium indicating that, for many species, continued habitat fragmentation is likely to outweigh any benefits from dispersal.     

Tetragonisca guard bees interpret expanding and contracting patterns as unintended displacement in space

Guard bees of the stingless bee Tetragonisca angustula (Apidae: Meliponinae) hover in stable positions in front of the nest to protect the flight corridor leading to the nest entrance against insect intruders. To unravel the visual control of station keeping, we exposed these hovering guards to expanding and contracting patterns at the nest front. The bees fly away from an expanding pattern and towards the centre of a contracting pattern along a line connecting their initial position and the centre of expansion regardless of where in the visual field they view the pattern. The response of bees to a spinning radial pattern is different: they fly parallel to the pattern, up and down or forward and backward depending on whether they initially hover to the side, above or below the centre of rotation. The bees respond to horizontal and to vertical expansion and contraction. They also adjust their distance relative to a rotating spiral which produces a realistic flow field and thus allowed us to test to what extent the bees minimize image motion speed. We find that guard bees indeed move in the appropriate direction to minimize the image motion speed they experience. A comparison of bees hovering at different distances from the nestfront at the onset of pattern motion and experiencing very different image velocities shows that the dynamics of the reaction is quite uniform. At the pattern velocities tested, we did not find evidence that guard bees use image motion to control their flight speed. The bees' response rather suggests that the underlying mechanism might be insensitive to the size of motion vectors. Accepted: 2 April 1997     

Can a Species Keep Pace with a Shifting Climate?

Consider a patch of favorable habitat surrounded by unfavorable habitat and assume that due to a shifting climate, the patch moves with a fixed speed in a one-dimensional universe. Let the patch be inhabited by a population of individuals that reproduce, disperse, and die. Will the population persist? How does the answer depend on the length of the patch, the speed of movement of the patch, the net population growth rate under constant conditions, and the mobility of the individuals? We will answer these questions in the context of a simple dynamic profile model that incorporates climate shift, population dynamics, and migration. The model takes the form of a growth-diffusion equation. We first consider a special case and derive an explicit condition by glueing phase portraits. Then we establish a strict qualitative dichotomy for a large class of models by way of rigorous PDE methods, in particular the maximum principle. The results show that mobility can both reduce and enhance the ability to track climate change that a narrow range can severely reduce this ability and that population range and total population size can both increase and decrease under a moving climate. It is also shown that range shift may be easier to detect at the expanding front, simply because it is considerably steeper than the retreating back.     

Fragmentation of clones: how does it influence dispersal and competitive ability?

We applied individual-based simulations to study the effect of physiological integration among ramets in clonal species that live in patchy habitats. Three strategies were compared: (1) Splitter, in which the genet was fragmented into independent ramets; (2) Transient Integrator, where only groups of ramets were connected; and (3) Permanent Integrator, in which fragmentation did not occur, and the whole genet was integrated. We studied the dynamics of spatial spreading and population growth in these strategies separately and in competition. Various habitat types were modeled by changing the density of favorable habitat patches. We found that the spatial pattern of good patches significantly influenced the growth of the populations. When the resource patches were scarce, a large proportion of the carrying capacity of the habitat was not utilized by any of the strategies. It was the Splitter that proved to be the most severely dispersal-limited. But at the same time, it could compete for the good patches most efficiently. The balance between these two contradictory effects was largely determined by the proportion of favorable to unfavorable areas. When this proportion was low or intermediate (up to ca. 50% good), integration was more advantageous. At higher proportions, fragmentation became beneficial. Fragmentation into groups of ramets (Transient Integration) was not sufficient, only radical splitting could ensure a significant selective advantage. Transient Integrators got fragmented according to the spatial pattern of ramet mortality. It was interesting that the enrichment of the area in good sites did not lead to larger fragment sizes. It merely raised the number of fragments. Nevertheless, these small fragments were more similar to integrated genets (in the Permanent Integrator) than to solitary ramets (in the Splitter) in terms of dispersal and competitive ability. This suggests that even a slightly integrated clonal species can be ecologically considered as an integrator.     

Woodland fragmentation affects space use of Eurasian red squirrels

When habitats become fragmented, variation in patch size and quality are expected to impose changes on the spacing pattern and social organization of animal populations. General theory predicts different possible responses including shrinking home ranges (fission response), increasing range overlap (fusion) and incorporation of multiple patches in the home range (expansion response) as fragmentation increases. We studied space use and social organization in a metapopulation of red squirrels (Sciurus vulgaris) in 15 woodland fragments differing in size and tree species composition. Home ranges and core areas of males were larger than females, and fragmentation had different and complex effects on the spacing pattern of both sexes. In food-supplemented patches, high densities led to increased intra-sexual overlap. In linear-shaped patches, squirrels used smaller home ranges and core areas and had lower male–male and male–female overlap levels, independent of patch quality or size. Home range and core area size of males increased with patch size, and male core areas overlapped extensively those of other males and females. Hence males seemed to show a fission response only in some patches. In contrast, home range and core area size of females was not related with patch size, but decreased with habitat quality, supporting predictions of a fusion response and intra-sexual defense of food-based core areas. Hence, where patch size and shape strongly affected space use of male red squirrels, social organization of females was only affected in small, food-supplemented patches, suggesting that the basic spatio-social organization of adult females is very resistant to fragmentation.     

Separating historic events from recurrent processes in cryptic species: phylogeography of mud snails (Hydrobia spp.)

The present study combines methods that were designed to infer intraspecific relationships (e.g. nested-clade analysis (NCA), mismatch distributions and maximum likelihood gene flow analysis) to analyse historic events and recurrent processes in the cryptic mud snail species Hydrobia acuta and H. glyca. Specifically, we test the proposed allopatry of cryptic species and whether the peculiar range-subdivision of the putative subspecies H. a. acuta and H. a. neglecta is a result of long-distance dispersal or continuous range expansion. The NCA indicates a past fragmentation of the two H. acuta subspecies as well as past fragmentations within H. glyca. Gene-flow analyses show extensive gene flow in an E-W direction (towards the Atlantic) in the Mediterranean H. a. acuta, generally low gene flow in a W-E direction in the Atlantic H. a. neglecta and complex gene-flow pattern in a N-S but also in a S-N direction (against the Gulf Stream) in H. glyca. Based on these data and supportive ecological and oceanographical data, we hypothesize that the separation of the two H. acuta subspecies was not caused by long-distance dispersal but by a range shift and/or range expansion of the closely related competitor H. glyca as a result of an interglacial warming with a subsequent range shift in H. acuta. Moreover, our data do not show evidence for a long-term, stable sympatry of Hydrobia species, supporting the concept of allopatric relationships within cryptic radiations. NCA and gene-flow analyses indicate that the only sympatric population found in our study is the result of a recent dispersal event from the nearby Mediterranean. It is assumed that allopatric relationships in ephemeral Hydrobia populations constitute an evolutionary advantage relative to competition, recruitment and re-establishment of habitats. Mechanisms that could be of relevance for maintaining allopatry are discussed.     

The effect of the last glacial age on speciation and population genetic structure of the endangered Ethiopian wolf (Canis simensis)

During the last glacial age, Afro-alpine habitats were widespread across the highlands of Ethiopia. A wolf-like canid ancestor is thought to have colonized this expanding habitat and given rise to a new species that was remarkably well adapted to the high altitude environment: the Ethiopian wolf Canis simensis. Here, we address the timing of genetic divergence and examine population genetic history and structure by investigating the distribution of mitochondrial DNA (mtDNA) sequence variation. The pattern of mtDNA variation and geographical distribution indicate an initial population expansion, probably immediately after divergence from the wolf-like ancestor, around 100,000 years ago. The partition of mtDNA haplotypes that followed was most likely the result of habitat reduction and fragmentation at the onset of deglaciation approximately 15,000 years ago. Phylogenetic and geographical associations suggest that the most likely genetic partitioning corresponds to three mountain areas, Arsi/Bale, Wollo/Shoa and Simien/Mt. Guna. Although there is a degree of clustering of haplotypes from both sides of the Rift Valley, the lack of reciprocal monophyly does not support the taxonomic classification of two subspecies. This study highlights the importance of populations north of the Rift Valley for the maintenance of genetic variability within the species and has consequent implications for conservation.     

Strategic Escape Direction: Orientation,Turning, and Escape Trajectories of Zebra‐Tailed Lizards (Callisaurus draconoides)

A prey's body orientation relative to a predator's approach path may affect risk of fleeing straight ahead. Consequently, prey often turn before fleeing. Relationships among orientation, turn, and escape angles and between these angles and predation risk have not been studied in terrestrial vertebrates and have rarely been studied in the field. Escape angles are expected to lead away from predators and be highly variable to avoid being predictable by predators. Using approach speed as a risk factor, we studied these issues in the zebra‐tailed lizard, Callisaurus draconoides. Lizards fled away from human simulated predators, but most did not flee straight away. Escape angles were variable, as expected under the unpredictability hypothesis, and had modes at nearly straight away (i.e., 0°) and nearly perpendicular to the predator's approach path (90°). The straight away mode suggests maximal distancing from the predator; the other mode suggests maintaining ability to monitor the predator or possibly an influence of habitat features such as obstacles and refuges that differ among directions. Turn angles were larger when orientation was more toward the predator, and escape angles were closer to straight away when turn angles were larger. Turning serves to reach a favorable fleeing direction. When orientation angle was more toward the predator, escape angle was unaffected, suggesting that turn angle compensates completely for increased risk of orientation toward the predator. When approached more rapidly, lizards fled more nearly straight away, as expected under greater predation risk. Turn angles were unrelated to approach speed.     

1980-2015年珠三角城市群城市扩张的时空特征分析

以珠三角城市群9座城市为研究对象,基于1980-2015年3期遥感影像提取的土地利用数据,通过扩张强度、扩张速率以及景观格局指数定量分析珠三角城市群尺度上城市扩张的时空变化特征。结果表明：（1）城市群扩张数量特征：1980-2015年珠三角城市群建设用地面积扩张约3倍,1980-2000和2000-2015期间分别扩张1513.1、3043.8 km²;扩张强度不断增强,扩张速率明显加快。（2）城市群扩张空间特征：珠三角城市群中扩张迅速的城市主要分布在研究区中部的城镇密集带及沿海地区;扩张较慢的城市主要分布在靠内陆的周边地带。（3）基于时间尺度发现珠三角城市群结构的演变呈现出由单核模式（广州,1980年）-双核模式（广州、深圳,2000年）-多中心、网络化模式演化的特征（东莞、中山等相邻城市,2015年）。（4）基于景观格局特征,珠三角区域扩张快速的城市形状复杂,结构趋于分散;扩张较慢的城市形状更规则,分布更聚集。（5）天然的人文联系与优越的地理位置、社会经济、政府政策以及基础设施的建设等因素共同驱动珠三角城市群的发展。该研究对提高城市群规划管理水平具有参考价值。     

Initial conditions and their effect on invasion velocity across heterogeneous landscapes

Accurate, time dependent control options are required to halt biological invasions prior to equilibrium establishment, beyond which control efforts are often impractical. Although invasions have been successfully modeled using diffusion theory, diffusion models are typically confined to providing simple range expansion estimates. In this work, we use a Susceptible/Infected cellular automaton (CA) to simulate diffusion. The CA model is coupled with a network model to track the speed and direction of simulated invasions across heterogeneous landscapes, allowing for identification of locations for targeted control in both time and space. We evaluated the role of the location of initial establishment insofar as it affected the pattern and rate of spread and how these are influenced by patch attributes such as size. Our results show that the location of initial establishment can significantly affect the temporal dynamics of an invasion. Traditional network metrics such as degree and measures of topological distance were insufficient for predicting the direction and speed of the invasion. Our coupled models allow the dynamic tracking of invasions across fragmented landscapes for both theoretical and practical applications.     

黄土高原人口密集区城镇扩张对生境质量的影响——以兰州、西安-咸阳及太原为例

快速城镇化过程极大改变了区域生境的空间格局和功能要素,深刻影响生境斑块之间的物质流和能量流,同时对生境和生物多样性造成严重威胁.本研究采用InVEST模型、景观指数及多元线性回归,系统分析了1990-2018年黄土高原全域及人口密集区城镇化对生境质量影响的时空格局特征与因素.结果表明:黄土高原城镇扩张显著影响生境质量,...     

不同栖息地破坏格局下具捕食偏爱似然竞争结局的模拟分析

使用元胞自动机模型,对具有捕食偏爱、不同栖息地破坏比例和不同空间破坏格局条件下的捕食-食饵系统中各物种的变化动态进行了模拟分析。在捕食者和两猎物物种共存时:栖息地破坏比例、栖息地破坏的聚集度对猎物物种间强弱关系产生相反的作用,若增加栖息地破坏比例不利于某一猎物生存,则提高聚集度对其有利;适当提高适宜栖息地的聚集度,对所有物种都有利,若聚集度过高,效果相当于减少了栖息地的破坏比例,可能对某些猎物物种不利,但对整体系统有利;被破坏栖息地的聚集度发生变化时,捕食者的反应更敏感;在一定条件下,增强弱势种群的捕食偏爱会有助于其生存。     

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.     

Population structure of pioneer specialist solitary bee Andrena vaga (Hymenoptera: Andrenidae) in central Europe: the effect of habitat fragmentation or evolutionary history?

Because patchiness of food sources or nesting opportunities frequently limits gene flow, specialists often exhibit distinct population structures in fragmented habitats. We studied the influence of habitat fragmentation on population structure in the solitary bee Andrena vaga, an early spring species that nests exclusively in sandy soil and feeds strictly on willows (Salix spp.). Because the homogenous habitat of the German floodplains, where the species was studied previously, resulted in the species’ weak population structure, we expected more structured populations in central Europe, where the sandy soils essential for nesting are highly fragmented. We analysed 387 females from 21 localities in the Czech Republic and Slovakia using nine microsatellite loci, and we inferred population structure using landscape genetics and Bayesian clustering methods. Contrary to our expectations, habitat fragmentation did not result in increased genetic isolation at the localities; however, two differentiated groups of localities, separated by a wide clinal zone of admixture, were detected within the study area. The observed pattern suggests that dispersive ability of A. vaga compensates the species dependence on unstable fragmented habitats. We propose that the population structure may mirror a secondary contact formed by the expansion of two populations that had been separated in the past. We emphasise the necessity of knowing the studied species’ population history before making conclusions concerning correlations between habitat and population structure, especially in areas of known suture zones created by the secondary contact of populations expanding from separate refugia.     

Changes in habitat associations during range expansion: disentangling the effects of climate and residence time

The distributions of many species are not at equilibrium with their environment. This includes spreading non-native species and species undergoing range shifts in response to climate change. The habitat associations of these species may change during range expansion as less favourable climatic conditions at expanding range margins constrain species to use only the most favourable habitats, violating the species distribution model assumption of stationarity. Alternatively, changes in habitat associations could result from density-dependent habitat selection; at range margins, population densities are initially low so species can exhibit density-independent selection of the most favourable habitats, while in the range core, where population densities are higher, species spread into less favourable habitat. We investigate if the habitat preferences of the non-native common waxbill Estrilda astrild changed as they spread in three directions (north, east and south-east) in the Iberian Peninsula. There are different degrees of climatic suitability and colonization speed across range expansion axes, allowing us to separate the effects of climate from residence time. In contrast to previous studies we find a stronger effect of residence time than climate in influencing the prevalence of common waxbills. As well as a strong additive effect of residence time, there were some changes in habitat associations, which were consistent with density-dependent habitat selection. The combination of broader habitat associations and higher prevalence in areas that have been colonised for longer means that species distribution models constructed early in the invasion process are likely to underestimate species’ potential distribution.     

Chemokinetic behavior of the infective third-stage larvae of Strongyloides ratti on a sodium chloride gradient.

The movements of the infective third-stage larvae (L3) of a rodent parasitic nematode Strongyloides ratti were examined on a sodium chloride (NaCl) gradient set up on agarose plates. The movements of larvae were followed by observing their tracks on the surface of the agarose. The direction of movement depended on the NaCl concentration at the point of their initial placement on the gradient. Larvae placed at between 230 and 370 mM NaCl tended to migrate towards areas of lower concentration. On the other hand, when placed at concentrations less than 20 mM NaCl, larvae tended to migrate initially towards higher concentrations but did not linger in areas where the concentration was over approximately 80 mM NaCl. It seems that S. ratti L3, tested in vitro, prefer regions with a concentration of NaCl below 80 mM NaCl. Two typical chemokinetic behaviors are seen; a unidirectional avoidance movement when initially placed in unfavorable environmental conditions and a random dispersal movement when placed within an area of favorable conditions. Track patterns were straight in the avoidance movement but included multiple changes of direction and loops in the dispersal movement. This study introduces an assay system suitable for studying chemokinetic behavior of larvae of Strongyloides ratti.     

Utilizing a multifaceted approach to assess the current distribution and conservation status of an uncommon species: the golden mouse (Ochrotomys nuttalli) in Florida

Aim

Our goal was to assess the conservation status of the understudied and naturally uncommon habitat specialist, the golden mouse (Ochrotomys nuttalli), at the edge of its range where its historically fragmented habitat has been subjected to severe loss.

Location

Peninsular Florida, north of approximately 27° latitude, USA.

Methods

We used data gathered from museum collections, regional biologists, geographic information systems (GIS) layers, field surveys and DNA sequencing to determine the habitats that best explain the distribution of the species, examine changes in the geographic extent of both the species and its habitats, and compare genetic differentiation between populations occupying disjunct regions. The results from these multiple analyses were combined to assess the conservation status of the species.

Results

Golden mouse occurrence records align well with the distribution of hardwood habitats in Florida. These habitats occur naturally as ‘islands’, but have become increasingly fragmented by anthropogenic land use. Despite habitat loss, the location of the southern range periphery has remained relatively unchanged in location over the past century. Genetic analysis reveals a history of limited dispersal of females among habitat ‘islands’ that likely predates anthropogenic landscape fragmentation. This pattern suggests that isolated populations that are extirpated will have little to no chance of successful recolonization.

Main conclusions

The combined results from multiple analyses produced a more complete picture of the threats faced by this previously data‐deficient species than any single analysis would have. Although the species' southern range limit cannot be shown to have retracted in the face of human expansion, habitat fragmentation clearly has put the species at increased risk. Conservation and management of hardwood habitats are critical to the persistence of the golden mouse at the edge of its range.