Inter-population Movement Research Articles

Changing-look Active Galactic Nuclei. I. Tracking the Transition on the Main Sequence of Quasars

This paper is the first in a series of papers that prepare and analyze spectral and other properties for a database of already-discovered changing-look active galactic nuclei (CL AGNs). Here we focus on the spectral fitting and analysis of broad emission lines in a sample of 93 CL AGNs collected from the literature with existing Sloan Digital Sky Survey (SDSS)/Baryon Oscillation Spectroscopic Survey (BOSS)/extended-BOSS spectroscopy where the Hβ emission-line profile does not completely disappear in any epochs. Additionally, we have gathered older/newer spectral epochs from all the available SDSS data releases to make the database more complete. We use PyQSOFit and perform a homogeneous spectral decomposition of all of our SDSS spectra and tabulate the AGN continuum and emission-line properties per epoch per source, chronologically. This further allows us to categorize the sources in our sample as Turn-On or Turn-Off and subsequently check for repeated occurrences of such phases. We then estimate the black hole mass (M BH) and the Eddington ratio (λ Edd) per epoch per source where the required parameters are available and well estimated. We demonstrate the movement of the source in the M BH–λ Edd plane, allowing us to check for systematic changes in the source’s fundamental properties. We then track their transition along the optical plane of the Eigenvector 1 schema and categorize sources that either stay within the same population (A or B) or make an interpopulation movement as a function of spectral epoch. We also test the Balmer decrement (Hα/Hβ) of a subset of our sample of CL AGNs as a function of time and AGN luminosity.

Using conservation genetics to prioritise management options for an endangered songbird

Genetic data can be highly informative for answering questions relevant to practical conservation efforts, but remain one of the most neglected aspects of species recovery plans. Framing genetic questions with reference to practical and tractable conservation objectives can help bypass this limitation of the application of genetics in conservation. Using a single-nucleotide polymorphism dataset from reduced-representation sequencing (DArTSeq), we conducted a genetic assessment of remnant populations of the endangered forty-spotted pardalote (Pardalotus quadragintus), a songbird endemic to Tasmania, Australia. Our objectives were to inform strategies for the conservation of genetic diversity in the species and estimate effective population sizes and patterns of inter-population movement to identify management units relevant to population conservation and habitat restoration. We show population genetic structure and identify two small populations on mainland Tasmania as ‘satellites’ of larger Bruny Island populations connected by migration. Our data identify management units for conservation objectives relating to genetic diversity and habitat restoration. Although our results do not indicate the immediate need to genetically manage populations, the small effective population sizes we estimated for some populations indicate that they are vulnerable to genetic drift, highlighting the urgent need to implement habitat restoration to increase population size and to conduct genetic monitoring. We discuss how our genetic assessment can be used to inform management interventions for the forty-spotted pardalote and show that by assessing contemporary genetic aspects, valuable information for conservation planning and decision-making can be produced to guide actions that account for genetic diversity and increase chances of recovery in species of conservation concern.

Spatial Memory and Taxis-Driven Pattern Formation in Model Ecosystems

Mathematical models of spatial population dynamics typically focus on the interplay between dispersal events and birth/death processes. However, for many animal communities, significant arrangement in space can occur on shorter timescales, where births and deaths are negligible. This phenomenon is particularly prevalent in populations of larger, vertebrate animals who often reproduce only once per year or less. To understand spatial arrangements of animal communities on such timescales, we use a class of diffusion–taxis equations for modelling inter-population movement responses between N ge 2 populations. These systems of equations incorporate the effect on animal movement of both the current presence of other populations and the memory of past presence encoded either in the environment or in the minds of animals. We give general criteria for the spontaneous formation of both stationary and oscillatory patterns, via linear pattern formation analysis. For N=2, we classify completely the pattern formation properties using a combination of linear analysis and nonlinear energy functionals. In this case, the only patterns that can occur asymptotically in time are stationary. However, for N ge 3, oscillatory patterns can occur asymptotically, giving rise to a sequence of period-doubling bifurcations leading to patterns with no obvious regularity, a hallmark of chaos. Our study highlights the importance of understanding between-population animal movement for understanding spatial species distributions, something that is typically ignored in species distribution modelling, and so develops a new paradigm for spatial population dynamics.

Assessing source-sink stability in the context of management and land-use change

Management actions and land-use change can disrupt interdependent population processes, re-define population networks, and change source-sink dynamics. Yet we know little about the types of changes that can de-stabilize source-sink dynamics and how such changes could affect management decisions. We examined the degree to which source-sink status and strength could change under a range of management actions and land-use change scenarios including different patterns and extents of habitat loss, restoration, demographic improvements from parasitism control, and increased frequencies inter-population movement. We developed an empirically-rich, spatially explicit, individual-based model for the formerly endangered Black-capped vireo in Texas. We simulated the network-wide consequences of different kinds of changes and compared the resulting source-sink strength, status, and regional abundance across scenarios. We gauged source-sink stability by the degree to which system changes caused the reversal of source or sink status. The stability of source-sink characterizations differed with the type of change. Source-sink dynamics were less responsive to small changes to population structure and changes that minimally affected demographic conditions. Source-sink status was most responsive to changes that affected habitat patterns and quality. Accurately classifying sources and sinks is challenging, particularly in variable and directionally changing systems. The stability of source-sink classifications depends on the type of management or land-use change. Management actions may need to weigh interventions that improve regional abundance against those that alter regional source-sink dynamics as abundance and source-sink states can be sensitive to different kinds of change.

Genetic evidence for spatial structuring in a continuous African elephant (Loxodonta africana) population

Earlier studies on savannah elephants (Loxodonta africana) investigated the genetic structure of fragmented or isolated populations. Contrastingly, this study aimed to determine if there was genetic evidence for spatial structuring in a continuous elephant population in the Kavango-Zambezi Transfrontier Conservation Area (KAZA-TFCA). We sequenced one mtDNA gene region for 88 individuals and genotyped 100 individuals for ten nuclear microsatellite loci. Bayesian Clustering Algorithms in Geneland identified groups of genetically similar individuals. An analysis of molecular variance determined if these groups (sub-populations) were significantly differentiated. We identified geographic areas with high genetic divergence (genetic barriers) between samples using a GIS landscape genetic toolbox. There were three significantly differentiated mtDNA sub-populations (Fst = 0.787) and two nDNA sub-populations that were not significantly differentiated (Fst = −0.02; Rst = −0.045), implying obstructed mtDNA, but high nDNA gene flow across the study region. The KAZA-TFCA population has a genetic diversity (mtDNA pairwise number of differences (π) = 2.59; nDNA mean alleles/locus and He = 7.5, 0.71) higher than other southern African populations, and inter-population movements may be responsible for maintaining this genetic diversity. We discount anthropogenic and geographic barriers as the primary drivers of genetic structuring in the KAZA-TFCA population and suggest that future studies should consider the influence of intrinsic factors (resource dependencies and social variables that limit movement) when investigating the genetic structure of elephant populations. We recommend continued support for conservation initiatives that aim at maintaining and restoring connectivity between populations, which in so doing may ensure inter-population gene flow and uphold the current genetic state of the KAZA-TFCA population.

Abstracts

Abstracts

Mating effort and space use of an alpine ungulate during the rut

In ungulates, the rut generally leads to increased intra and interpopulation movements for males. Because movements induce energetic costs and missed feeding opportunities, they could be an indication of male mating effort. We studied space use of 44 male mountain goats (Oreamnos americanus) from three neighboring subpopulations in southeast Alaska, during the rut from 2005 to 2008. Using mixed models and an information theoretic approach with AIC, we analyzed the relationships between individual traits of males and their space use. We found no indication of breeding migration between subpopulations. Distances between individual seasonal ranges were not related to any individual trait. Daily movements, home range sizes, and total distance traveled during the rut did not vary with mass or age of individuals. As such, effects of individual traits on male space use during the rut appear weak and observed space use patterns do not support any of the main mating effort hypotheses.

Inter-Population Movements of Steller Sea Lions in Alaska with Implications for Population Separation

Genetic studies and differing population trends support the separation of Steller sea lions (Eumetopias jubatus) into a western distinct population segment (WDPS) and an eastern DPS (EDPS) with the dividing line between populations at 144° W. Despite little exchange for thousands of years, the gap between the breeding ranges narrowed during the past 15–30 years with the formation of new rookeries near the DPS boundary. We analyzed >22,000 sightings of 4,172 sea lions branded as pups in each DPS from 2000–2010 to estimate probabilities of a sea lion born in one DPS being seen within the range of the other DPS (either ‘West’ or ‘East’). Males from both populations regularly traveled across the DPS boundary; probabilities were highest at ages 2–5 and for males born in Prince William Sound and southern Southeast Alaska. The probability of WDPS females being in the East at age 5 was 0.067 but 0 for EDPS females which rarely traveled to the West. Prince William Sound-born females had high probabilities of being in the East during breeding and non-breeding seasons. We present strong evidence that WDPS females have permanently emigrated to the East, reproducing at two ‘mixing zone’ rookeries. We documented breeding bulls that traveled >6,500 km round trip from their natal rookery in southern Alaska to the northern Bering Sea and central Aleutian Islands and back within one year. WDPS animals began moving East in the 1990s, following steep population declines in the central Gulf of Alaska. Results of our study, and others documenting high survival and rapid population growth in northern Southeast Alaska suggest that conditions in this mixing zone region have been optimal for sea lions. It is unclear whether eastward movement across the DPS boundary is due to less-optimal conditions in the West or a reflection of favorable conditions in the East.

Human-caused mortality influences spatial population dynamics: Pumas in landscapes with varying mortality risks

An understanding of how stressors affect dispersal attributes and the contribution of local populations to multi-population dynamics are of immediate value to basic and applied ecology. Puma (Puma concolor) populations are expected to be influenced by inter-population movements and susceptible to human-induced source–sink dynamics. Using long-term datasets we quantified the contribution of two puma populations to operationally define them as sources or sinks. The puma population in the Northern Greater Yellowstone Ecosystem (NGYE) was largely insulated from human-induced mortality by Yellowstone National Park. Pumas in the western Montana Garnet Mountain system were exposed to greater human-induced mortality, which changed over the study due to the closure of a 915km2 area to hunting. The NGYE’s population growth depended on inter-population movements, as did its ability to act as a source to the larger region. The heavily hunted Garnet area was a sink with a declining population until the hunting closure, after which it became a source with positive intrinsic growth and a 16× increase in emigration. We also examined the spatial and temporal characteristics of individual dispersal attributes (emigration, dispersal distance, establishment success) of subadult pumas (N=126). Human-caused mortality was found to negatively impact all three dispersal components. Our results demonstrate the influence of human-induced mortality on not only within population vital rates, but also inter-population vital rates, affecting the magnitude and mechanisms of local population’s contribution to the larger metapopulation.

Genetic population structure of the paper waspPolistes olivaceus(Hymenoptera: Vespidae) in Bangladesh

AbstractDispersal triggers gene flow, which in turn strongly affects the ensuing genetic population structure of a species. Using nuclear microsatellite loci and mitochondrial DNA (mtDNA), we estimated the genetic population structure of the waspPolistes olivaceusthroughout Bangladesh. The level of population differentiation using nuclear markers (FST) appeared to be much lower than that estimated using mtDNA haplotype sequences (ФST), even after correcting for effective population size differences between the two markers. These results suggest a philopatric tendency, in which gynes disperse less than males. We observed no isolation by distance among the study populations at either the nuclear or mtDNA level, suggesting nonequilibrium between gene flow and drift as a result of very frequent interpopulation movement. For the nuclear markers, an individual assignment test showed no genetically and geographically distinct groups. Instead, phylogenetic analyses as well as a minimum spanning network using mtDNA haplotypes consistently revealed two distinct lineages. The distribution of haplotypes indicated western populations with a single lineage and offered clear evidence for restricted gene flow across the Jamuna–Padma–Upper Meghna river system. Mismatch distributions exhibited a unimodal distribution, which along with a starlike haplotype network, suggested a population expansion in lineage I but not in lineage II. Overall, these results suggest that gene flow among populations ofP. olivaceuswas affected by both female philopatry and a major river system across Bangladesh.

Genetic analysis reveals demographic fragmentation of grizzly bears yielding vulnerably small populations

Ecosystem conservation requires the presence of native carnivores, yet in North America, the distributions of many larger carnivores have contracted. Large carnivores live at low densities and require large areas to thrive at the population level. Therefore, if human-dominated landscapes fragment remaining carnivore populations, small and demographically vulnerable populations may result. Grizzly bear range contraction in the conterminous USA has left four fragmented populations, three of which remain along the Canada-USA border. A tenet of grizzly bear conservation is that the viability of these populations requires demographic linkage (i.e. inter-population movement of both sexes) to Canadian bears. Using individual-based genetic analysis, our results suggest this demographic connection has been severed across their entire range in southern Canada by a highway and associated settlements, limiting female and reducing male movement. Two resulting populations are vulnerably small (< or =100 animals) and one of these is completely isolated. Our results suggest that these trans-border bear populations may be more threatened than previously thought and that conservation efforts must expand to include international connectivity management. They also demonstrate the ability of genetic analysis to detect gender-specific demographic population fragmentation in recently disturbed systems, a traditionally intractable yet increasingly important ecological measurement worldwide.

MODELING INTERPOPULATION DISPERSAL BY BANNER-TAILED KANGAROO RATS

Many metapopulation models assume rules of population connectivity that are implicitly based on what we know about within-population dispersal, but especially for vertebrates, few data exist to assess whether interpopulation dispersal is just within-population dispersal “scaled up.” We extended existing multi-stratum mark–release–recapture models to incorporate the robust design, allowing us to compare patterns of within- and between-population movement in the banner-tailed kangaroo rat (Dipodomys spectabilis). Movement was rare among eight populations separated by only a few hundred meters: seven years of twice-annual sampling captured >1200 individuals but only 26 interpopulation dispersers. We developed a program that implemented models with parameters for capture, survival, and interpopulation movement probability and that evaluated competing hypotheses in a model selection framework. We evaluated variants of the island, stepping-stone, and isolation-by-distance models of interpopulation movement, incorporating effects of age, season, and habitat (short or tall grass). For both sexes, QAICc values clearly favored isolation-by-distance models, or models combining the effects of isolation by distance and habitat. Models with probability of dispersal expressed as linear-logistic functions of distance and as negative exponentials of distance fit the data equally well. Interpopulation movement probabilities were similar among sexes (perhaps slightly biased toward females), greater for juveniles than adults (especially for females), and greater before than during the breeding season (especially for females). These patterns resemble those previously described for within-population dispersal in this species, which we interpret as indicating that the same processes initiate both within- and between-population dispersal.

Implications of Anthropogenic Landscape Change on Inter-Population Movements of the Desert Tortoise (Gopherus agassizii)

In the Sonoran Desert of North America, populations of the desert tortoise (Gopherus agassizii) occur in rocky foothills throughout southwestern Arizona and northwestern Mexico. Although tortoise populations appear to be isolated from each other by low desert valleys, individuals occasionally move long distances between populations. Increasingly, these movements are hindered by habitat fragmentation due to anthropogenic landscape changes. We used molecular techniques and radiotelemetry to examine movement patterns of desert tortoises in southern Arizona. We collected blood samples from 170 individuals in nine mountain ranges and analyzed variability in seven microsatellite loci to determine genetic differentiation among populations. Gene flow estimates between populations indicate that populations exchanged individuals historically at a rate greater than one migrant per generation, and positive correlation between genetic and geographic distance of population pairs suggests that the limiting factor for gene flow among populations is isolation by distance. Life history traits of the desert tortoise, a long-lived species with delayed sexual maturity, may severely constrain the ability of small populations to respond to disturbances that increase adult mortality. Historic gene flow estimates among populations suggests that recovery of declining populations may rely heavily on the immigration of new individuals from adjacent mountain ranges. Management strategies compatible with the evolutionary history of gene flow among disjunct populations will help ensure the long-term persistence of Sonoran desert tortoise populations.

Synchronisation Between Individuals and the Dynamics of Linked Populations

Recent research into the dynamics of linked populations, using lattices of difference equations, suggest that coupling by migration between populations often has no effect on the stability of the populations within the system. An underlying assumption of these models is that inter-population movement is segregated in time from other population processes. Here I present an individual-based model which does not require this restriction, and show that the behaviour of the model is largely unaffected by reducing the synchrony between migration events. This holds even if migration and reproduction events are completely randomly mixed. Another restriction of several previous models is that migration only occurs to nearest-neighbour sites. Again, an individual-based model is presented which allows individuals to move several times per generation and hence allow them to reproduce at a site far removed from their position at the start of the generation. This generalisation is shown to have little effect on the stability of local population. Local coupling does not appear to affect whether the equilibrium is locally stable or unstable, however, it does affect qualitative features of unstable dynamics.

Movements of a Polar Bear from Northern Alaska to Northern Greenland

Using satellite telemetry, we monitored the movements of an adult female polar bear ( Ursus maritimus ) as she traveled from the Alaskan Beaufort Sea coast to northern Greenland. She is the first polar bear known to depart the Beaufort Sea region for an extended period, and the first polar bear known to move between Alaska and Greenland. This bear traveled for four months across the polar basin and came within 2 degrees of the North Pole. During the first year following her capture, she traveled 5256 km. Evidence to suggest her use of maternity dens in northern Alaska and in northern Greenland demonstrates the potential for genetic exchange between two widely separate populations of polar bears. The long life spans of polar bears and the rarity of their long-range movements means the significance of interpopulation movement can be assessed after long-term monitoring of individuals. Key words: polar bear, Ursus maritimus , satellite telemetry, movements, Beaufort Sea, populations, Alaska, Greenland, polar basin

Genetic and Ecological Guidelines for Species Reintroduction Programs: Application to Great Lakes Fishes

Complete restoration of the Great Lakes is unlikely, due to naturalization of exotic species, habitat degradation and destruction, heavy fishing mortality, lack of native gene pools, and complicated political jurisdictions that rarely work toward a common vision. A more realistic goal is rehabilitation, a movement along the trajectory toward complete restoration. Proper rehabilitation employs an evolutionary-genetic perspective, which protects and works with the remaining genetic variation available in lake trout or other species of concern. A difficult question is how to define the units of genetic conservation; one answer is to determine Evolutionary Significant Units. To do this, population structure must be defined, and various conceptual models are here presented that determine genetic population structure as a function of geographic structure of the habitat. Several concepts from the developing field of conservation biology should also be incorporated into lake trout rehabilitation. These include metapopulations—groups of populations that experience some degree of regular or intermittent gene flow, and that serve as recolonization sources after local extinctions. Related to this is “source-sink dynamics,” a recognition that habitats typically are unequal in quality. Habitat fragmentation is also relevant because it can stop inter-population movement and disrupt metapopulation structure and source-sink dynamics. Finally, hatcheries should be used in rehabilitation only with great caution and skepticism. Hatcheries tend to address the symptoms, rather than the causes, of fish declines, and may mask the underlying problems by continually replacing declining fish. Improved habitat quality, rather than artificial production, is the key to rehabilitation of the Great Lakes for lake trout and other native species, and hatcheries should only be used as a stopgap measure to prevent further losses of genetic diversity.