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1.
Sci Adv ; 9(50): eadh4584, 2023 Dec 15.
Article in English | MEDLINE | ID: mdl-38100594

ABSTRACT

Antarctic krill, crucial to the Southern Ocean ecosystem and a vital fisheries resource, is endangered by climate change. Identifying drivers of krill biomass is therefore essential for determining catch limits and designating protection zones. We present a modeling approach to pinpointing effects of sea surface temperature, ice cover, chlorophyll levels, climate indices, and intraspecific competition. Our study reveals that larval recruitment is driven by both competition among age classes and chlorophyll levels. In addition, while milder ice and temperature in spring and summer favor reproduction and early larval survival, both larvae and juveniles strongly benefit from heavier ice and colder temperatures in winter. We conclude that omitting top-down control of resources by krill is only acceptable for retrospective or single-year prognostic models that use field chlorophyll data but that incorporating intraspecific competition is essential for longer-term forecasts. Our findings can guide future krill modeling strategies, reinforcing the sustainability of this keystone species.


Subject(s)
Ecosystem , Euphausiacea , Animals , Retrospective Studies , Ice Cover , Chlorophyll , Larva , Antarctic Regions
2.
Mov Ecol ; 11(1): 37, 2023 Jul 05.
Article in English | MEDLINE | ID: mdl-37408064

ABSTRACT

BACKGROUND: For many migratory species, inexperienced (naïve) individuals reach remote non-breeding areas independently using one or more inherited compass headings and, potentially, magnetic signposts to gauge where to switch between compass headings. Inherited magnetic-based migration has not yet been assessed as a population-level process, particularly across strong geomagnetic gradients or where long-term geomagnetic shifts (hereafter, secular variation) could create mismatches with magnetic headings. Therefore, it remains unclear whether inherited magnetic headings and signposts could potentially adapt to secular variation under natural selection. METHODS: To address these unknowns, we modelled migratory orientation programs using an evolutionary algorithm incorporating global geomagnetic data (1900-2023). Modelled population mixing incorporated both natal dispersal and trans-generational inheritance of magnetic headings and signposts, including intrinsic (stochastic) variability in inheritance. Using the model, we assessed robustness of trans-hemispheric migration of a migratory songbird whose Nearctic breeding grounds have undergone rapid secular variation (mean 34° clockwise drift in declination, 1900-2023), and which travels across strong geomagnetic gradients via Europe to Africa. RESULTS: Model-evolved magnetic-signposted migration was overall successful throughout the 124-year period, with 60-90% mean successful arrival across a broad range in plausible precision in compass headings and gauging signposts. Signposted migration reduced trans-Atlantic flight distances and was up to twice as successful compared with non-signposted migration. Magnetic headings shifted plastically in response to the secular variation (mean 16°-17° among orientation programs), whereas signpost latitudes were more constrained (3°-5° mean shifts). This plasticity required intrinsic variability in inheritance (model-evolved σ ≈ 2.6° standard error), preventing clockwise secular drift from causing unsustainable open-ocean flights. CONCLUSIONS: Our study supports the potential long-term viability of inherited magnetic migratory headings and signposts, and illustrates more generally how inherited migratory orientation programs can both mediate and constrain evolution of routes, in response to global environmental change.

3.
Proc Natl Acad Sci U S A ; 119(43): e2118156119, 2022 10 25.
Article in English | MEDLINE | ID: mdl-36256813

ABSTRACT

The twin crises of climate change and biodiversity loss define a strong need for functional diversity monitoring. While the availability of high-quality ecological monitoring data is increasing, the quantification of functional diversity so far requires the identification of species traits, for which data are harder to obtain. However, the traits that are relevant for the ecological function of a species also shape its performance in the environment and hence, should be reflected indirectly in its spatiotemporal distribution. Thus, it may be possible to reconstruct these traits from a sufficiently extensive monitoring dataset. Here, we use diffusion maps, a deterministic and de facto parameter-free analysis method, to reconstruct a proxy representation of the species' traits directly from monitoring data and use it to estimate functional diversity. We demonstrate this approach with both simulated data and real-world phytoplankton monitoring data from the Baltic Sea. We anticipate that wider application of this approach to existing data could greatly advance the analysis of changes in functional biodiversity.


Subject(s)
Biodiversity , Phytoplankton , Climate Change , Phenotype , Baltic States , Ecosystem
4.
Chaos ; 32(9): 093124, 2022 Sep.
Article in English | MEDLINE | ID: mdl-36182390

ABSTRACT

A class of nucleation and growth models of a stable phase is investigated for various different growth velocities. It is shown that for growth velocities v ≈ s ( t ) / t and v ≈ x / τ ( x ), where s ( t ) and τ are the mean domain size of the metastable phase (M-phase) and the mean nucleation time, respectively, the M-phase decays following a power law. Furthermore, snapshots at different time t that are taken to collect data for the distribution function c ( x , t ) of the domain size x of the M-phase are found to obey dynamic scaling. Using the idea of data-collapse, we show that each snapshot is a self-similar fractal. However, for v = const ., such as in the classical Kolmogorov-Johnson-Mehl-Avrami model, and for v ≈ 1 / t, the decays of the M-phase are exponential and they are not accompanied by dynamic scaling. We find a perfect agreement between numerical simulation and analytical results.

5.
Commun Biol ; 5(1): 1058, 2022 10 04.
Article in English | MEDLINE | ID: mdl-36195660

ABSTRACT

Migratory orientation of many animals is inheritable, enabling inexperienced (naïve) individuals to migrate independently using a geomagnetic or celestial compass. It remains unresolved how naïve migrants reliably reach remote destinations, sometimes correcting for orientation error or displacement. To assess naïve migratory performance (successful arrival), we simulate and assess proposed compass courses for diverse airborne migratory populations, accounting for spherical-geometry effects, compass precision, cue transfers (e.g., sun to star compass), and geomagnetic variability. We formulate how time-compensated sun-compass headings partially self-correct, according to how inner-clocks are updated. For the longest-distance migrations simulated, time-compensated sun-compass courses are most robust to error, and most closely resemble known routes. For shorter-distance nocturnal migrations, geomagnetic or star-compass courses are most robust, due to not requiring nightly cue-transfers. Our predictive study provides a basis for assessment of compass-based naïve migration and mechanisms of self-correction, and supports twilight sun-compass orientation being key to many long-distance inaugural migrations.


Subject(s)
Animal Migration , Orientation , Animals
6.
Math Biosci Eng ; 19(6): 5482-5508, 2022 03 28.
Article in English | MEDLINE | ID: mdl-35603365

ABSTRACT

Bacteria, in contrast to eukaryotic cells, contain two types of genes: chromosomal genes that are fixed to the cell, and plasmids, smaller loops of DNA capable of being passed from one cell to another. The sharing of plasmid genes between individual bacteria and between bacterial lineages has contributed vastly to bacterial evolution, allowing specialized traits to 'jump ship' between one lineage or species and the next. The benefits of this generosity from the point of view of both recipient cell and plasmid are generally understood: plasmids receive new hosts and ride out selective sweeps across the population, recipient cells gain new traits (such as antibiotic resistance). Explaining this behavior from the point of view of donor cells is substantially more difficult. Donor cells pay a fitness cost in order to share plasmids, and run the risk of sharing advantageous genes with their competition and rendering their own lineage redundant, while seemingly receiving no benefit in return. Using both compartment based models and agent based simulations we demonstrate that 'secretive' genes which restrict horizontal gene transfer are favored over a wide range of models and parameter values, even when sharing carries no direct cost. 'Generous' chromosomal genes which are more permissive of plasmid transfer are found to have neutral fitness at best, and are generally disfavored by selection. Our findings lead to a peculiar paradox: given the obvious benefits of keeping secrets, why do bacteria share information so freely?


Subject(s)
Bacteria , Gene Transfer, Horizontal , Bacteria/genetics , Drug Resistance, Microbial , Phenotype , Plasmids/genetics
7.
R Soc Open Sci ; 9(3): 220181, 2022 Mar.
Article in English | MEDLINE | ID: mdl-35345436

ABSTRACT

Clinicians prescribing antibiotics in a hospital context follow one of several possible 'treatment protocols'-heuristic rules designed to balance the immediate needs of patients against the long-term threat posed by the evolution of antibiotic resistance and multi-resistant bacteria. Several criteria have been proposed for assessing these protocols; unfortunately, these criteria frequently conflict with one another, each providing a different recommendation as to which treatment protocol is best. Here, we review and compare these optimization criteria. We are able to demonstrate that criteria focused primarily on slowing evolution of resistance are directly antagonistic to patient health both in the short and long term. We provide a new optimization criteria of our own, intended to more meaningfully balance the needs of the future and present. Asymptotic methods allow us to evaluate this criteria and provide insights not readily available through the numerical methods used previously in the literature. When cycling antibiotics, we find an antibiotic switching time which proves close to optimal across a wide range of modelling assumptions.

8.
Front Netw Physiol ; 2: 834057, 2022.
Article in English | MEDLINE | ID: mdl-36926111

ABSTRACT

A distinguishing feature of many ecological networks in the microbial realm is the diversity of substrates that could potentially serve as energy sources for microbial consumers. The microorganisms are themselves the agents of compound diversification via metabolite excretion or overflow metabolism. It has been suggested that the emerging richness of different substrates is an important condition for the immense biological diversity in microbial ecosystems. In this work, we study how complex cross-feeding networks (CFN) of microbial species may develop from a simple initial community given some elemental evolutionary mechanisms of resource-dependent speciation and extinctions using a network flow model. We report results of several numerical experiments and report an in-depth analysis of the evolutionary dynamics. We find that even in stable environments, the system is subject to persisting turnover, indicating an ongoing co-evolution. Further, we compare the impact of different parameters, such as the ratio of mineralization, as well as the metabolic versatility and variability on the evolving community structure. The results imply that high microbial and molecular diversity is an emergent property of evolution in cross-feeding networks, which affects transformation and accumulation of substrates in natural systems, such as soils and oceans, with potential relevance to biotechnological applications.

9.
PLoS Biol ; 19(12): e3001449, 2021 12.
Article in English | MEDLINE | ID: mdl-34932574

ABSTRACT

Plasmid transfer contributes significantly to bacterial evolution, but the forces selecting such generosity are poorly understood; this Formal Comment revisits a study which examined these forces both analytically and experimentally, making a correction to the algebra and reaching strikingly different results.


Subject(s)
Bacteria , Bacteria/genetics , Plasmids/genetics
11.
Sci Rep ; 11(1): 7483, 2021 04 05.
Article in English | MEDLINE | ID: mdl-33820912

ABSTRACT

In a warming climate, rising seawater temperatures and declining primary and secondary production will drastically affect growth and fitness of marine invertebrates in the northern Atlantic Ocean. To study the ecological performance of juvenile hydroids Hydractinia echinata we exposed them to current and predicted water temperatures which reflect the conditions in the inter- and subtidal in combination with changing food availability (high and low) in laboratory experiments. Here we show, that the interplay between temperature stress and diminished nutrition affected growth and vitality of juvenile hydroids more than either factor alone, while high food availability mitigated their stress responses. Our numerical growth model indicated that the growth of juvenile hydroids at temperatures beyond their optimum is a saturation function of energy availability. We demonstrated that the combined effects of environmental stressors should be taken into consideration when evaluating consequences of climate change. Interactive effects of ocean warming, decreasing resource availability and increasing organismal energy demand may have major impacts on biodiversity and ecosystem function.


Subject(s)
Hydrozoa/growth & development , Nutrients/analysis , Temperature , Animals , Models, Biological , Statistics as Topic , Water
12.
Ecol Lett ; 24(4): 847-861, 2021 Apr.
Article in English | MEDLINE | ID: mdl-33471443

ABSTRACT

Size and shape profoundly influence an organism's ecophysiological performance and evolutionary fitness, suggesting a link between morphology and diversity. However, not much is known about how body shape is related to taxonomic richness, especially in microbes. Here we analyse global datasets of unicellular marine phytoplankton, a major group of primary producers with an exceptional diversity of cell sizes and shapes and, additionally, heterotrophic protists. Using two measures of cell shape elongation, we quantify taxonomic diversity as a function of cell size and shape. We find that cells of intermediate volume have the greatest shape variation, from oblate to extremely elongated forms, while small and large cells are mostly compact (e.g. spherical or cubic). Taxonomic diversity is strongly related to cell elongation and cell volume, together explaining up to 92% of total variance. Taxonomic diversity decays exponentially with cell elongation and displays a log-normal dependence on cell volume, peaking for intermediate-volume cells with compact shapes. These previously unreported broad patterns in phytoplankton diversity reveal selective pressures and ecophysiological constraints on the geometry of phytoplankton cells which may improve our understanding of marine ecology and the evolutionary rules of life.


Subject(s)
Marine Biology , Phytoplankton , Biological Evolution , Cell Size
13.
Phys Rev E ; 102(4-1): 042301, 2020 Oct.
Article in English | MEDLINE | ID: mdl-33212633

ABSTRACT

The rise of the World Airline Network over the past century has led to sharp changes in our notions of "distance" and "closeness"-in terms of both trade and travel, but also (less desirably) with respect to the spread of disease. When novel pathogens are discovered, countries, cities, and hospitals are caught trying to predict how much time they have to prepare. In this paper, by considering the early stages of epidemic spread as a simple branching process, we derive the full probability distribution of arrival times. We are able to rederive a number of past arrival time results (in suitable limits) and demonstrate the robustness of our approach, both to parameter values far outside the traditionally considered regime and to errors in the parameter values used. The branching process approach provides some theoretical justification to the "effective distance" introduced by Brockmann and Helbing [Science 342, 1337 (2013)SCIEAS0036-807510.1126/science.1245200]; however, we also observe that when compared to real-world data, the predictive power of all methods in this class is significantly lower than has been previously reported.

14.
Philos Trans R Soc Lond B Biol Sci ; 375(1814): 20190455, 2020 12 21.
Article in English | MEDLINE | ID: mdl-33131442

ABSTRACT

Dispersal and foodweb dynamics have long been studied in separate models. However, over the past decades, it has become abundantly clear that there are intricate interactions between local dynamics and spatial patterns. Trophic meta-communities, i.e. meta-foodwebs, are very complex systems that exhibit complex and often counterintuitive dynamics. Over the past decade, a broad range of modelling approaches have been used to study these systems. In this paper, we review these approaches and the insights that they have revealed. We focus particularly on recent papers that study trophic interactions in spatially extensive settings and highlight the common themes that emerged in different models. There is overwhelming evidence that dispersal (and particularly intermediate levels of dispersal) benefits the maintenance of biodiversity in several different ways. Moreover, some insights have been gained into the effect of different habitat topologies, but these results also show that the exact relationships are much more complex than previously thought, highlighting the need for further research in this area. This article is part of the theme issue 'Integrative research perspectives on marine conservation'.


Subject(s)
Biodiversity , Food Chain , Models, Biological , Animals , Conservation of Natural Resources
15.
Chaos ; 30(9): 093123, 2020 Sep.
Article in English | MEDLINE | ID: mdl-33003939

ABSTRACT

COVID-19 is an emerging respiratory infectious disease caused by the coronavirus SARS-CoV-2. It was first reported on in early December 2019 in Wuhan, China and within three months spread as a pandemic around the whole globe. Here, we study macro-epidemiological patterns along the time course of the COVID-19 pandemic. We compute the distribution of confirmed COVID-19 cases and deaths for countries worldwide and for counties in the US and show that both distributions follow a truncated power-law over five orders of magnitude. We are able to explain the origin of this scaling behavior as a dual-scale process: the large-scale spread of the virus between countries and the small-scale accumulation of case numbers within each country. Assuming exponential growth on both scales, the critical exponent of the power-law is determined by the ratio of large-scale to small-scale growth rates. We confirm this theory in numerical simulations in a simple meta-population model, describing the epidemic spread in a network of interconnected countries. Our theory gives a mechanistic explanation why most COVID-19 cases occurred within a few epicenters, at least in the initial phase of the outbreak. By combining real world data, modeling, and numerical simulations, we make the case that the distribution of epidemic prevalence might follow universal rules.


Subject(s)
Coronavirus Infections/epidemiology , Models, Theoretical , Pneumonia, Viral/epidemiology , Betacoronavirus , COVID-19 , Humans , Pandemics , Population Dynamics , SARS-CoV-2
16.
Nature ; 577(7789): 226-230, 2020 01.
Article in English | MEDLINE | ID: mdl-31853064

ABSTRACT

Predator-prey cycles rank among the most fundamental concepts in ecology, are predicted by the simplest ecological models and enable, theoretically, the indefinite persistence of predator and prey1-4. However, it remains an open question for how long cyclic dynamics can be self-sustained in real communities. Field observations have been restricted to a few cycle periods5-8 and experimental studies indicate that oscillations may be short-lived without external stabilizing factors9-19. Here we performed microcosm experiments with a planktonic predator-prey system and repeatedly observed oscillatory time series of unprecedented length that persisted for up to around 50 cycles or approximately 300 predator generations. The dominant type of dynamics was characterized by regular, coherent oscillations with a nearly constant predator-prey phase difference. Despite constant experimental conditions, we also observed shorter episodes of irregular, non-coherent oscillations without any significant phase relationship. However, the predator-prey system showed a strong tendency to return to the dominant dynamical regime with a defined phase relationship. A mathematical model suggests that stochasticity is probably responsible for the reversible shift from coherent to non-coherent oscillations, a notion that was supported by experiments with external forcing by pulsed nutrient supply. Our findings empirically demonstrate the potential for infinite persistence of predator and prey populations in a cyclic dynamic regime that shows resilience in the presence of stochastic events.


Subject(s)
Chlorophyta/physiology , Food Chain , Models, Biological , Rotifera/physiology , Animals , Biota , Chlorella vulgaris/growth & development , Chlorella vulgaris/physiology , Chlorophyta/growth & development , Rotifera/growth & development
17.
Appl Environ Microbiol ; 85(24)2019 12 15.
Article in English | MEDLINE | ID: mdl-31604772

ABSTRACT

Growth energetics and metabolic efficiency contribute to the lifestyle and habitat imprint of microorganisms. Roseobacters constitute one of the most abundant and successful marine bacterioplankton groups. Here, we reflect on the energetics and metabolic efficiency of Phaeobacter inhibens DSM 17395, a versatile heterotrophic roseobacter. Fourteen different substrates (five sugars and nine amino acids) and their degradation pathways were assessed for energetic efficiencies based on catabolic ATP yields, calculated from net formed ATP and reducing equivalents. The latter were converted into ATP by employing the most divergent coupling ratios (i.e., ions per ATP) currently known for F1Fo ATP synthases in heterotrophic bacteria. The catabolic ATP yields of the pathways studied in P. inhibens differed ∼3-fold. The actual free energy costs for ATP synthesis were estimated at 81.6 kJ per mol ATP (3.3 ions per ATP) or 104.2 kJ per mol ATP (4.3 ions per ATP), yielding an average thermodynamic efficiency of ∼37.7% or ∼29.5%, respectively. Growth performance (rates, yields) and carbon assimilation efficiency were determined for P. inhibens growing in process-controlled bioreactors with 10 different single substrates (Glc, Man, N-acetylglucosamine [Nag], Phe, Trp, His, Lys, Thr, Val, or Leu) and with 2 defined substrate mixtures. The efficiencies of carbon assimilation into biomass ranged from ∼28% to 61%, with His/Trp and Thr/Leu yielding the lowest and highest levels. These efficiencies correlated with catabolic and ATP yields only to some extent. Substrate-specific metabolic demands and/or functions, as well as the compositions of the substrate mixtures, apparently affected the energetic costs of growth. These include energetic burdens associated with, e.g., slow growth, stress, and/or the production of tropodithietic acid.IMPORTANCE Heterotrophic members of the bacterioplankton serve the marine ecosystem by transforming organic matter, an activity that is governed by the bacterial growth efficiencies (BGEs) obtained under given environmental conditions. In marine ecology, the concept of BGE refers to the carbon assimilation efficiency within natural communities. The marine bacterium studied here, Phaeobacter inhibens DSM 17395, is a copiotrophic representative of the globally abundant Roseobacter group, and the 15 catabolic pathways investigated are widespread among these marine heterotrophs. Combining pathway-specific catabolic ATP yields with in-depth quantitative physiological data could (i) provide a new baseline for the study of growth energetics and efficiency in further Roseobacter group members and other copiotrophic marine bacteria in productive coastal ecosystems and (ii) contribute to a better understanding of the factors controlling BGE (including the additional energetic burden arising from widespread secondary-metabolite formation) based on laboratory studies with pure cultures.


Subject(s)
Amino Acids/metabolism , Heterotrophic Processes/physiology , Rhodobacteraceae/metabolism , Sugars/metabolism , Adenosine Triphosphate/metabolism , Biomass , Bioreactors , Carbohydrate Metabolism , Metabolic Networks and Pathways , Rhodobacteraceae/growth & development , Roseobacter/metabolism , Tropolone/analogs & derivatives
18.
Proc Biol Sci ; 286(1901): 20190036, 2019 04 24.
Article in English | MEDLINE | ID: mdl-31014215

ABSTRACT

The number of released individuals, which is a component of propagule pressure, is considered to be a major driver for the establishment success of non-native species. However, propagule pressure is often assumed to result from single or few release events, which does not necessarily apply to the frequent releases of invertebrates or other taxa through global transport. For instance, the high intensity of global shipping may result in frequent releases of large numbers of individuals, and the complexity of shipping dynamics impedes predictions of invasion dynamics. Here, we present a mathematical model for the spread of planktonic organisms by global shipping, using the history of movements by 33 566 ships among 1477 ports to simulate population dynamics for the comb jelly Mnemiopsis leidyi as a case study. The degree of propagule pressure at one site resulted from the coincident arrival of individuals from other sites with native or non-native populations. Key to sequential spread in European waters was a readily available source of propagules and a suitable recipient environment. These propagules were derived from previously introduced 'bridgehead' populations supplemented with those from native sources. Invasion success is therefore determined by the complex interaction of global shipping and local population dynamics. The general findings probably hold true for the spread of species in other complex systems, such as insects or plant seeds exchanged via commercial trade or transport.


Subject(s)
Ctenophora/physiology , Introduced Species , Zooplankton/physiology , Animals , Ecosystem , Models, Biological , Population Dynamics , Ships
19.
FEMS Microbiol Ecol ; 94(10)2018 10 01.
Article in English | MEDLINE | ID: mdl-30124819

ABSTRACT

Reduced nitrogen species are key nutrients for biological productivity in the oceans. Ammonium is often present in low and growth-limiting concentrations, albeit peaks occur during collapse of algal blooms or via input from deep sea upwelling and riverine inflow. Autotrophic phytoplankton exploit ammonium peaks by storing nitrogen intracellularly. In contrast, the strategy of heterotrophic bacterioplankton to acquire ammonium is less well understood. This study revealed the marine bacterium Phaeobacter inhibens DSM 17395, a Roseobacter group member, to have already depleted the external ammonium when only ∼⅓ of the ultimately attained biomass is formed. This was paralleled by a three-fold increase in cellular nitrogen levels and rapid buildup of various nitrogen-containing intracellular metabolites (and enzymes for their biosynthesis) and biopolymers (DNA, RNA and proteins). Moreover, nitrogen-rich cells secreted potential RTX proteins and the antibiotic tropodithietic acid, perhaps to competitively secure pulses of external ammonium and to protect themselves from predation. This complex response may ensure growing cells and their descendants exclusive provision with internal nitrogen stocks. This nutritional strategy appears prevalent also in other roseobacters from distant geographical provenances and could provide a new perspective on the distribution of reduced nitrogen in marine environments, i.e. temporary accumulation in bacterioplankton cells.


Subject(s)
Ammonium Compounds/metabolism , Nitrogen/metabolism , Plankton/metabolism , Roseobacter/metabolism , Seawater/microbiology , Ammonium Compounds/analysis , Anti-Bacterial Agents/metabolism , Bacterial Proteins/metabolism , Biomass , Heterotrophic Processes , Plankton/chemistry , Roseobacter/chemistry , Seawater/chemistry , Tropolone/analogs & derivatives , Tropolone/metabolism
20.
R Soc Open Sci ; 5(6): 180438, 2018 Jun.
Article in English | MEDLINE | ID: mdl-30110431

ABSTRACT

Regular, long-distance migrations of thousands of animal species have consequences for the ecosystems that they visit, modifying trophic interactions and transporting many non-pathogenic and pathogenic organisms. The spatial structure and dynamic properties of animal migrations and population flyways largely determine those trophic and transport effects, but are yet poorly studied. As a basis, we propose a periodic Markov model on the spatial migration network of breeding, stopover and wintering sites to formally describe the process of animal migration on the population level. From seasonally changing transition rates we derived stable, seasonal densities of animals at the network nodes. We parametrized the model with high-quality GPS and satellite telemetry tracks of white storks (Ciconia ciconia) and greater white-fronted geese (Anser a. albifrons). Topological and network flow properties of the two derived networks conform to migration properties like seasonally changing connectivity and shared, directed movement. Thus, the model realistically describes the migration movement of complete populations and can become an important tool to study the effects of climate and habitat change and pathogen spread on migratory animals. Furthermore, the property of periodically changing transition rates makes it a new type of complex model and we need to understand its dynamic properties.

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