The movement ecology of seagrasses

A movement ecology framework is applied to enhance our understanding of the causes, mechanisms and consequences of movement in seagrasses: marine, clonal, flowering plants. Four life-history stages of seagrasses can move: pollen, sexual propagules, vegetative fragments and the spread of individuals through clonal growth. Movement occurs on the water surface, in the water column, on or in the sediment, via animal vectors and through spreading clones. A capacity for long-distance dispersal and demographic connectivity over multiple timeframes is the novel feature of the movement ecology of seagrasses with significant evolutionary and ecological consequences. The space–time movement footprint of different life-history stages varies. For example, the distance moved by reproductive propagules and vegetative expansion via clonal growth is similar, but the timescales range exponentially, from hours to months or centuries to millennia, respectively. Consequently, environmental factors and key traits that interact to influence movement also operate on vastly different spatial and temporal scales. Six key future research areas have been identified.     

Dispersal movements of non-native and native terrestrial slugs in an urban environment

Animal movement varies from undirected dispersal to directed migration. Movement rates may have implications for conservation and resource management, as well as pest control, and they play a key role in invasion success. In slugs, long-distance dispersal is typically passive, whereas active movement is critical for local dispersal and determines access to resources such as food and shelter. Telemetry has recently been used to study individual slug movements in the wild, whereas movement in arena tests has explored mechanisms of interspecific competition and invasiveness in slugs. Studies that relate the performance of individual slugs in arena tests to their post-release behavior in nature are lacking. We measured individual short-term movement speed of commonly occurring native and non-native slugs of the genera Arion and Limax in arena tests and tracked their post-release dispersal movements in a garden by PIT telemetry. We demonstrate clear differences in movement behavior among the species, but non-native slugs did not display higher movement rates than their native congeners. In the arena test, slugs of the genus Limax displayed a higher short-term speed than slugs of the genus Arion, whereas in the field, individuals of Limax maximus showed lower dispersal rates compared to the other slug species. Moreover, there was a positive correlation between short-term speed in the arena test and movement in the field among individuals of L. cinereoniger, indicating the possible existence of behavioral syndromes in slugs, which may link movement ecology, animal personality, and the invasion ecology of pest species.     

Animal movements in fire‐prone landscapes

Movement is a trait of fundamental importance in ecosystems subject to frequent disturbances, such as fire‐prone ecosystems. Despite this, the role of movement in facilitating responses to fire has received little attention. Herein, we consider how animal movement interacts with fire history to shape species distributions. We consider how fire affects movement between habitat patches of differing fire histories that occur across a range of spatial and temporal scales, from daily foraging bouts to infrequent dispersal events, and annual migrations. We review animal movements in response to the immediate and abrupt impacts of fire, and the longer‐term successional changes that fires set in train. We discuss how the novel threats of altered fire regimes, landscape fragmentation, and invasive species result in suboptimal movements that drive populations downwards. We then outline the types of data needed to study animal movements in relation to fire and novel threats, to hasten the integration of movement ecology and fire ecology. We conclude by outlining a research agenda for the integration of movement ecology and fire ecology by identifying key research questions that emerge from our synthesis of animal movements in fire‐prone ecosystems.     

Movement ecology of Afrotropical birds: Functional traits provide complementary insights to species identity

Effects of anthropogenic activities on habitats and species communities and populations are complex and vary across species depending on their ecological traits. Movement ecology may provide important insights into species’ responses to habitat structures and quality. We investigated how movement behavior across a human‐modified landscape depends on species identity and species traits, with particular focus on habitat specialization, feeding guilds, and dispersal behavior. We tracked 34 individuals of nine Afrotropical bird species during three years in an anthropogenic riparian landscape of East Africa. We investigated whether species’ functional traits predicted their habitat use and movement behavior better than species’ identities. Our results indicate that habitat specialists mainly occur in dense riparian thickets, while habitat generalists do occur in agricultural land. Home‐ranges of omnivorous habitat generalists are larger than of frugivorous and insectivorous generalists and omnivorous and insectivorous specialists. Movement speed was highest in settlement areas for all species, with activity peaks during morning and afternoon for habitat specialists. Our results reveal that functional traits and species identity provide complementary insights into responses of organisms to habitat structures and habitat quality.     

Finding habitat patches and directional connectivity

For animal species inhabiting patchy environments, the search behavior of individuals and the distance from which they can detect suitable habitat (perceptual range) are key determinants of the functional connectivity of landscapes. We examined the movement behavior and perceptual range of adult cactus bugs ( Chelinidea vittiger ), which are habitat specialists that feed and reproduce on Opuntia cactus. Movement pathways of walking individuals released into unsuitable matrix habitat (30–3000 m from Opuntia ) were highly directional. These results supported predictions of optimal search behavior from published simulation models. A release experiment within natural patch networks indicated that the perceptual range of C. vittiger depended on size of the target patch, matrix structure, and direction of the target patch relative to prevailing winds. A strong effect of wind direction on orientation behavior (and presumed perceptual range) was evident in a release experiment using 'artificial' patches of potted Opuntia . In these two experiments, individuals released 50–100 cm from Opuntia patches were more likely to orientate toward patches located upwind than to those located crosswind or downwind. A reexamination of the pathways of individuals walking in the matrix also revealed a strong bias for movement upwind. Such upwind movement by individuals, both within and outside of patch networks, suggests that C. vittiger uses olfaction to navigate and it complicates our ability to interpret search behavior and to estimate perceptual range. Current techniques for assessing perceptual range have limitations for olfactory-based species. Furthermore, we need to broaden our view of perceptual range and of patch and landscape connectivity. Perceptual range may be anisotropic and connectivity may be directional. An organism-based approach to spatial ecology requires that we consider the dominant senses used by species when navigating around patchy landscapes.     

Going through the motions: incorporating movement analyses into disease research

Though epidemiology dates back to the 1700s, most mathematical representations of epidemics still use transmission rates averaged at the population scale, especially for wildlife diseases. In simplifying the contact process, we ignore the heterogeneities in host movements that complicate the real world, and overlook their impact on spatiotemporal patterns of disease burden. Movement ecology offers a set of tools that help unpack the transmission process, letting researchers more accurately model how animals within a population interact and spread pathogens. Analytical techniques from this growing field can also help expose the reverse process: how infection impacts movement behaviours, and therefore other ecological processes like feeding, reproduction, and dispersal. Here, we synthesise the contributions of movement ecology in disease research, with a particular focus on studies that have successfully used movement‐based methods to quantify individual heterogeneity in exposure and transmission risk. Throughout, we highlight the rapid growth of both disease and movement ecology and comment on promising but unexplored avenues for research at their overlap. Ultimately, we suggest, including movement empowers ecologists to pose new questions, expanding our understanding of host–pathogen dynamics and improving our predictive capacity for wildlife and even human diseases.     

Commensal ecology, urban landscapes, and their influence on the genetic characteristics of city-dwelling Norway rats (Rattus norvegicus)

Movement of individuals promotes colonization of new areas, gene flow among local populations, and has implications for the spread of infectious agents and the control of pest species. Wild Norway rats ( Rattus norvegicus ) are common in highly urbanized areas but surprisingly little is known of their population structure. We sampled individuals from 11 locations within Baltimore, Maryland, to characterize the genetic structure and extent of gene flow between areas within the city. Clustering methods and a neighbour-joining tree based on pairwise genetic distances supported an east–west division in the inner city, and a third cluster comprised of historically more recent sites. Most individuals (~95%) were assigned to their area of capture, indicating strong site fidelity. Moreover, the axial dispersal distance of rats (62 m) fell within typical alley length. Several rats were assigned to areas 2–11.5 km away, indicating some, albeit infrequent, long-distance movement within the city. Although individual movement appears to be limited (30–150 m), locations up to 1.7 km are comprised of relatives. Moderate F _ST, differentiation between identified clusters, and high allelic diversity indicate that regular gene flow, either via recruitment or migration, has prevented isolation. Therefore, ecology of commensal rodents in urban areas and life-history characteristics of Norway rats likely counteract many expected effects of isolation or founder events. An understanding of levels of connectivity of rat populations inhabiting urban areas provides information about the spatial scale at which populations of rats may spread disease, invade new areas, or be eradicated from an existing area without reinvasion.     

State-space models of individual animal movement

Detailed observation of the movement of individual animals offers the potential to understand spatial population processes as the ultimate consequence of individual behaviour, physiological constraints and fine-scale environmental influences. However, movement data from individuals are intrinsically stochastic and often subject to severe observation error. Linking such complex data to dynamical models of movement is a major challenge for animal ecology. Here, we review a statistical approach, state-space modelling, which involves changing how we analyse movement data and draw inferences about the behaviours that shape it. The statistical robustness and predictive ability of state-space models make them the most promising avenue towards a new type of movement ecology that fuses insights from the study of animal behaviour, biogeography and spatial population dynamics.     

Movement ecology and sex are linked to barn owl microbial community composition

The behavioural ecology of host species is likely to affect their microbial communities, because host sex, diet, physiology, and movement behaviour could all potentially influence their microbiota. We studied a wild population of barn owls (Tyto alba) and collected data on their microbiota, movement, diet, size, coloration, and reproduction. The composition of bacterial species differed by the sex of the host and female owls had more diverse bacterial communities than their male counterparts. The abundance of two families of bacteria, Actinomycetaceae and Lactobacillaceae, also varied between the sexes, potentially as a result of sex differences in hormones and immunological function, as has previously been found with Lactobacillaceae in the microbiota of mice. Male and female owls did not differ in the prey they brought to the nest, which suggests that dietary differences are unlikely to underlie the differences in their microbiota. The movement behaviour of the owls was associated with the host microbiota in both males and females because owls that moved further from their nest each day had more diverse bacterial communities than owls that stayed closer to their nests. This novel result suggests that the movement ecology of hosts can impact their microbiota, potentially on the basis of their differential encounters with new bacterial species as the hosts move and forage across the landscape. Overall, we found that many aspects of the microbial community are correlated with the behavioural ecology of the host and that data on the microbiota can aid in generating new hypotheses about host behaviour.     

Temperature- and size-dependent growth and movement of the North Pacific giant octopus (Enteroctopus dofleini) in the Bering Sea

Octopus growth and movement occurs during all life stages and have implications for survival, food web dynamics and reproduction. From 2009 to 2011, 1714 North Pacific giant octopus (Enteroctopus dofleini) were tagged and recaptured in the eastern Bering Sea with visible implant elastomers to better understand the ecology of this data-poor species. Over this period, 246 of the tagged individuals were recaptured. In autumn, when temperatures were warmest, E. dofleini had higher growth rates and moved more than in the colder winter months. For both short- and long-term recaptures, small octopus grew faster than large octopus. Movement of octopus over short- and long-term periods was low for very small and very large octopus and high for median-sized octopus, which is likely to be a function of maturity status and reproductive activities. Approximately 80% of recaptures moved less than 2?km from the initial tagging location for time periods up to a year, suggesting long-term site fidelity. As temperature and size may be tightly linked to growth and movement rates for E. dofleini in the eastern Bering Sea, predicted climate warming will likely alter ecological processes for the species and impact their distribution.     

Modeling Taxa-Abundance Distributions in Microbial Communities using Environmental Sequence Data

We show that inferring the taxa-abundance distribution of a microbial community from small environmental samples alone is difficult. The difficulty stems from the disparity in scale between the number of genetic sequences that can be characterized and the number of individuals in communities that microbial ecologists aspire to describe. One solution is to calibrate and validate a mathematical model of microbial community assembly using the small samples and use the model to extrapolate to the taxa-abundance distribution for the population that is deemed to constitute a community. We demonstrate this approach by using a simple neutral community assembly model in which random immigrations, births, and deaths determine the relative abundance of taxa in a community. In doing so, we further develop a neutral theory to produce a taxa-abundance distribution for large communities that are typical of microbial communities. In addition, we highlight that the sampling uncertainties conspire to make the immigration rate calibrated on the basis of small samples very much higher than the true immigration rate. This scale dependence of model parameters is not unique to neutral theories; it is a generic problem in ecology that is particularly acute in microbial ecology. We argue that to overcome this, so that microbial ecologists can characterize large microbial communities from small samples, mathematical models that encapsulate sampling effects are required.     

Aggregative group behavior in insect parasitic nematode dispersal

Movement behavior of foraging animals is critical to the determination of their spatial ecology and success in exploiting resources. Individuals sometimes gain advantages by foraging in groups to increase their efficiency in garnering these resources. Group movement behavior has been studied in various vertebrates. In this study we explored the propensity for innate group movement behavior among insect parasitic nematodes. Given that entomopathogenic nematodes benefit from group attack and infection, we hypothesised that the populations would tend to move in aggregate in the absence of extrinsic cues. Movement patterns of entomopathogenic nematodes in sand were investigated when nematodes were applied to a specific locus or when the nematodes emerged naturally from infected insect hosts; six nematode species in two genera were tested (Heterorhabditis bacteriophora, Heterorhabditis indica, Steinernema carpocapsae, Steinernema feltiae, Steinernema glaseri and Steinernema riobrave). Nematodes were applied in aqueous suspension via filter paper discs or in infected insect host cadavers (to mimic emergence in nature). We discovered that nematode dispersal resulted in an aggregated pattern rather than a random or uniform distribution; the only exception was S. glaseri when emerging directly from infected hosts. The group movement may have been continuous from the point of origin, or it may have been triggered by a propensity to aggregate after a short period of random movement. To our knowledge, this is the first report of group movement behavior in parasitic nematodes in the absence of external stimuli (e.g., without an insect or other apparent biotic or abiotic cue). These findings have implications for nematode spatial distribution and suggest that group behavior is involved in nematode foraging.     

Gut microbial ecology of the Critically Endangered Fijian crested iguana (Brachylophus vitiensis): Effects of captivity status and host reintroduction on endogenous microbiomes

Animals often exhibit distinct microbial communities when maintained in captivity as compared to when in the wild. Such differentiation may be significant in headstart and reintroduction programs where individuals spend some time in captivity before release into native habitats. Using 16S rRNA gene sequencing, we (i) assessed differences in gut microbial communities between captive and wild Fijian crested iguanas (Brachylophus vitiensis) and (ii) resampled gut microbiota in captive iguanas released onto a native island to monitor microbiome restructuring in the wild. We used both cloacal swabs and fecal samples to further increase our understanding of gut microbial ecology in this IUCN Critically Endangered species. We found significant differentiation in gut microbial community composition and structure between captive and wild iguanas in both sampling schemes. Approximately two months postrelease, microbial communities in cloacal samples from formerly captive iguanas closely resembled wild counterparts. Interestingly, microbial communities in fecal samples from these individuals remained significantly distinct from wild conspecifics. Our results indicate that captive upbringings can lead to differences in microbial assemblages in headstart iguanas as compared to wild individuals even after host reintroduction into native conditions. This investigation highlights the necessity of continuous monitoring of reintroduced animals in the wild to ensure successful acclimatization and release.     

Understanding movement data and movement processes: current and emerging directions

Animal movement has been the focus on much theoretical and empirical work in ecology over the last 25 years. By studying the causes and consequences of individual movement, ecologists have gained greater insight into the behavior of individuals and the spatial dynamics of populations at increasingly higher levels of organization. In particular, ecologists have focused on the interaction between individuals and their environment in an effort to understand future impacts from habitat loss and climate change. Tools to examine this interaction have included: fractal analysis, first passage time, Lévy flights, multi‐behavioral analysis, hidden markov models, and state‐space models. Concurrent with the development of movement models has been an increase in the sophistication and availability of hierarchical bayesian models. In this review we bring these two threads together by using hierarchical structures as a framework for reviewing individual models. We synthesize emerging themes in movement ecology, and propose a new hierarchical model for animal movement that builds on these emerging themes. This model moves away from traditional random walks, and instead focuses inference on how moving animals with complex behavior interact with their landscape and make choices about its suitability.     

Long-Distance and Frequent Movements of the Flying-Fox Pteropus poliocephalus: Implications for Management

Flying-foxes (Pteropodidae) are large bats capable of long-distance flight. Many species are threatened; some are considered pests. Effective conservation and management of flying-foxes are constrained by lack of knowledge of their ecology, especially of movement patterns over large spatial scales. Using satellite telemetry, we quantified long-distance movements of the grey-headed flying-fox Pteropus poliocephalus among roost sites in eastern Australia. Fourteen adult males were tracked for 2-40 weeks (mean 25 weeks). Collectively, these individuals utilised 77 roost sites in an area spanning 1,075 km by 128 km. Movement patterns varied greatly between individuals, with some travelling long distances. Five individuals travelled cumulative distances >1,000 km over the study period. Five individuals showed net displacements >300 km during one month, including one movement of 500 km within 48 hours. Seasonal movements were consistent with facultative latitudinal migration in part of the population. Flying-foxes shifted roost sites frequently: 64% of roost visits lasted <5 consecutive days, although some individuals remained at one roost for several months. Modal 2-day distances between consecutive roosts were 21-50 km (mean 45 km, range 3-166 km). Of 13 individuals tracked for >12 weeks, 10 moved >100 km in one or more weeks. Median cumulative displacement distances over 1, 10 and 30 weeks were 0 km, 260 km and 821 km, respectively. On average, over increasing time-periods, one additional roost site was visited for each additional 100 km travelled. These findings explain why culling and relocation attempts have had limited success in resolving human-bat conflicts in Australia. Flying-foxes are highly mobile between camps and regularly travel long distances. Consequently, local control actions are likely to have only temporary effects on local flying-fox populations. Developing alternative methods to manage these conflicts remains an important challenge that should be informed by a better understanding of the species' movement patterns.     

Restricted movement by mottled sculpin (pisces: cottidae) in a southern Appalachian stream

1. We used direct observation and mark‐recapture techniques to quantify movements by mottled sculpins (Cottus bairdi) in a 1 km segment of Shope Fork in western North Carolina. Our objectives were to: (i) quantify the overall rate of sculpin movement, (ii) assess variation in movement among years, individuals, and sculpin size classes, (iii) relate movement to variation in stream flow and population size structure, and (iv) quantify relationships between movement and individual growth rates. 2. Movements were very restricted: median and mean movement distances for all sculpin size classes over a 45 day period were 1.3 and 4.4 m respectively. Nevertheless, there was a high degree of intrapopulation and temporal variation in sculpin movement. Movement of juveniles increased with discharge and with the density of large adults. Movement by small and large adults was not influenced by stream flow, but large adults where more mobile when their own density was high. Finally, there were differences in the growth rates of mobile and sedentary sculpins. Mobile juveniles grew faster than sedentary individuals under conditions of low flow and high density of large adults, whereas adults exhibited the opposite pattern. 3. Our results support the hypothesis that juvenile movement and growth is influenced by both intraspecific interactions with adults and stream flow. In contrast, adult movement appears to be influenced by competitive interactions among residents for suitable space. The relationship between movement and growth may provide a negative feedback mechanism regulating mottled sculpin populations in this system.     

Microbial diversity-productivity relationships in aquatic ecosystems

Thanks to recent advances in molecular biology, one's knowledge of microbial co-occurrence patterns, microbial biogeography and microbial biodiversity is expanding rapidly. This MiniReview explores microbial diversity-productivity relationships in the light of what is known from the general ecology literature. Analyses of microbial diversity-productivity relationships from 70 natural, experimental, and engineered aquatic ecosystems reveal patterns that are strikingly similar to those that have long been documented for communities of macroorganisms. Microbial ecology and the general science of ecology are thus continuing to converge.     

Paradox lost: variable colour-pattern geometry is associated with differences in movement in aposematic frogs

Aposematic signal variation is a paradox: predators are better at learning and retaining the association between conspicuousness and unprofitability when signal variation is low. Movement patterns and variable colour patterns are linked in non-aposematic species: striped patterns generate illusions of altered speed and direction when moving linearly, affecting predators'' tracking ability; blotched patterns benefit instead from unpredictable pauses and random movement. We tested whether the extensive colour-pattern variation in an aposematic frog is linked to movement, and found that individuals moving directionally and faster have more elongated patterns than individuals moving randomly and slowly. This may help explain the paradox of polymorphic aposematism: variable warning signals may reduce protection, but predator defence might still be effective if specific behaviours are tuned to specific signals. The interacting effects of behavioural and morphological traits may be a key to the evolution of warning signals.     

Habitat utilization by an invasive herbivorous fish (<Emphasis Type="Italic">Siganus rivulatus</Emphasis>) in its native and invaded range

Movement is essential for understanding the distribution and abundance of animals. While it has been suggested that invasion success can be facilitated by species’ ability to adapt to novel environments, direct comparisons of movement patterns between native and invaded ranges of animals in their natural habitat are rare. The rivulated rabbitfish Siganus rivulatus was introduced from the Red Sea into the Mediterranean, where it is now found in extremely high abundances, and has overgrazed the coastal marine ecosystem in many locations. Through a continuous acoustic tracking system, we found that the movement of S. rivulatus individuals at a Mediterranean site differed substantially from those at a Red Sea site, with individuals in the Mediterranean having larger overall home ranges and lower site fidelity. However, no variation between sites was found in daily home range sizes. Results show that at the Mediterranean site S. rivulatus individuals have a larger spatial footprint, which may contribute to their impact and ability to expand their distribution. This study demonstrates a potential shift in individual movement of a marine invasive species between its native and invaded range, and highlights the role of movement in understanding biological invasions.