Movement behavior of swordfish provisions connectivity between the temperate and tropical southwest Pacific Ocean.

Swordfish (Xiphias gladius) are a widely distributed (45°N-45°S) large pelagic fish targeted by fisheries worldwide. Swordfish that occur at high latitudes tend to disproportionately be large adults, so their movements have implications for population dynamics and fisheries management. In the southwest Pacific, little is known about this subset of the stock and existing evidence suggests limited movement from the subtropics into cooler high latitude waters. Here, we capitalize on the recent emergence of a recreational swordfish fishery off temperate southeast Australia to characterize movements of swordfish caught in the fishery with pop-up satellite archival transmitting tags. Data were recovered from tags deployed for 56-250 days on 11 swordfish (50-350 kg) tagged between 38 and 43°S in the western Tasman Sea. Five swordfish entered the Coral Sea (< 30°S), with four reaching north to 11-24°S, up to 3275 km away from location of capture. Behavior modelling suggests these four individuals rapidly transited north until encountering 23-27 °C water, at which point they lingered in the area for several months, consistent with spawning-related partial migration. One migrating swordfish still carrying a tag after the spawning season returned to ~ 120 km of its release location, suggesting site fidelity. Movements toward the central south Pacific were confined to two individuals crossing 165°E. Swordfish predominantly underwent normal diel vertical migration, descending into the mesopelagic zone at dawn (median daytime depth 494.9 m, 95% CI 460.4-529.5 m). Light attenuation predicted daytime depth, with swordfish rising by up to 195 m in turbid water. At night, swordfish were deeper during the full moon, median night-time depth 45.8 m (37.8-55.5) m versus 18.0 m (14.9-21.8) m at new moon. Modelling fine-scale (10 min-1) swordfish depth revealed dynamic effects of moon phase varying predictably across time of night with implications for fisheries interactions. Studying highly migratory fishes near distribution limits allows characterization of the full range of movement phenotypes within a population, a key consideration for important fish stocks in changing oceans.

Assessing the multiple benefits of partially protected marine protected areas in Australia: A systematic review protocol.

BACKGROUND: There is global pressure to protect more of the world's oceans, primarily to protect biodiversity, and to fulfill the "30 by 30" goal set by the International Union for the Conservation of Nature (IUCN) that has recently been ratified under the Kunming-Montreal Global Biodiversity Framework at the fifteenth Conference of Parties (COP-15). Fully protected marine protected areas (MPAs) provide the highest level of protection for biodiversity from destructive or extractive practices and may limit access to the area itself. Fully protected MPAs (also commonly referred to as 'no-take MPAs') ban all fishing activities, thereby removing the realisation of direct economic and social benefits from resource extraction within these areas. However, fully protected MPAs can still act as source of productivity to surrounding areas, while also providing an important scientific reference role for off-reserve management thereby providing indirect economic and social outcomes, as well as biodiversity benefits. Sustainable marine resource management strives to achieve 'triple-bottom line' benefits, where economic, social, and biodiversity benefits are maximised in managed areas of the ocean. Implementing 'partially protected' areas (PPAs) in areas of high biodiversity value (i.e., inshore, productive areas of the ocean) that allow for some extractive activities, may allow us to supplement fully MPAs to meet IUCN conservation goals, while maximising social and economic benefits. However, our current understanding lacks explicit quantitative assessments of whether and how PPAs can benefit (or otherwise) biodiversity, while also providing economic and social benefits. This study provides a method to systematically review the scientific and legislative literature to understand how PPAs may contribute to conserving biodiversity while also providing social and economic benefits to Australia. METHODS AND EXPECTED OUTPUTS: The implementation of partially protected areas (PPAs) requires careful consideration of many potentially competing factors, and an understanding of the types of partial protection already in place in a region. We have developed a systematic literature review protocol focussing on the primary research question: "What is the current state of partially protected area (PPA) implementation across Australian marine areas?". The aim of the review is to provide marine resource managers with a comprehensive overview of PPAs in Australia, including associated goals and stated management strategies to achieve these goals, and a methodological approach that may be utilised globally. The review protocol was designed by the research team for a Fisheries Resource and Development Corporation (FRDC) strategic research grant and will seek input from a project steering committee for the project on aggregation of the initial results. The steering committee is made up of stakeholders from a wide range of backgrounds and interests, covering marine conservation, fisheries management, Indigenous values, and academic research in Australia. Multiple academic databases, alongside Australian Federal, State, and Territory legislation and related policies will be reviewed using Boolean keyword search strings for both academic databases and relevant grey literature. Results from eligible documents will be compiled and insights from the review collated to provide information on the status of PPA implementation in Australia.

Lobsters with pre-existing damage to their mechanosensory statocyst organs do not incur further damage from exposure to seismic air gun signals.

Staotcysts, the mechanosensory organs common to many marine invertebrates, have shown sensitivity to aquatic noise. Previously, rock lobsters (Jasus edwardsii) from a remote site with little exposure to anthropogenic noise incurred persistent damage to the statocyst and righting reflex following exposure to seismic air gun signals. Here, J. edwardsii collected from a site subject to high levels of anthropogenic noise were exposed to an equivalent seismic air gun signal regime as the previous study of noise-naïve lobsters. Following exposure, both control and exposed treatments were found to have damage to the statocyst equivalent to that of noise-naïve lobsters following seismic exposure, which led to the conclusion that the damage was pre-existing and not exacerbated by seismic exposure. The source of the damage in the lobsters in this study could not be ascertained, but the soundscape comparisons of the collection sites showed that the noisy site had a 5-10 dB greater level of noise, equivalent to a 3-10 times greater intensity, in the 10-700 Hz range than was found at the remote collection site. In addition to the lack of further damage following seismic exposure, no disruption to the righting reflex was observed. Indeed, compared to the noise naïve lobsters, the lobsters here demonstrated an ability to cope with or adapt to the mechanosensory damage, indicating a need for better understanding of the ecological impacts of the damage caused by low frequency noise on marine organisms. More broadly, this study raises historical exposure to noise as a previously unrecognised but vitally important consideration for studies of aquatic noise.

Home range, site fidelity and synchronous migrations of three co-occurring, morphologically distinct estuarine fish species.

Determining the movement behaviours of animals is essential for understanding population dynamics. This is fundamental for developing effective spatial management strategies and in assessing the response of species to anthropogenic disturbance. This study uses a Bayesian state-space model applied to acoustic transmitter data to describe the temporal and spatial movement patterns of three estuarine fish species commonly targeted by recreational anglers in southeast Australia: Black Bream (Acanthopagrus butcheri), Sand Flathead (Platycephalus bassensis), and Brown Trout (Salmo trutta). Despite morphological differences between the three species, several common traits were observed in their movement and behaviour. Of the 50 individuals across all three species that were tracked, the vast majority remained within the estuary where they were tagged for the duration of the study. While the home ranges of the three species differed in size, all individuals remained resident around the mid-estuary where the majority of fish were tagged were released. Each of the species also displayed seasonal migrations, presumably linked to spawning. The timing of the beginning of these migrations was well synchronised both within and among species, starting in late spring/early summer. This suggests that environmental factors such as water temperature and day length may play an important role in cueing spawning behaviour for each of the species. These migratory behaviours suggest adverse changes to estuarine conditions such as reduced river flows may have potential consequences for spawning success for some species and hence implications for fisheries management.

Seismic air guns damage rock lobster mechanosensory organs and impair righting reflex.

The effects of anthropogenic aquatic noise on marine invertebrates are poorly understood. We investigated the impact of seismic surveys on the righting reflex and statocyst morphology of the palinurid rock lobster, Jasus edwardsii, using field-based exposure to air gun signals. Following exposure equivalent to a full-scale commercial assay passing within 100-500 m, lobsters showed impaired righting and significant damage to the sensory hairs of the statocyst. Reflex impairment and statocyst damage persisted over the course of the experiments-up to 365 days post-exposure and did not improved following moulting. These results indicate that exposure to air gun signals caused morphological damage to the statocyst of rock lobsters, which can in turn impair complex reflexes. This damage and impairment adds further evidence that anthropogenic aquatic noise has the potential to harm invertebrates, necessitating a better understanding of possible ecological and economic impacts.

Exposure to seismic air gun signals causes physiological harm and alters behavior in the scallop Pecten fumatus.

Seismic surveys map the seabed using intense, low-frequency sound signals that penetrate kilometers into the Earth's crust. Little is known regarding how invertebrates, including economically and ecologically important bivalves, are affected by exposure to seismic signals. In a series of field-based experiments, we investigate the impact of exposure to seismic surveys on scallops, using measurements of physiological and behavioral parameters to determine whether exposure may cause mass mortality or result in other sublethal effects. Exposure to seismic signals was found to significantly increase mortality, particularly over a chronic (months postexposure) time scale, though not beyond naturally occurring rates of mortality. Exposure did not elicit energetically expensive behaviors, but scallops showed significant changes in behavioral patterns during exposure, through a reduction in classic behaviors and demonstration of a nonclassic "flinch" response to air gun signals. Furthermore, scallops showed persistent alterations in recessing reflex behavior following exposure, with the rate of recessing increasing with repeated exposure. Hemolymph (blood analog) physiology showed a compromised capacity for homeostasis and potential immunodeficiency, as a range of hemolymph biochemistry parameters were altered and the density of circulating hemocytes (blood cell analog) was significantly reduced, with effects observed over acute (hours to days) and chronic (months) scales. The size of the air gun had no effect, but repeated exposure intensified responses. We postulate that the observed impacts resulted from high seabed ground accelerations driven by the air gun signal. Given the scope of physiological disruption, we conclude that seismic exposure can harm scallops.

Marine foods sourced from farther as their use of global ocean primary production increases.

The growing human population must be fed, but historic land-based systems struggle to meet expanding demand. Marine production supports some of the world's poorest people but increasingly provides for the needs of the affluent, either directly by fishing or via fodder-based feeds for marine and terrestrial farming. Here we show the expanding footprint of humans to utilize global ocean productivity to feed themselves. Our results illustrate how incrementally each year, marine foods are sourced farther from where they are consumed and moreover, require an increasing proportion of the ocean's primary productivity that underpins all marine life. Though mariculture supports increased consumption of seafood, it continues to require feeds based on fully exploited wild stocks. Here we examine the ocean's ability to meet our future demands to 2100 and find that even with mariculture supplementing near-static wild catches our growing needs are unlikely to be met without significant changes.

Age-dependent speciation can explain the shape of empirical phylogenies.

Tens of thousands of phylogenetic trees, describing the evolutionary relationships between hundreds of thousands of taxa, are readily obtainable from various databases. From such trees, inferences can be made about the underlying macroevolutionary processes, yet remarkably these processes are still poorly understood. Simple and widely used evolutionary null models are problematic: Empirical trees show very different imbalance between the sizes of the daughter clades of ancestral taxa compared to what models predict. Obtaining a simple evolutionary model that is both biologically plausible and produces the imbalance seen in empirical trees is a challenging problem, to which none of the existing models provide a satisfying answer. Here we propose a simple, biologically plausible macroevolutionary model in which the rate of speciation decreases with species age, whereas extinction rates can vary quite generally. We show that this model provides a remarkable fit to the thousands of trees stored in the online database TreeBase. The biological motivation for the identified age-dependent speciation process may be that recently evolved taxa often colonize new regions or niches and may initially experience little competition. These new taxa are thus more likely to give rise to further new taxa than a taxon that has remained largely unchanged and is, therefore, well adapted to its niche. We show that age-dependent speciation may also be the result of different within-species populations following the same laws of lineage splitting to produce new species. As the fit of our model to the tree database shows, this simple biological motivation provides an explanation for a long standing problem in macroevolution.

When is spillover from marine reserves likely to benefit fisheries?

The net movement of individuals from marine reserves (also known as no-take marine protected areas) to the remaining fishing grounds is known as spillover and is frequently used to promote reserves to fishers on the grounds that it will benefit fisheries. Here we consider how mismanaged a fishery must be before spillover from a reserve is able to provide a net benefit for a fishery. For our model fishery, density of the species being harvested becomes higher in the reserve than in the fished area but the reduction in the density and yield of the fished area was such that the net effect of the closure was negative, except when the fishery was mismanaged. The extent to which effort had to exceed traditional management targets before reserves led to a spillover benefit varied with rates of growth and movement of the model species. In general, for well-managed fisheries, the loss of yield from the use of reserves was less for species with greater movement and slower growth. The spillover benefit became more pronounced with increasing mis-management of the stocks remaining available to the fishery. This model-based result is consistent with the literature of field-based research where a spillover benefit from reserves has only been detected when the fishery is highly depleted, often where traditional fisheries management controls are absent. We conclude that reserves in jurisdictions with well-managed fisheries are unlikely to provide a net spillover benefit.

Global distribution and conservation of evolutionary distinctness in birds.

BACKGROUND: Integrated, efficient, and global prioritization approaches are necessary to manage the ongoing loss of species and their associated function. "Evolutionary distinctness" measures a species' contribution to the total evolutionary history of its clade and is expected to capture uniquely divergent genomes and functions. Here we demonstrate how such a metric identifies species and regions of particular value for safeguarding evolutionary diversity. RESULTS: Among the world's 9,993 recognized bird species, evolutionary distinctness is very heterogeneously distributed on the phylogenetic tree and varies little with range size or threat level. Species representing the most evolutionary history over the smallest area (those with greatest "evolutionary distinctness rarity") as well as some of the most imperiled distinct species are often concentrated outside the species-rich regions and countries, suggesting they may not be well captured by current conservation planning. We perform global cross-species and spatial analyses and generate minimum conservation sets to assess the benefits of the presented species-level metrics. We find that prioritizing imperiled species by their evolutionary distinctness and geographic rarity is a surprisingly effective and spatially economical way to maintain the total evolutionary information encompassing the world's birds. We identify potential conservation gaps in relation to the existing reserve network that in particular highlight islands as effective priority areas. CONCLUSIONS: The presented distinctness metrics are effective yet easily communicable and versatile tools to assist objective global conservation decision making. Given that most species will remain ecologically understudied, combining growing phylogenetic and spatial data may be an efficient way to retain vital aspects of biodiversity.

The equivalence of two phylogenetic biodiversity measures: the Shapley value and Fair Proportion index.

Most biodiversity conservation programs are forced to prioritise species in order to allocate their funding. This paper contains a mathematical proof that provides biological support for one common approach based on phylogenetic indices. Phylogenetic trees describe the evolutionary relationships between a group of taxa. Two indices for computing the distinctiveness of each taxon in a phylogenetic tree are considered here-the Shapley value and the Fair Proportion index. These indices provide a measure of the importance of each taxon for overall biodiversity and have been used to prioritise taxa for conservation. The Shapley value is the biodiversity contribution a taxon is expected to make if all taxa are equally likely to become extinct. This interpretation makes it appealing to use the Shapley value in biodiversity conservation applications. The Fair Proportion index lacks a convenient interpretation, however it is significantly easier to calculate and understand. It has been empirically observed that there is a high correlation between the two indices. This paper shows the mathematical basis for this correlation and proves that as the number of taxa increases, the indices become equivalent. Consequently in biodiversity prioritisation the simpler Fair Proportion index can be used whilst retaining the appealing interpretation of the Shapley value.

BIO::Phylo-phyloinformatic analysis using perl.

BACKGROUND: Phyloinformatic analyses involve large amounts of data and metadata of complex structure. Collecting, processing, analyzing, visualizing and summarizing these data and metadata should be done in steps that can be automated and reproduced. This requires flexible, modular toolkits that can represent, manipulate and persist phylogenetic data and metadata as objects with programmable interfaces. RESULTS: This paper presents Bio::Phylo, a Perl5 toolkit for phyloinformatic analysis. It implements classes and methods that are compatible with the well-known BioPerl toolkit, but is independent from it (making it easy to install) and features a richer API and a data model that is better able to manage the complex relationships between different fundamental data and metadata objects in phylogenetics. It supports commonly used file formats for phylogenetic data including the novel NeXML standard, which allows rich annotations of phylogenetic data to be stored and shared. Bio::Phylo can interact with BioPerl, thereby giving access to the file formats that BioPerl supports. Many methods for data simulation, transformation and manipulation, the analysis of tree shape, and tree visualization are provided. CONCLUSIONS: Bio::Phylo is composed of 59 richly documented Perl5 modules. It has been deployed successfully on a variety of computer architectures (including various Linux distributions, Mac OS X versions, Windows, Cygwin and UNIX-like systems). It is available as open source (GPL) software from http://search.cpan.org/dist/Bio-Phylo.

Sampling trees from evolutionary models.

A wide range of evolutionary models for species-level (and higher) diversification have been developed. These models can be used to test evolutionary hypotheses and provide comparisons with phylogenetic trees constructed from real data. To carry out these tests and comparisons, it is often necessary to sample, or simulate, trees from the evolutionary models. Sampling trees from these models is more complicated than it may appear at first glance, necessitating careful consideration and mathematical rigor. Seemingly straightforward sampling methods may produce trees that have systematically biased shapes or branch lengths. This is particularly problematic as there is no simple method for determining whether the sampled trees are appropriate. In this paper, we show why a commonly used simple sampling approach (SSA)-simulating trees forward in time until n species are first reached-should only be applied to the simplest pure birth model, the Yule model. We provide an alternative general sampling approach (GSA) that can be applied to most other models. Furthermore, we introduce the constant-rate birth-death model sampling approach, which samples trees very efficiently from a widely used class of models. We explore the bias produced by SSA and identify situations in which this bias is particularly pronounced. We show that using SSA can lead to erroneous conclusions: When using the inappropriate SSA, the variance of a gradually evolving trait does not correlate with the age of the tree; when the correct GSA is used, the trait variance correlates with tree age. The algorithms presented here are available in the Perl Bio::Phylo package, as a stand-alone program TreeSample, and in the R TreeSim package.

Stochastic properties of generalised Yule models, with biodiversity applications.

The Yule model is a widely used speciation model in evolutionary biology. Despite its simplicity many aspects of the Yule model have not been explored mathematically. In this paper, we formalise two analytic approaches for obtaining probability densities of individual branch lengths of phylogenetic trees generated by the Yule model. These methods are flexible and permit various aspects of the trees produced by Yule models to be investigated. One of our methods is applicable to a broader class of evolutionary processes, namely the Bellman-Harris models. Our methods have many practical applications including biodiversity and conservation related problems. In this setting the methods can be used to characterise the expected rate of biodiversity loss for Yule trees, as well as the expected gain of including the phylogeny in conservation management. We briefly explore these applications.

Evolutionarily distinctive species often capture more phylogenetic diversity than expected.

Evolutionary distinctiveness measures of how evolutionarily isolated a species is relative to other members of its clade. Recently, distinctiveness metrics that explicitly incorporate time have been proposed for conservation prioritization. However, we found that such measures differ qualitatively in how well they capture the total amount of evolution (termed phylogenetic diversity, or PD) represented by a set of species. We used simulation and simple graph theory to explore this relationship with reference to phylogenetic tree shape. Overall, the distinctiveness measures capture more PD on more unbalanced trees and on trees with many splits near the present. The rank order of performance was robust across tree shapes, with apportioning measures performing best and node-based measures performing worst. A sample of 50 ultrametric trees from the literature showed the same patterns. Taken together, this suggests that distinctiveness metrics may be a useful addition to other measures of value for conservation prioritization of species. The simplest measure, the age of a species, performed surprisingly well, suggesting that new measures that focus on tree shape near the tips may provide a transparent alternative to more complicated full-tree approaches.

Maximizing phylogenetic diversity in biodiversity conservation: Greedy solutions to the Noah's Ark problem.

The Noah's Ark Problem (NAP) is a comprehensive cost-effectiveness methodology for biodiversity conservation that was introduced by Weitzman (1998) and utilizes the phylogenetic tree containing the taxa of interest to assess biodiversity. Given a set of taxa, each of which has a particular survival probability that can be increased at some cost, the NAP seeks to allocate limited funds to conserving these taxa so that the future expected biodiversity is maximized. Finding optimal solutions using this framework is a computationally difficult problem to which a simple and efficient "greedy" algorithm has been proposed in the literature and applied to conservation problems. We show that, although algorithms of this type cannot produce optimal solutions for the general NAP, there are two restricted scenarios of the NAP for which a greedy algorithm is guaranteed to produce optimal solutions. The first scenario requires the taxa to have equal conservation cost; the second scenario requires an ultrametric tree. The NAP assumes a linear relationship between the funding allocated to conservation of a taxon and the increased survival probability of that taxon. This relationship is briefly investigated and one variation is suggested that can also be solved using a greedy algorithm.