Avoiding bias in estimates of population size for translocation management.

Mark-recapture surveys are commonly used to monitor translocated populations globally. Data gathered are then used to estimate demographic parameters, such as abundance and survival, using Jolly-Seber (JS) models. However, in translocated populations initial population size is known and failure to account for this may bias parameter estimates, which are important for informing conservation decisions during population establishment. Here, we provide methods to account for known initial population size in JS models by incorporating a separate component likelihood for translocated individuals, using a maximum-likelihood estimation, with models that can be fitted using either R or MATLAB. We use simulated data and a case study of a threatened lizard species with low capture probability to demonstrate that unconstrained JS models may overestimate the size of translocated populations, especially in the early stages of post-release monitoring. Our approach corrects this bias; we use our simulations to demonstrate that overestimates of population size between 78% and 130% can occur in the unconstrained JS models when the detection probability is below 0.3 compared to 1%-8.9% for our constrained model. Our case study did not show an overestimate; however accounting for the initial population size greatly reduced error in all parameter estimates and prevented boundary estimates. Adopting the corrected JS model for translocations will help managers to obtain more robust estimates of the population sizes of translocated animals, better informing future management including reinforcement decisions, and ultimately improving translocation success.

Seabird morphology determines operational wind speeds, tolerable maxima, and responses to extremes.

Storms can cause widespread seabird stranding and wrecking,1,2,3,4,5 yet little is known about the maximum wind speeds that birds are able to tolerate or the conditions they avoid. We analyzed >300,000 h of tracking data from 18 seabird species, including flapping and soaring fliers, to assess how flight morphology affects wind selectivity, both at fine scales (hourly movement steps) and across the breeding season. We found no general preference or avoidance of particular wind speeds within foraging tracks. This suggests seabird flight morphology is adapted to a "wind niche," with higher wing loading being selected in windier environments. In support of this, wing loading was positively related to the median wind speeds on the breeding grounds, as well as the maximum wind speeds in which birds flew. Yet globally, the highest wind speeds occur in the tropics (in association with tropical cyclones) where birds are morphologically adapted to low median wind speeds. Tropical species must therefore show behavioral responses to extreme winds, including long-range avoidance of wind speeds that can be twice their operable maxima. By contrast, Procellariiformes flew in almost all wind speeds they encountered at a seasonal scale. Despite this, we describe a small number of cases where albatrosses avoided strong winds at close range, including by flying into the eye of the storm. Extreme winds appear to pose context-dependent risks to seabirds, and more information is needed on the factors that determine the hierarchy of risk, given the impact of global change on storm intensity.6,7.

Ecological inference using data from accelerometers needs careful protocols.

Accelerometers in animal-attached tags are powerful tools in behavioural ecology, they can be used to determine behaviour and provide proxies for movement-based energy expenditure. Researchers are collecting and archiving data across systems, seasons and device types. However, using data repositories to draw ecological inference requires a good understanding of the error introduced according to sensor type and position on the study animal and protocols for error assessment and minimisation.Using laboratory trials, we examine the absolute accuracy of tri-axial accelerometers and determine how inaccuracies impact measurements of dynamic body acceleration (DBA), a proxy for energy expenditure, in human participants. We then examine how tag type and placement affect the acceleration signal in birds, using pigeons Columba livia flying in a wind tunnel, with tags mounted simultaneously in two positions, and back- and tail-mounted tags deployed on wild kittiwakes Rissa tridactyla. Finally, we present a case study where two generations of tag were deployed using different attachment procedures on red-tailed tropicbirds Phaethon rubricauda foraging in different seasons.Bench tests showed that individual acceleration axes required a two-level correction to eliminate measurement error. This resulted in DBA differences of up to 5% between calibrated and uncalibrated tags for humans walking at a range of speeds. Device position was associated with greater variation in DBA, with upper and lower back-mounted tags varying by 9% in pigeons, and tail- and back-mounted tags varying by 13% in kittiwakes. The tropicbird study highlighted the difficulties of attributing changes in signal amplitude to a single factor when confounding influences tend to covary, as DBA varied by 25% between seasons.Accelerometer accuracy, tag placement and attachment critically affect the signal amplitude and thereby the ability of the system to detect biologically meaningful phenomena. We propose a simple method to calibrate accelerometers that can be executed under field conditions. This should be used prior to deployments and archived with resulting data. We also suggest a way that researchers can assess accuracy in previously collected data, and caution that variable tag placement and attachment can increase sensor noise and even generate trends that have no biological meaning.

The role of wingbeat frequency and amplitude in flight power.

Body-mounted accelerometers provide a new prospect for estimating power use in flying birds, as the signal varies with the two major kinematic determinants of aerodynamic power: wingbeat frequency and amplitude. Yet wingbeat frequency is sometimes used as a proxy for power output in isolation. There is, therefore, a need to understand which kinematic parameter birds vary and whether this is predicted by flight mode (e.g. accelerating, ascending/descending flight), speed or morphology. We investigate this using high-frequency acceleration data from (i) 14 species flying in the wild, (ii) two species flying in controlled conditions in a wind tunnel and (iii) a review of experimental and field studies. While wingbeat frequency and amplitude were positively correlated, R2 values were generally low, supporting the idea that parameters can vary independently. Indeed, birds were more likely to modulate wingbeat amplitude for more energy-demanding flight modes, including climbing and take-off. Nonetheless, the striking variability, even within species and flight types, highlights the complexity of describing the kinematic relationships, which appear sensitive to both the biological and physical context. Notwithstanding this, acceleration metrics that incorporate both kinematic parameters should be more robust proxies for power than wingbeat frequency alone.

Individual consistency in migration strategies of a tropical seabird, the Round Island petrel.

BACKGROUND: In migratory species, the extent of within- and between-individual variation in migratory strategies can influence potential rates and directions of responses to environmental changes. Quantifying this variation requires tracking of many individuals on repeated migratory journeys. At temperate and higher latitudes, low levels of within-individual variation in migratory behaviours are common and may reflect repeated use of predictable resources in these seasonally-structured environments. However, variation in migratory behaviours in the tropics, where seasonal predictability of food resources can be weaker, remains largely unknown. METHODS: Round Island petrels (Pterodroma sp.) are tropical, pelagic seabirds that breed all year round and perform long-distance migrations. Using multi-year geolocator tracking data from 62 individuals between 2009 and 2018, we quantify levels of within- and between-individual variation in non-breeding distributions and timings. RESULTS: We found striking levels of between-individual variation in at-sea movements and timings, with non-breeding migrations to different areas occurring across much of the Indian Ocean and throughout the whole year. Despite this, repeat-tracking of individual petrels revealed remarkably high levels of spatial and temporal consistency in within-individual migratory behaviour, particularly for petrels that departed at similar times in different years and for those departing in the austral summer. However, while the same areas were used by individuals in different years, they were not necessarily used at the same times during the non-breeding period. CONCLUSIONS: Even in tropical systems with huge ranges of migratory routes and timings, our results suggest benefits of consistency in individual migratory behaviours. Identifying the factors that drive and maintain between-individual variation in migratory behaviour, and the consequences for breeding success and survival, will be key to understanding the consequences of environmental change across migratory ranges.

DNA metabarcoding reveals introduced species predominate in the diet of a threatened endemic omnivore, Telfair's skink (Leiolopisma telfairii).

Introduced species can exert disproportionately negative effects on island ecosystems, but their potential role as food for native consumers is poorly studied. Telfair's skinks are endemic omnivores living on Round Island, Mauritius, a globally significant site of biodiversity conservation. We aimed to determine the dietary diversity and key trophic interactions of Telfair's skinks, whether introduced species are frequently consumed, and if diet composition changes seasonally between male and female skinks. We used DNA metabarcoding of skink fecal samples to identify animals (COI) and plants (ITS2) consumed by skinks. There were 389 dietary presence counts belonging to 77 dietary taxa found across the 73 Telfair's skink fecal samples. Introduced taxa were cumulatively consumed more frequently than other categories, accounting for 49.4% of all detections, compared to cryptogenic (20.6%), native (20.6%), and endemic taxa (9.5%). The most frequently consumed introduced species was the ant, Pheidole megacephala, present in 40% of samples. Blue latan palm, Latania loddigesii, was the most frequently consumed endemic species, present in 33% of samples but was only detected in the dry season, when fruits are produced. We found a strong seasonal difference in diet composition explained by the presence of certain plant species solely or primarily in one season and a marked increase in the consumption of animal prey in the dry season. Male and female skinks consumed several taxa at different frequencies. These results present a valuable perspective on the role of introduced species in the trophic network of their invaded ecosystem. Both native and introduced species provide nutritional resources for skinks, and this may have management implications in the context of species conservation and island restoration.

How often should dead-reckoned animal movement paths be corrected for drift?

BACKGROUND: Understanding what animals do in time and space is important for a range of ecological questions, however accurate estimates of how animals use space is challenging. Within the use of animal-attached tags, radio telemetry (including the Global Positioning System, 'GPS') is typically used to verify an animal's location periodically. Straight lines are typically drawn between these 'Verified Positions' ('VPs') so the interpolation of space-use is limited by the temporal and spatial resolution of the system's measurement. As such, parameters such as route-taken and distance travelled can be poorly represented when using VP systems alone. Dead-reckoning has been suggested as a technique to improve the accuracy and resolution of reconstructed movement paths, whilst maximising battery life of VP systems. This typically involves deriving travel vectors from motion sensor systems and periodically correcting path dimensions for drift with simultaneously deployed VP systems. How often paths should be corrected for drift, however, has remained unclear. METHODS AND RESULTS: Here, we review the utility of dead-reckoning across four contrasting model species using different forms of locomotion (the African lion Panthera leo, the red-tailed tropicbird Phaethon rubricauda, the Magellanic penguin Spheniscus magellanicus, and the imperial cormorant Leucocarbo atriceps). Simulations were performed to examine the extent of dead-reckoning error, relative to VPs, as a function of Verified Position correction (VP correction) rate and the effect of this on estimates of distance moved. Dead-reckoning error was greatest for animals travelling within air and water. We demonstrate how sources of measurement error can arise within VP-corrected dead-reckoned tracks and propose advancements to this procedure to maximise dead-reckoning accuracy. CONCLUSIONS: We review the utility of VP-corrected dead-reckoning according to movement type and consider a range of ecological questions that would benefit from dead-reckoning, primarily concerning animal-barrier interactions and foraging strategies.

Persistent organic pollutants in sea bird eggs from the Indian Ocean's Mascarene Basin.

We report the concentrations of persistent organic pollutants (POPs) in seabird eggs from St. Brandon's Atoll, a tropical island system in the western Indian Ocean. Ten eggs each of sooty terns (Onychoprion fuscatus), fairy terns (Gygis alba), and common noddies (Anous stolidus) were collected from the atoll. For a terrestrial reference, we analysed three feral chicken (Gallus gallus domesticus) eggs from the same location. Sooty tern eggs contained the highest mean concentrations of three chemical classes: Æ©CHL3 (0.21 ng/g wm; wet mass), Æ©PCB10 (1.5 ng/g wm), and Æ©PBDE6 (1.1 ng/g wm). Fairy tern eggs contained the highest mean concentrations of HCB (0.68 ng/g wm) and Æ©CHB5 (0.83 ng/g wm). The chicken eggs contained the highest mean concentrations of Æ©DDT3 (2.6 ng/g wm), while common noddy eggs contained the highest mean concentrations of Æ©HCH2 (0.5 ng/g wm). We surmise that the differences in chemical composition between species reflect different pollutant compositions in prey from the bird's different foraging ranges. The sooty terns foraging offshore contained higher POPs concentrations than the nearshore-foraging common noddies. Fairy tern eggs contained intermediate concentrations, commensurate with their intermediate foraging. Inter-island differences in contaminant concentrations were seen between eggs of the common noddies from St. Brandon's Atoll and Rodrigues Island, 520 km to the south-east. Concentrations of contaminants found in this study were lower than values quantified by other studies, making St. Brandon's Atoll an ideal reference site to monitor background concentrations of POPs in the tropical Indian Ocean.

An "orientation sphere" visualization for examining animal head movements.

Animal behavior is elicited, in part, in response to external conditions, but understanding how animals perceive the environment and make the decisions that bring about these behavioral responses is challenging.Animal heads often move during specific behaviors and, additionally, typically have sensory systems (notably vision, smell, and hearing) sampling in defined arcs (normally to the front of their heads). As such, head-mounted electronic sensors consisting of accelerometers and magnetometers, which can be used to determine the movement and directionality of animal heads (where head "movement" is defined here as changes in heading [azimuth] and/or pitch [elevation angle]), can potentially provide information both on behaviors in general and also clarify which parts of the environment the animals might be prioritizing ("environmental framing").We propose a new approach to visualize the data of such head-mounted tags that combines the instantaneous outputs of head heading and pitch in a single intuitive spherical plot. This sphere has magnetic heading denoted by "longitude" position and head pitch by "latitude" on this "orientation sphere" (O-sphere).We construct the O-sphere for the head rotations of a number of vertebrates with contrasting body shape and ecology (oryx, sheep, tortoises, and turtles), illustrating various behaviors, including foraging, walking, and environmental scanning. We also propose correcting head orientations for body orientations to highlight specific heading-independent head rotation, and propose the derivation of O-sphere-metrics, such as angular speed across the sphere. This should help identify the functions of various head behaviors.Visualizations of the O-sphere provide an intuitive representation of animal behavior manifest via head orientation and rotation. This has ramifications for quantifying and understanding behaviors ranging from navigation through vigilance to feeding and, when used in tandem with body movement, should provide an important link between perception of the environment and response to it in free-ranging animals.

Perfluoroalkyl substances (PFAS) in tern eggs from St. Brandon's Atoll, Indian Ocean.

Per- and polyfluoroalkyl substances (PFAS) are anthropogenic fluorinated compounds of concern for human and environmental health. There is no data on PFAS concentrations in marine bird eggs from the Western Indian Ocean. We analysed eight PFAS in eggs of fairy terns (Gygis alba), sooty terns (Onychoprion fuscatus), and common noddies (Anous stolidus) from St. Brandon's Atoll. Fairy tern eggs contained the highest concentrations, followed by sooty terns and common noddies. Perfluoroundecanoic acid (PFUdA) had the highest mean concentration (2.3 ng/g wm), followed by perfluorooctane sulfonic acid (PFOS) (2.0 ng/g wm), and perfluorononanoic acid (PFNA) (0.93 ng/g wm) in fairy tern eggs. Concentrations of all PFAS were lower than values found in literature. PFOS and PFOA concentrations were three orders of magnitude lower than toxicity reference values and levels of lowest-observed-adverse-effect-level concentrations. Eggs from St. Brandon's would be useful to monitor background changes on a regional and perhaps global scale.

New universal ITS2 primers for high-resolution herbivory analyses using DNA metabarcoding in both tropical and temperate zones.

DNA metabarcoding is a rapidly growing technique for obtaining detailed dietary information. Current metabarcoding methods for herbivory, using a single locus, can lack taxonomic resolution for some applications. We present novel primers for the second internal transcribed spacer of nuclear ribosomal DNA (ITS2) designed for dietary studies in Mauritius and the UK, which have the potential to give unrivalled taxonomic coverage and resolution from a short-amplicon barcode. In silico testing used three databases of plant ITS2 sequences from UK and Mauritian floras (native and introduced) totalling 6561 sequences from 1790 species across 174 families. Our primers were well-matched in silico to 88% of species, providing taxonomic resolution of 86.1%, 99.4% and 99.9% at the species, genus and family levels, respectively. In vitro, the primers amplified 99% of Mauritian (n = 169) and 100% of UK (n = 33) species, and co-amplified multiple plant species from degraded faecal DNA from reptiles and birds in two case studies. For the ITS2 region, we advocate taxonomic assignment based on best sequence match instead of a clustering approach. With short amplicons of 187-387 bp, these primers are suitable for metabarcoding plant DNA from faecal samples, across a broad geographic range, whilst delivering unparalleled taxonomic resolution.

The flip-or-flop boutique: Marine debris on the shores of St Brandon's rock, an isolated tropical atoll in the Indian Ocean.

Isolated coral atolls are not immune from marine debris accumulation. We identified Southeast Asia, the Indian sub-continent, and the countries on the Arabian Sea as most probable source areas of 50 000 items on the shores of St. Brandon's Rock (SBR), Indian Ocean. 79% of the debris was plastics. Flip-flops, energy drink bottles, and compact fluorescent lights (CFLs) were notable item types. The density of debris (0.74 m(-)(1) shore length) is comparable to similar islands but less than mainland sites. Intact CFLs suggests product-facilitated long-range transport of mercury. We suspect that aggregated marine debris, scavenged by the islands from currents and gyres, could re-concentrate pollutants. SBR islets accumulated debris types in different proportions suggesting that many factors act variably on different debris types. Regular cleaning of selected islets will take care of most of the accumulated debris and may improve the ecology and tourism potential. However, arrangements and logistics require more study.

High risks of losing genetic diversity in an endemic Mauritian gecko: implications for conservation.

Genetic structure can be a consequence of recent population fragmentation and isolation, or a remnant of historical localised adaptation. This poses a challenge for conservationists since misinterpreting patterns of genetic structure may lead to inappropriate management. Of 17 species of reptile originally found in Mauritius, only five survive on the main island. One of these, Phelsuma guimbeaui (lowland forest day gecko), is now restricted to 30 small isolated subpopulations following severe forest fragmentation and isolation due to human colonisation. We used 20 microsatellites in ten subpopulations and two mitochondrial DNA (mtDNA) markers in 13 subpopulations to: (i) assess genetic diversity, population structure and genetic differentiation of subpopulations; (ii) estimate effective population sizes and migration rates of subpopulations; and (iii) examine the phylogenetic relationships of haplotypes found in different subpopulations. Microsatellite data revealed significant population structure with high levels of genetic diversity and isolation by distance, substantial genetic differentiation and no migration between most subpopulations. MtDNA, however, showed no evidence of population structure, indicating that there was once a genetically panmictic population. Effective population sizes of ten subpopulations, based on microsatellite markers, were small, ranging from 44 to 167. Simulations suggested that the chance of survival and allelic diversity of some subpopulations will decrease dramatically over the next 50 years if no migration occurs. Our DNA-based evidence reveals an urgent need for a management plan for the conservation of P. guimbeaui. We identified 18 threatened and 12 viable subpopulations and discuss a range of management options that include translocation of threatened subpopulations to retain maximum allelic diversity, and habitat restoration and assisted migration to decrease genetic erosion and inbreeding for the viable subpopulations.

Ecological effects of the invasive giant madagascar day gecko on endemic mauritian geckos: applications of binomial-mixture and species distribution models.

The invasion of the giant Madagascar day gecko Phelsuma grandis has increased the threats to the four endemic Mauritian day geckos (Phelsuma spp.) that have survived on mainland Mauritius. We had two main aims: (i) to predict the spatial distribution and overlap of P. grandis and the endemic geckos at a landscape level; and (ii) to investigate the effects of P. grandis on the abundance and risks of extinction of the endemic geckos at a local scale. An ensemble forecasting approach was used to predict the spatial distribution and overlap of P. grandis and the endemic geckos. We used hierarchical binomial mixture models and repeated visual estimate surveys to calculate the abundance of the endemic geckos in sites with and without P. grandis. The predicted range of each species varied from 85 km2 to 376 km2. Sixty percent of the predicted range of P. grandis overlapped with the combined predicted ranges of the four endemic geckos; 15% of the combined predicted ranges of the four endemic geckos overlapped with P. grandis. Levin's niche breadth varied from 0.140 to 0.652 between P. grandis and the four endemic geckos. The abundance of endemic geckos was 89% lower in sites with P. grandis compared to sites without P. grandis, and the endemic geckos had been extirpated at four of ten sites we surveyed with P. grandis. Species Distribution Modelling, together with the breadth metrics, predicted that P. grandis can partly share the equivalent niche with endemic species and survive in a range of environmental conditions. We provide strong evidence that smaller endemic geckos are unlikely to survive in sympatry with P. grandis. This is a cause of concern in both Mauritius and other countries with endemic species of Phelsuma.

Patterns of coccidial prevalence in lizards of Mauritius.

This paper reports prevalence of coccidial oocysts in fecal samples from 6 endemic and 2 introduced lizard species on Mauritius, an island nation in the Indian Ocean. Total coccidian prevalence was 54% (n = 341) for the endemic 'ornate day gecko,' Phelsuma ornata; 48% (n = 159) for the endemic 'Durrell's night gecko,' Nactus durrelli; 53% (n = 15) for the endemic 'Serpent Island night gecko,' N. serpensinsula; and 78% (n = 248) for the introduced gecko, Hemidactylus frenatus. These high prevalences may reflect lack of long-lasting immune response to coccidial infection. There were few significant differences in prevalence among age, island, sex, or body condition within species, suggesting that these coccidia are relatively nonpathogenic and have little effect on host fitness. Prevalence was higher in the dry season than the wet season. These data suggest other factors, such as low host immune response to reinfection, affect overall prevalence more significantly than the effect of humidity on oocyst survival on Mauritius. No coccidia were found in samples from the endemic 'Gunner's Quoin night gecko,' N. coindemirensis (n = 155), probably reflecting parasite extinction due to a host population bottleneck following historical introduction of rats. There was no evidence of competitive or facilitative interactions between Eimeria sp. and Isospora sp., but evidence of competition between 2 Eimeria species in the 'ornate day gecko,' Phelsuma ornata. No evidence was found of cross-species infection, suggesting that reptile coccidia have high host specificity and are, therefore, poor subjects for studies of parasite-mediated competition and the evolution of sex.