Species delimitation using genomic data to resolve taxonomic uncertainties in a speciation continuum of pelagic seabirds.

Speciation is a continuous and complex process shaped by the interaction of numerous evolutionary forces. Despite the continuous nature of the speciation process, the implementation of conservation policies relies on the delimitation of species and evolutionary significant units (ESUs). Puffinus shearwaters are globally distributed and threatened pelagic seabirds. Due to remarkable morphological status the group has been under intense taxonomic debate for the past three decades. Here, we use double digest Restriction-Site Associated DNA sequencing (ddRAD-Seq) to genotype species and subspecies of North Atlantic and Mediterranean Puffinus shearwaters across their entire geographical range. We assess the phylogenetic relationships and population structure among and within the group, evaluate species boundaries, and characterise the genomic landscape of divergence. We find that current taxonomies are not supported by genomic data and propose a more accurate taxonomy by integrating genomic information with other sources of evidence. Our results show that several taxon pairs are at different stages of a speciation continuum. Our study emphasises the potential of genomic data to resolve taxonomic uncertainties, which can help to focus management actions on relevant taxa, even if they do not necessarily coincide with the taxonomic rank of species.

Underwater visibility constrains the foraging behaviour of a diving pelagic seabird.

Understanding the sensory ecology of species is vital if we are to predict how they will function in a changing environment. Visual cues are fundamentally important for many predators when detecting and capturing prey. However, many marine areas have become more turbid through processes influenced by climate change, potentially affecting the ability of marine predators to detect prey. We performed the first study that directly relates a pelagic seabird species's foraging behaviour to oceanic turbidity. We collected biologging data from 79 foraging trips and 5472 dives of a visually dependent, pursuit-diving seabird, the Manx shearwater (Puffinus puffinus). Foraging behaviour was modelled against environmental variables affecting underwater visibility, including water turbidity, cloud cover and solar angle. Shearwaters were more likely to initiate area-restricted search and foraging dives in clearer waters. Underwater visibility also strongly predicted dive rate and depth, suggesting that fine-scale prey capture was constrained by the detectability of prey underwater. Our novel use of dynamic descriptors of underwater visibility suggests that visual cues are vital for underwater foraging. Our data indicate that climate change could negatively impact seabird populations by making prey more difficult to detect, compounded by the widely reported effects of reduced prey populations.

Marine subsidies likely cause gigantism of iguanas in the Bahamas.

We utilized natural experiment opportunities presented by differential conditions (presence/absence of seabirds and invasive species) on cays in the Bahamas to study whether interisland variations in food resources contributed to gigantism in Allen Cays Rock Iguanas (Cyclura cychlura inornata). We analyzed the stable carbon (δ13C) and nitrogen (δ15N) isotope values from iguana tissues and resources from each island food web to test the predictions that (1) food webs on islands with seabirds exhibit the influence of marine subsidies from seabird guano, whereas those from non-seabird islands do not, and (2) size differences in iguanas among cays were due to either (a) supplemental food availability from mice and/or seabird carcasses killed by barn owls (Tyto alba) and/or (b) access to more nutrient-rich vegetation fertilized by seabird guano. Food web components from the seabird island (Allen Cay) had 5-9‰ higher δ15N values than those on the other cays and Allen Cay plants contained nearly two times more nitrogen. Bayesian stable isotope mixing models indicated that C3 plants dominated iguana diets on all islands and showed no evidence for consumption of mice or shearwaters. The iguanas on Allen Cay were ~ 2 times longer (48.3 ± 11.6 cm) and ~ 6 times heavier (5499 ± 2847 g) than iguanas on other cays and this was likely from marine-derived subsidies from seabird guano which caused an increase in nitrogen concentration in the plants and a resultant increase in the δ15N values across the entire food web relative to non-seabird islands.

Lévy patterns in seabirds are multifaceted describing both spatial and temporal patterning.

BACKGROUND: The flight patterns of albatrosses and shearwaters have become a touchstone for much of Lévy flight research, spawning an extensive field of enquiry. There is now compelling evidence that the flight patterns of these seabirds would have been appreciated by Paul Lévy, the mathematician after whom Lévy flights are named. Here we show that Lévy patterns (here taken to mean spatial or temporal patterns characterized by distributions with power-law tails) are, in fact, multifaceted in shearwaters being evident in both spatial and temporal patterns of activity. RESULTS: We tested for Lévy patterns in the at-sea behaviours of two species of shearwater breeding in the North Atlantic Ocean (Calonectris borealis) and the Mediterranean sea (C. diomedea) during their incubating and chick-provisioning periods. We found that distributions of flight durations, on/in water durations and inter-dive time-intervals have power-law tails and so bear the hallmarks of Lévy patterns. CONCLUSIONS: The occurrence of these statistical laws is remarkable given that bird behaviours are strongly shaped by an individual's motivational state and by complex environmental interactions. Our observations could take Lévy patterns as models of animal behaviour to a new level by going beyond the characterisation of spatial movements to characterise how different behaviours are interwoven throughout daily animal life.

Exploring the pathology of liver, kidney, muscle, and stomach of fledgling seabirds associated with plastic ingestion.

There remain significant gaps in knowledge about 'sub-lethal' impacts of plastic ingestion, particularly chronic impacts on cells, tissues, or organs. Few studies have applied traditional animal health tools, such as histopathology, to assess physiological damage to wildlife, with fewer still providing information on the dosage or exposure to plastics needed to elicit negative effects. Our study seeks to investigate a common hypothesis in plastic pollution research; that an increasing plastics burden will have an impact on an animal's health, examining two wild species with high levels of environmental exposure to plastic through their diet. Here we assess the histopathology of the muscle, upper digestive tract, liver and kidney of two seabird species that are known to be commonly exposed to plastic, comparing exposed and non-exposed individuals. Fledgling seabirds showed histopathological evidence of cumulative pressures such as starvation, disease, and endoparasite burden. However, we observed no evidence of chronic harm that could be explicitly linked to the plastics. We found one case of haemorrhage, reaffirming that large/sharp plastic foreign bodies may cause acute physical damage. Given the numerous interacting pressures on the health of fledging seabirds, including exposure to plastic, this study highlights the need to scrutinise plastic-animal interactions and research though a One Health lens.

Analysis of organochlorines and polycyclic aromatic hydrocarbons designed for pollutant biomonitoring in three seabird matrices.

Pollutant biomonitoring demands analytical methods to cover a wide range of target compounds, work with minimal sample amounts, and apply least invasive and reproducible sampling procedures. We developed a method to analyse 68 bioaccumulative organic pollutants in three seabird matrices: plasma, liver, and stomach oil, representing different exposure phases. Extraction efficiency was assessed based on recoveries of spiked surrogate samples, then the method was applied to environmental samples collected from Scopoli's shearwater (Calonectris diomedea). Extraction was performed in an ultrasonic bath, purification with Florisil cartridges (5 g, 20 mL), and analysis by GC-Orbitrap-MS. Quality controls at 5 ng yielded satisfactory recoveries (80-120%) although signal intensification was found for some compounds. The method permitted the detection of 28 targeted pollutants in the environmental samples. The mean sum of organic pollutants was 4.25 ± 4.83 ng/g in plasma, 1634 ± 2990 ng/g in liver, and 233 ± 111 ng/g in stomach oil (all wet weight). Pollutant profiles varied among the matrices, although 4,4'-DDE was the dominant compound overall. This method is useful for pollutant biomonitoring in seabirds and discusses the interest of analysing different matrices.

Birds of a feather eat plastic together: high levels of plastic ingestion in Great Shearwater adults and juveniles across their annual migratory cycle.

Limited work to date has examined plastic ingestion in highly migratory seabirds like Great Shearwaters (Ardenna gravis) across the their entire migratory range, although this species is prone to ingest plastic as a wide-ranging procellariiform. We examined 217 Great Shearwaters obtained from 2008-2019 at multiple locations spanning their yearly migration cycle across the Northwest and South Atlantic to assess accumulation of ingested plastic as well as trends over time and between locations. A total of 2,328 plastic fragments were documented in the ventriculus portion of the gastrointestinal tract, with an average of 9 plastic fragments per bird. The mass, count, and frequency of plastic occurrence (FO) varied by location, with higher plastic burdens but lower FO in South Atlantic individuals from the breeding colonies. No fragments of the same size or morphology were found in the primary forage fish prey, the sand lance, (Ammodytes spp., n = 202) that supports Great Shearwaters in Massachusetts Bay, USA, suggesting the birds directly ingest the bulk of their plastic loads rather than accumulating via trophic transfer. Fourier-transform infrared spectroscopy indicated that low- and high-density polyethylene were the most common polymers ingested, within all years and locations. Individuals from the South Atlantic contained a higher proportion of larger plastic items and fragments compared to juveniles and non-breeding adults from the NW Atlantic, possibly due to increased use of remote, pelagic areas subject to reduced inputs of smaller, more diverse, and potentially less buoyant plastics found adjacent to coastal margins. Different signatures of polymer type, size, and category between similar life stages at different locations suggests rapid turnover of ingested plastics commensurate with migratory stage and location, though more empirical evidence is needed to ground-truth this hypothesis. This work is the first to comprehensively measure the accumulation of ingested plastics by Great Shearwaters over the last decade and across multiple locations spanning their yearly trans-equatorial migration cycle, and underscores their utility as sentinels of plastic pollution in Atlantic ecosystems.

Wedge-tailed Shearwater (Ardenna pacifica) nesting success in human-dominated coastal environments.

Many seabird populations are declining globally, but successful conservation efforts have led to population expansion of some species into human-dominated landscapes. Thus, there is an increased potential for direct human and seabird interactions for certain species in human-occupied areas, with nest-site characteristics potentially affecting the susceptibility of nests to human disturbance. We assessed the effect of human activity and nest-site characteristics on Wedge-tailed Shearwater (Ardenna pacifica, 'ua'u kani) nesting success at two breeding colonies, one with human exposure and one without, located in Kailua, O'ahu, Hawai'i. Human activity was measured by recording the frequency of people who entered a 5 m buffer around each nest. Nests were checked every two to three days to monitor nest success. The effect of human activity and nest-site characteristics on nesting success was determined using a variety of combinations of variables within binomial logistic regression models and AICc model selection. Nest-site characteristics among nests at both sites and human activity at the human-exposed site did not show a significant effect on nesting success. Our results suggest Wedge-tailed Shearwaters may experience some tolerance of human activity immediately around their nests-as long as burrow collapse does not occur. Given the small sample sizes and a single season of data collection, additional studies are needed to better understand the effect of human disturbance on Wedge-tailed Shearwaters. Infrastructure, such as fencing and signage, may be effective at reducing human-caused nest failure and may allow humans and disturbance-tolerant seabird species to coexist in shared coastal environments.

A baseline for POPs contamination in Australian seabirds: little penguins vs. short-tailed shearwaters.

While globally distributed throughout the world's ecosystems, there is little baseline information on persistent organic pollutants (POPs) in marine environments in Australia and, more broadly, the Southern Hemisphere. To fill this knowledge gap, we collected baseline information on POPs in migratory short-tailed shearwaters (Ardenna tenuirostris) from Fisher Island, Tasmania, and resident little penguins (Eudyptula minor) from Phillip Island, Victoria. Levels of polychlorinated biphenyls (PCBs), organochlorine pesticides (OCPs) and brominated flame retardants (BFRs) were determined from blood samples, with total contamination ranging 7.6-47.7 ng/g ww for short-tailed shearwaters and 0.12-46.9 ng/g ww for little penguins. In both species contamination followed the same pattern where PCBs>OCPs>BFRs. BFR levels included the presence of the novel flame retardant hexabromobenzene (HBB). These novel results of POPs in seabirds in southeast Australia provide important information on the local (penguins) and global (shearwaters) distribution of POPs in the marine environment.

Inter-annual changes in oceanic conditions drives spatial and trophic consistency of a tropical marine predator.

Pelagic seabirds exhibit plasticity in foraging characteristics in relation to oceanographic conditions. This should be particularly relevant in tropical marine environments where food resources are naturally more unpredictable. We studied how inter-annual variations (2013-2018) in tropical oceanographic conditions (driver of oceanic productivity) can influence the spatial and trophic ecology of Cape Verde shearwater (Calonectris edwardsii) during the breeding season. During years of poor oceanographic conditions around the colony, birds engaged in longer trips to West Africa, showed higher spatial and behavioural consistency, and presented a wider isotopic niche. Opposite patterns were generally found for years of good oceanographic conditions, when birds foraged more on their colony surroundings. New foraging areas off West Africa were highlighted as relevant, especially during years of poor environmental conditions. This study highlights the need for long-term studies to assess variation in foraging areas and foraging decisions by seabird populations.

Are pelagic seabirds exposed to amnesic shellfish poisoning toxins?

Marine birds have been hypothesized to be underreported victims of harmful algal blooms (HABs). Toxic blooms of Pseudo-nitzschia spp., the primary amnesic toxin producer microalgae, domoic acid (DA) are known to cause massive mortalities of coastal seabirds and marine mammals around the world. However, these fatalities are only detected when birds die nearby the coastline and little is known about possible outbreaks of pelagic seabirds in oceanic areas. Here we aim to understand whether pelagic seabirds are exposed to amnesic shellfish poisoning (ASP) toxins. For this purpose, we tracked pelagic seabirds feeding on small epipelagic fish and squid, reported to be vectors of DA, which are obtained in high productivity zones where intense Pseudo-nitzschia blooms regularly occur. In particular, we tracked Cory's (Calonectris borealis) and Scopoli's (C. diomedea) shearwaters breeding in Gran Canaria (Canary Is.) and in Menorca (Balearic Is.) and feeding on the Canary Current region and the Catalonian coast, respectively. We sampled birds for blood at the recovery of the GPS (Global Positioning System) and analyzed it for DA determination by Liquid Chromatography coupled with Tandem Mass Spectrometry (LC-MS/MS). Among the 61 samples analyzed from Gran Canaria, and 87 from Menorca, 31 (50.8%) and 28 (32.2%) from each location presented detectable levels of DA ranging 1.0-10.6 ng mL-1. This work reveals that DA can be detected at variable levels in the blood of ASP-asymptomatic shearwaters and suggests a chronic exposure of shearwaters to DA, highlighting the need for further studies on DA effects. These results are of high relevance due to the vulnerability of these marine birds, which populations are in continuous decline. Since global warming is expected to alter and increase the occurrence of HABs, marine toxins might become an additional stressor for seabirds and exacerbate the already precarious conservation status of many species.

The effect of parental condition on egg-size and reproductive success in short-tailed shearwaters Puffinus tenuirostris.

Eggs were exchanged between 50 pairs of shorttailed shearwaters Puffinus tenuirostris on Great Dog Island, Bass Strait, Australia, in an attempt to distinguish the intrinsic effects of egg-size from any effects stemming from differential quality of parental care. At 64 "experimental" nests, large and small eggs were exchanged whereas at 36 "control" nests, eggs of equivalent, medium, size were exchanged. Egg-size appeared independent of maternal effects. In both groups, hatching and fledging success were independent both of eggsize and of the body condition of the attending parents. This suggests that breeding success in these birds may be more closely related to the behavioural traits of parents than to physiological factors.