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Summary The concordance of evolutionary histories and extant species interactions provides a useful metric for addressing questions of how the structure of ecological communities is influenced by macro-evolutionary processes. We introduce paco (v0.3.1), an R package to perform Procrustean Approach to Cophylogeny. This method assesses the phylogenetic congruence, or evolutionary dependence, of two groups of interacting species using both ecological interaction networks and their phylogenetic history. We demonstrate the functionality of paco through its application to empirical host-parasite and plant-pollinator communities. Although the package is intended to assess the phylogenetic congruence between groups of interacting species, the method is also directly applicable to other scenarios that may show phylogenetic congruence including historical biogeography, molecular systematics, and cultural evolution.

What determines whether or not a species is a generalist or a specialist? Evidence that the environment can influence species interactions is rapidly accumulating. However, a systematic link between environment and the number of partners a species interacts with has been elusive so far. Presumably, because environmental gradients appear to have contrasting effects on species depending on the environmental variable. Here, we test for a relationship between the stresses imposed by the environment, instead of environmental gradients directly, and species specialisation using a global dataset of plant-pollinator interactions. We found that the environment can play a significant effect on specialisation, even when accounting for community composition, likely by interacting with species’ traits and evolutionary history. Species that have a large number of interactions are more likely to focus on a smaller number of, presumably higher-quality, interactions under stressful environmental conditions. Contrastingly, the specialists present in multiple locations are more likely to broaden their niche, presumably engaging in opportunistic interactions to cope with increased environmental stress. Indeed, many apparent specialists effectively behave as facultative generalists. Overall, many of the species we analysed are not inherently generalist or specialist. Instead, species’ level of specialisation should be considered on a relative scale depending on where they are found and the environmental conditions at that location.

Whale sharks (Rhincodon typus) are typically dispersed throughout their circumtropical range, but the species is also known to aggregate in specific coastal areas. Accurate site descriptions associated with these aggregations are essential for the conservation of R. typus, an Endangered species. Although aggregations have become valuable hubs for research, most site descriptions rely heavily on sightings data. In the present study, visual census, passive acoustic monitoring, and long range satellite telemetry were combined to track the movements of R. typus from Shib Habil, a reef-associated aggregation site in the Red Sea. An array of 63 receiver stations was used to record the presence of 84 acoustically tagged sharks (35 females, 37 males, 12 undetermined) from April 2010 to May 2016. Over the same period, identification photos were taken for 76 of these tagged individuals and 38 were fitted with satellite transmitters. In total of 37,461 acoustic detections, 210 visual encounters, and 33 satellite tracks were analyzed to describe the sharks’ movement ecology. The results demonstrate that the aggregation is seasonal, mostly concentrated on the exposed side of Shib Habil, and seems to attract sharks of both sexes in roughly equal numbers. The combined methodologies also tracked 15 interannual homing-migrations, demonstrating that many sharks leave the area before returning in later years. When compared to acoustic studies from other aggregations, these results demonstrate that R. typus exhibits diverse, site-specific ecologies across its range. Sightings-independent data from acoustic telemetry and other sources are an effective means of validating more common visual surveys.

Many different concepts have been used to describe species' roles in food webs (i.e., the ways in which species participate in their communities as consumers and resources). As each concept focuses on a different aspect of food-web structure, it can be difficult to relate these concepts to each other and to other aspects of ecology. Here we use the Eltonian niche as an overarching framework, within which we summarize several commonly-used role concepts (degree, trophic level, motif roles, and centrality). We focus mainly on the topological versions of these concepts but, where dynamical versions of a role concept exist, we acknowledge these as well. Our aim is to highlight areas of overlap and ambiguity between different role concepts and to describe how these roles can be used to group species according to different strategies (i.e., equivalence and functional roles). The existence of “gray areas” between role concepts make it essential for authors to carefully consider both which role concept(s) are most appropriate for the analyses they wish to conduct and what aspect of species' niches (if any) they wish to address. The ecological meaning of differences between species' roles can change dramatically depending on which role concept(s) are used.

Background Off Mafia Island, Tanzania, whale sharks are seen in a small bay that also boasts one of the country's largest coastal fisheries. We set out to determine the population structure of whale sharks here, examine their feeding behaviour and residency, and investigate the interaction between the sharks and fishers. Approach We conducted 261 boat-based surveys at Mafia Island between Oct 2012 and Dec 2015, recording all whale shark sightings as well as counting the numbers of tourist and fishing boats in the survey area during each trip. We estimated shark size visually and with laser photogrammetry and recorded sex, scars and behaviour of each shark. To examine residency, we deployed 67 acoustic tags on whale sharks and 19 receiver stations within the bay, and analysed recapture probability (both visual- and acoustic-based) with generalised additive mixing models. To investigate feeding plankton biomass and community structure, we collected 20 plankton samples when whale sharks were feeding and 202 background samples for comparison. We interviewed fishers and they recorded their fishing locations using a GPS. Results We identified 129 different whale sharks at Mafia Island, 54% of which were resighted in two or more years. Tagged sharks were acoustically detected throughout the year even though sightings were largely limited to Oct-Mar. Sharks ranged in total length from 2.5–9.7 m, were mostly male (87%), and were usually seen feeding (75%) at or near the surface. Dense patches (25 mg m–3) of large sergestids Lucifer hanseni prompted feeding, while background plankton tows were dominated by small copepods (2.5 mg m–3). Small and medium-sized planktivorous fishes also fed on the same patches of sergestids, often together with whale sharks. These smaller species were the main target of the local fishers. Fishing boats operated in the survey area on 90% of the time, but also made catches further afield. Fishers reported a conflict between whale sharks and their fishing operations on 36% of the time, although sharks were usually released from the nets without injury. A mean of 2.6 tourist boats per day searched the same survey area for whale sharks. Conclusions This small, resident group of whale sharks is routinely exposed to tourists, boat traffic and high fishing pressure. Whale sharks are commonly encircled in fishing nets and released, but scars indicate that they do occasionally get injured during this process. Mortalities are rare, based on interviews with fishers and long-term observers. Whale sharks at Mafia Island are unusually resident and face clear local threats. No defined management measures are currently in place, but we present some potential strategies for mitigating threats to whale sharks here.

Background A recent global study of whale shark population genetics has allowed for better understanding of genetic connections between aggregations in both the Indo-Pacific and Atlantic. This overview included an aggregation found within the Red Sea near Al Lith, Saudi Arabia, however the Mafia Island, Tanzania, aggregation was not part of the study. The ecological behavior of these aggregations differs with the Saudi Arabian individuals showing strong seasonality, while acoustic telemetry data revealed cryptic residency at Mafia Island. Approach Genetic analysis using 11 microsatellite markers was performed on whale sharks from both locations. A combination of primers sourced from previous studies and newly designed primers were used to compare both aggregations and the individuals within. The Red Sea population was compared between 5 seasons spanning 6 years from 2010–2015. The Tanzanian population was compared for 2 field seasons from 2012–2014. Temporal genetic diversity was examined using allelic richness on only the Saudi Arabian individuals due to a short sampling period in Tanzania. Kinship for both aggregations was tested using COLONY and KINALYZER. Results Over a 6 year period, genetic diversity in the Red Sea showed no significant change. Contrasting to other whale shark aggregations, allelic richness in the Red Sea shows no sign of reduction. Kinship analysis using COLONY found two potential sibling pairs in Tanzania. One pair had a high probability (.993) of being a full sibling dyad while the other had a lower probability (.357). There were no sibling pairs identified in the Red Sea. Conclusions The lack of significant change of genetic diversity in Al Lith, Saudi Arabia, differs from a trend at Ningaloo Reef, Australia that showed a decrease in genetic diversity. Although these differences could be driven by location, this should encourage further long term genetic sampling at aggregations to better understand whale shark population trends. The potential of sibling pairs being found within one aggregation warrants further investigation into kinship within and between aggregations throughout the Indo-Pacific.

Although whale sharks ( Rhincodon typus ) have been documented to move thousands of kilometres, they are most frequently observed at a few predictable seasonal aggregation sites. The absence of sharks at the surface during visual surveys has led to the assumption that sharks disperse to places unknown during the long ‘off-seasons’ at most of these locations. Here we compare 2 years of R. typus visual sighting records from Mafia Island in Tanzania to concurrent acoustic telemetry of tagged individuals. Sightings revealed a clear seasonal pattern with a peak between October and February and no sharks observed at other times. By contrast, acoustic telemetry demonstrated year-round residency of R. typus . The sharks use a different habitat in the off-season, swimming deeper and further away from shore, presumably in response to prey distributions. This behavioural change reduces the sharks' visibility, giving the false impression that they have left the area. We demonstrate, for the first time to our knowledge, year-round residency of unprovisioned, individual R. typus at an aggregation site, and highlight the importance of using multiple techniques to study the movement ecology of marine megafauna.

Large planktivores require high-density prey patches to make feeding energetically viable. This is a major challenge for species living in tropical and subtropical seas, such as whale sharks Rhincodon typus. Here, we characterize zooplankton biomass, size structure and taxonomic composition from whale shark feeding events and background samples at Mafia Island, Tanzania. The majority of whale sharks were feeding (73%, 380 of 524 observations), with the most common behaviour being active surface feeding (87%). We used 20 samples collected from immediately adjacent to feeding sharks and an additional 202 background samples for comparison to show that plankton biomass was ∼10 times higher in patches where whale sharks were feeding (25 vs. 2.6 mg m−3). Taxonomic analyses of samples showed that the large sergestid Lucifer hanseni (∼10 mm) dominated while sharks were feeding, accounting for ∼50% of identified items, while copepods (