Quantifying movement patterns for shark conservation at remote coral atolls in the Indian Ocean

Grey reef sharks (Carcharhinus amblyrhynchos) are apex predators found on many Indo-Pacific coral reefs, but little is known about their movement patterns and habitat requirements. We used acoustic telemetry to determine movements and habitat use of these sharks at the isolated Rowley Shoals atolls, 250 km off the coast of north-western Australia. We equipped 12 male and 14 female sharks ranging from 0.79 to 1.69 m in total length with transmitters that were detected by an array of 11 strategically placed receivers on two atoll reefs. Over 26,000 detections were recorded over the 325 days of receiver deployment. No sharks were observed to move between reefs. Receivers on the outer slopes of reefs provided nearly all (99%) of the detections. We found no differences in general attendance parameters due to size, sex or reef, except for maximum period of detection where larger sharks were detected over a longer period than smaller sharks. Male and female sharks were often detected at separate receivers at the outer slope habitat of one reef, suggesting sexual segregation, but this pattern did not occur at the second reef where males and females were detected at similar frequencies. We identified two patterns of daily behaviour: (1) sharks were present at the reef both day and night or (2) sharks spent more time in attendance during day than at night. Fast Fourier transforms identified 24-h cycles of attendance at the reef and a secondary peak of attendance at 12 h for most sharks, although no individuals shared the same attendance patterns. Our study provides baseline data that can be used to optimise the minimum area and habitat requirements for conservation of these apex predators.     

Performance of remote acoustic receivers within a coral reef habitat: implications for array design

Remote monitoring technologies are increasingly being implemented in the marine environment to better understand the movement patterns of taxa. Coral reefs are no exception. However, there is a paucity of information relating to the performance of acoustic receivers on coral reefs. Our results suggest that the detection performance of acoustic receivers may be significantly impacted by the unique nature of the reef environment. This study assessed the performance of passive acoustic receivers on a typical inner-shelf fringing reef, Orpheus Island, on the Great Barrier Reef, Australia. The detection range and diel performance variability of acoustic receivers was assessed using two parallel lines of 5 VR2W receivers spanning 125?m, deployed on the reef base and reef crest. Two 9-mm acoustic transmitters were moored at opposite ends of each receiver line. The working detection range for receivers was found to be approximately 90?m for the transmitter moored on the reef base and just 60?m for the transmitter moored on the reef crest. However, the detection range on the reef crest increased to 90?m when just the reef crest receivers were considered, highlighting importance of optimal receiver deployment. No diel patterns in receiver performance or detection capacities were detected, suggesting that no corrections are required when interpreting nocturnal versus diurnal activity patterns. We suggest that studies aiming for complete coverage of a site within a reef environment will require receivers in close (<100?m) proximity, and that the placement depth of receivers must be a major consideration, with shallow receivers exhibiting a greater detection range than those on the reef slope. Our results highlight the challenges imposed by coral reefs for acoustic telemetry and the importance of receiver placement for studies conducted within these habitats.     

Markov models and network analysis reveal sex‐specific differences in the space‐use of a coastal apex predator

Understanding the links between external variables such as habitat and interactions with conspecifics and animal space‐use is fundamental to developing effective management measures. In the marine realm, automated acoustic tracking has become a widely used method for monitoring the movement of free‐ranging animals, yet researchers generally lack robust methods for analysing the resulting spatial‐usage data. In this study, acoustic tracking data from male and female broadnose sevengill sharks Notorynchus cepedianus, collected in a system of coastal embayments in southeast Tasmania were analyzed to examine sex‐specific differences in the sharks’ coastal space‐use and test novel methods for the analysis of acoustic telemetry data. Sex‐specific space‐use of the broadnose sevengill shark from acoustic telemetry data was analysed in two ways: The recently proposed spatial network analysis of between‐receiver movements was employed to identify sex‐specific space‐use patterns. To include the full breadth of temporal information held in the data, movements between receivers were furthermore considered as transitions between states of a Markov chain, with the resulting transition probability matrix allowing the ranking of the relative importance of different parts of the study area. Both spatial network and Markov chain analysis revealed sex‐specific preferences of different sites within the study area. The identification of priority areas differed for the methods, due to the fact that in contrast to network analysis, our Markov chain approach preserves the chronological sequence of detections and accounts for both residency periods and movements. In addition to adding to our knowledge of the ecology of a globally distributed apex predator, this study presents a promising new step towards condensing the vast amounts of information collected with acoustic tracking technology into straightforward results which are directly applicable to the management and conservation of any species that meet the assumptions of our model.     

Large–Scale Movement and Reef Fidelity of Grey Reef Sharks

Despite an Indo-Pacific wide distribution, the movement patterns of grey reef sharks (Carcharhinus amblyrhynchos) and fidelity to individual reef platforms has gone largely unstudied. Their wide distribution implies that some individuals have dispersed throughout tropical waters of the Indo-Pacific, but data on large-scale movements do not exist. We present data from nine C. amblyrhynchos monitored within the Great Barrier Reef and Coral Sea off the coast of Australia. Shark presence and movements were monitored via an array of acoustic receivers for a period of six months in 2008. During the course of this monitoring few individuals showed fidelity to an individual reef suggesting that current protective areas have limited utility for this species. One individual undertook a large-scale movement (134 km) between the Coral Sea and Great Barrier Reef, providing the first evidence of direct linkage of C. amblyrhynchos populations between these two regions. Results indicate limited reef fidelity and evidence of large-scale movements within northern Australian waters.     

Can you hear me now? Range‐testing a submerged passive acoustic receiver array in a Caribbean coral reef habitat

Submerged passive acoustic technology allows researchers to investigate spatial and temporal movement patterns of many marine and freshwater species. The technology uses receivers to detect and record acoustic transmissions emitted from tags attached to an individual. Acoustic signal strength naturally attenuates over distance, but numerous environmental variables also affect the probability a tag is detected. Knowledge of receiver range is crucial for designing acoustic arrays and analyzing telemetry data. Here, we present a method for testing a relatively large‐scale receiver array in a dynamic Caribbean coastal environment intended for long‐term monitoring of multiple species. The U.S. Geological Survey and several academic institutions in collaboration with resource management at Buck Island Reef National Monument (BIRNM), off the coast of St. Croix, recently deployed a 52 passive acoustic receiver array. We targeted 19 array‐representative receivers for range‐testing by submersing fixed delay interval range‐testing tags at various distance intervals in each cardinal direction from a receiver for a minimum of an hour. Using a generalized linear mixed model (GLMM), we estimated the probability of detection across the array and assessed the effect of water depth, habitat, wind, temperature, and time of day on the probability of detection. The predicted probability of detection across the entire array at 100 m distance from a receiver was 58.2% (95% CI: 44.0–73.0%) and dropped to 26.0% (95% CI: 11.4–39.3%) 200 m from a receiver indicating a somewhat constrained effective detection range. Detection probability varied across habitat classes with the greatest effective detection range occurring in homogenous sand substrate and the smallest in high rugosity reef. Predicted probability of detection across BIRNM highlights potential gaps in coverage using the current array as well as limitations of passive acoustic technology within a complex coral reef environment.     

Fine-scale movement and activity patterns of Caribbean reef sharks (<Emphasis Type="Italic">Carcharhinus perezi</Emphasis>) in the Bahamas

Knowledge of the spatial ecology and movement of animals contributes to our understanding of intra- and inter-specific interactions and ecosystem dynamics, and can inform conservation actions. Here we assessed the space use and activity levels of a marine predator, the Caribbean reef shark (Carcharhinus perezi), in coastal regions of Eleuthera, The Bahamas over a 60-day period using acoustic telemetry. Of the 14 adult sharks (eight males, six females) tagged with acoustic transmitters (equipped with accelerometer sensor), nine were detected in a 14 km² gridded receiver array. Male sharks were significantly less likely to be detected over time relative to females. Given post-release survival is typically high in C. perezi, this finding may indicate that males have larger home ranges and may exhibit lower site fidelity compared to females. Patterns of space use indicated C. perezi primarily occupied the outer reef shelf and were rarely detected on the interior of the reef. Shark activity levels (inferred from acceleration profiles) were highest in close proximity to the reef shelf. Our findings indicate C. perezi individuals frequently occupy deeper water habitats, but make forays into reef shelf habitats where high activity levels are likely related to foraging.     

Conventional tagging and acoustic telemetry of a small surgeonfish, <Emphasis Type="Italic">Zebrasoma flavescens</Emphasis>, in a structurally complex coral reef environment

Passive acoustic telemetry and conventional tag/re-sight techniques were used to study daily movement patterns of adult yellow tang, Zebrasoma flavescens, over a period of months. Range testing and visual observations revealed the limitations of using small acoustic transmitters to monitor movements of small coral reef fish in a topographically complex and noisy coral reef environment. Visual observations of conventionally tagged and albino fish suggest individuals return each day to forage over the same few hundred m² of shallow, turf algae dominated boulder and reef flat habitat for periods of at least weeks to months. Acoustic telemetry data suggest lower frequency of repeated use of daytime foraging, nighttime refuge and sunset spawning sites. However, integration of observation and acoustic telemetry data revealed that many fish were not detected while they were within the empirically tested range of the receivers. These observations indicate that data from passive acoustic telemetry can underestimate the frequency and duration of repeated use of specific areas. Yellow tang adults made daily crepuscular migrations of up to 600 m between foraging and spawning or sheltering sites at consistent times relative to sunset and sunrise. While there was high individual variability in migration distance, almost all individuals moved in the same direction (from south to north) at sunset. This study provided valuable information for evaluating ongoing fishery management efforts using marine protected areas in Hawaii.     

Ecology of the introduced snapper Lutjanus kasmiva (Forsskal) in the reef fish assemblage of a Hawaiian bay

Since its first introduction in 1955, the snapper Lutjanus kasmira (taape) has developed large populations in shallow coastal waters of Hawaii. Visual abundance estimates and trap catches of taape were different between the main habitat types of the reef. Overall, taape was the second most abundant species by numbers and biomass over a hard substratum and was patchily distributed, even within habitat type. The largest individuals occurred singly or in small groups on shallow reef slopes; on the deep slopes and in spur-and-groove habitat, taape occurred in larger groups of smaller sized fish. Catches in fish traps showed a negative relationship with the complexity of the adjacent natural substratum, suggesting that traps are more productive where they offer additional cover in areas with less natural cover. In all areas in which traps were used, from fringing reef to open sand, the catch was dominated by taape. Tagging confirmed the theory that taape tend to maintain a limited range over long periods. Tagging results also indicated occasional long-range movements among major habitats and provided data confirming patterns of movement between reef habitat by day and sand habitat by night. Taape collected over open sand substratum by trap and by hook-and-line were smaller than those collected by the same means at the reef. Observations and collections suggested an ontogenetic trend in habitat use by taape from initial settlement in fringe areas toward ultimate residence on the main reef. Other common fish species in the reef assemblages were tested for statistical associations with taape. Although several positive species associations were found, multivariate analysis did not reveal patterns that indicated strong ecological relationships.     

Lunar and seasonal patterns in fecundity of an indeterminate,multiple‐spawning surgeonfish,the yellow tang Zebrasoma flavescens

Adult bull sharks Carcharhinus leucas were monitored with electronic tags to investigate horizontal and vertical movements in the Atlantic and Pacific Oceans. In both locations, C. leucas showed some fidelity to specific coastal areas with only limited horizontal movements away from the tagging sites after tag attachment. Fish tagged in the Bahamas were detected mostly in the upper 20 m of the water column in water 25-26° C, whereas C. leucas tagged in Fiji spent most of their time below 20 m in water usually >26° C. The results highlight the importance of coastal inshore habitats for this species.     

Mobile receivers: releasing the mooring to ‘see’ where fish go

Much has been learned from the large scale deployment of acoustic tags on aquatic species and associated networks of riverine and marine receivers. While effective in the linear environment of river systems, marine systems limit the ability to provide spatial information on fish movements and distributions due to a combination of costs, logistics, and lack of off-shore technology. At the same time, each year millions of dollars worth of tags are being released into the aquatic environment with extended battery/transmission life, yet detections are limited to coastal arrays. Here we explore new methods of tracking acoustically tagged species in the marine environment. A new miniaturized acoustic receiver, the Vemco Mobile Transceiver (VMT) can be carried by large marine organisms. In combination with satellite and archival tag technology, VMTs were deployed on northern elephant seals to monitor acoustic tags encountered during their migrations across the Northeast Pacific. Early results include acoustic detections of tagged great white sharks, salmon sharks, Chinook salmon, steelhead, lingcod, green sturgeon and other elephant seals. We also propose several alternative directions for future effort: 1) analyzing the growing number of passive acoustic survey recordings made from hydrophone arrays for acoustic tag detections, 2) working with acoustic technology providers to develop hull-mounted receiver systems for the thousands of ocean going vessels around the world and 3) integrating acoustic receiver technology into the thousands of moored and drifting oceanographic buoy arrays.     

Movements,Home Range and Site Fidelity of Snapper (Chrysophrys auratus) within a Temperate Marine Protected Area

Understanding the movement dynamics of marine fish provides valuable information that can assist with species management, particularly regarding protection within marine protected areas (MPAs). We performed an acoustic tagging study implemented within the Port Stephens-Great Lakes Marine Park on the mid-north coast of New South Wales, Australia, to assess the movement patterns, home range and diel activity of snapper (Chrysophrys auratus; Sparidae); a species of significant recreational and commercial fishing importance in Australia. The study focused on C. auratus movements around Cabbage Tree Island, which is predominantly a no-take sanctuary zone (no fishing), with an array of acoustic stations deployed around the island and adjacent reefs and islands. Thirty C. auratus were tagged with internal acoustic tags in November 2010 with their movements recorded until September 2014. Both adult and juvenile C. auratus were observed to display strong site fidelity to Cabbage Tree Island with a mean 12-month residency index of 0.83 (range = 0 low to 1 high). Only three fish were detected on acoustic receivers away from Cabbage Tree Island, with one fish moving a considerable distance of ~ 290 kms over a short time frame (46 days). The longest period of residency recorded at the island was for three fish occurring regularly at the site for a period of 1249 days. Chrysophrys auratus displayed strong diurnal behaviour and detection frequency was significantly higher during the day than at night; however, there was no significant difference in detection frequency between different hours. This study demonstrates that even small-scale protected areas can benefit C. auratus during multiple life-history stages as it maintains a small home range and displays strong site fidelity over a period of 3 years.     

Seasonal occurrence and population structure of the broadnose sevengill shark Notorynchus cepedianus in coastal habitats of south-east Tasmania

Research longline sampling was conducted seasonally from December 2006 to February 2009 to investigate the occurrence and population structure of the broadnose sevengill shark Notorynchus cepedianus in coastal areas of south-east Tasmania. Notorynchus cepedianus showed a consistent temporal trend in seasonal occurrence in Norfolk Bay characterized by high abundances in summer to near absence in winter. This pattern was less pronounced in the Derwent Estuary, where fish were still caught during winter. The absence of smaller total length (L(T) ) classes (<80 cm) from the catches suggests that N. cepedianus are not using these coastal habitats as nursery areas. Of the 457 individuals tagged, 68 (15%) were recaptured. Time at liberty ranged from 6 days to almost 4 years and all but one of the recaptures were caught in its original tagging location, suggesting site fidelity. The large number of N. cepedianus in these coastal systems over summer indicates that these areas are important habitats for this species and that N. cepedianus may have a significant influence on community dynamics through both direct and indirect predator-prey interactions.     

Ascending speed and nocturnal activity of adult masu salmon (Oncorhynchus masou Brevoort) during upstream migration

We released five adult masu salmon (Oncorhynchus masou) tagged with external transmitters to track their ascending behaviour. The signals of all specimens were recorded in the upper area of the river system. Two patterns of ascending behaviour were recognized: ascending upward immediately after release and ascending during increased river discharge. The fastest ascending speed was about 1000 m h^–1. Active movements were detected at night. The signal recording duration at each receiver for each fish was generally brief. Most fish did not stay at the pools where the receivers were installed.     

Evidence of Partial Migration in a Large Coastal Predator: Opportunistic Foraging and Reproduction as Key Drivers?

Understanding animal movement decisions that involve migration is critical for evaluating population connectivity, and thus persistence. Recent work on sharks has shown that often only a portion of the adult population will undertake migrations, while the rest may be resident in an area for long periods. Defining the extent to which adult sharks use specific habitats and their migratory behaviour is essential for assessing their risk of exposure to threats such as fishing and habitat degradation. The present study used acoustic telemetry to examine residency patterns and migratory behaviour of adult bull sharks (Carcharhinus leucas) along the East coast of Australia. Fifty-six VR2W acoustic receivers were used to monitor the movements of 33 bull sharks in the central Great Barrier Reef (GBR). Both males and females were detected year-round, but their abundance and residency peaked between September and December across years (2012–2014). High individual variability in reef use patterns was apparent, with some individuals leaving the array for long periods, whereas others (36%) exhibited medium (0.20–0.40) or high residency (> 0.50). A large portion of the population (51%) undertook migrations of up to 1,400 km to other coral reefs and/or inshore coastal habitats in Queensland and New South Wales. Most of these individuals (76%) were mature females, and the timing of migrations coincided with the austral summer (Dec-Feb). All migrating individuals (except one) returned to the central GBR, highlighting its importance as a potential foraging ground. Our findings suggest that adult bull sharks appear to be highly dependent on coral reef resources and provide evidence of partial migration, where only a portion of the female population undertook seasonal migrations potentially to give birth. Given that estuarine habitats face constant anthropogenic pressures, understanding partial migration and habitat connectivity of large coastal predators should be a priority for their management.     

Fish with Chips: Tracking Reef Fish Movements to Evaluate Size and Connectivity of Caribbean Marine Protected Areas

Coral reefs and associated fish populations have experienced rapid decline in the Caribbean region and marine protected areas (MPAs) have been widely implemented to address this decline. The performance of no-take MPAs (i.e., marine reserves) for protecting and rebuilding fish populations is influenced by the movement of animals within and across their boundaries. Very little is known about Caribbean reef fish movements creating a critical knowledge gap that can impede effective MPA design, performance and evaluation. Using miniature implanted acoustic transmitters and a fixed acoustic receiver array, we address three key questions: How far can reef fish move? Does connectivity exist between adjacent MPAs? Does existing MPA size match the spatial scale of reef fish movements? We show that many reef fishes are capable of traveling far greater distances and in shorter duration than was previously known. Across the Puerto Rican Shelf, more than half of our 163 tagged fish (18 species of 10 families) moved distances greater than 1 km with three fish moving more than 10 km in a single day and a quarter spending time outside of MPAs. We provide direct evidence of ecological connectivity across a network of MPAs, including estimated movements of more than 40 km connecting a nearshore MPA with a shelf-edge spawning aggregation. Most tagged fish showed high fidelity to MPAs, but also spent time outside MPAs, potentially contributing to spillover. Three-quarters of our fish were capable of traveling distances that would take them beyond the protection offered by at least 40–64% of the existing eastern Caribbean MPAs. We recommend that key species movement patterns be used to inform and evaluate MPA functionality and design, particularly size and shape. A re-scaling of our perception of Caribbean reef fish mobility and habitat use is imperative, with important implications for ecology and management effectiveness.     

Reef-Fidelity and Migration of Tiger Sharks,Galeocerdo cuvier,across the Coral Sea

Knowledge of the habitat use and migration patterns of large sharks is important for assessing the effectiveness of large predator Marine Protected Areas (MPAs), vulnerability to fisheries and environmental influences, and management of shark–human interactions. Here we compare movement, reef-fidelity, and ocean migration for tiger sharks, Galeocerdo cuvier, across the Coral Sea, with an emphasis on New Caledonia. Thirty-three tiger sharks (1.54 to 3.9 m total length) were tagged with passive acoustic transmitters and their localised movements monitored on receiver arrays in New Caledonia, the Chesterfield and Lord Howe Islands in the Coral Sea, and the east coast of Queensland, Australia. Satellite tags were also used to determine habitat use and movements among habitats across the Coral Sea. Sub-adults and one male adult tiger shark displayed year-round residency in the Chesterfields with two females tagged in the Chesterfields and detected on the Great Barrier Reef, Australia, after 591 and 842 days respectively. In coastal barrier reefs, tiger sharks were transient at acoustic arrays and each individual demonstrated a unique pattern of occurrence. From 2009 to 2013, fourteen sharks with satellite and acoustic tags undertook wide-ranging movements up to 1114 km across the Coral Sea with eight detected back on acoustic arrays up to 405 days after being tagged. Tiger sharks dove 1136 m and utilised three-dimensional activity spaces averaged at 2360 km³. The Chesterfield Islands appear to be important habitat for sub-adults and adult male tiger sharks. Management strategies need to consider the wide-ranging movements of large (sub-adult and adult) male and female tiger sharks at the individual level, whereas fidelity to specific coastal reefs may be consistent across groups of individuals. Coastal barrier reef MPAs, however, only afford brief protection for large tiger sharks, therefore determining the importance of other oceanic Coral Sea reefs should be a priority for future research.     

Archival and acoustic tags reveal the post-spawning migrations, diving behavior, and thermal habitat of hatchery-origin Sacramento River steelhead kelts (Oncorhynchus mykiss)

Acoustic and geolocating archival tags were deployed on steelhead kelts to study their post-spawning migrations, diving behavior and thermal habitat. Fourteen reconditioned steelhead kelts were surgically implanted with LAT2510 archival and V7-2 L acoustic tags, and released from Coleman National Fish Hatchery. An array of acoustic receivers in the Sacramento River and Delta, and the San Francisco Bay estuary detected the downstream movement of the steelhead. Two steelhead kelts (J and M) with archival tags were recovered at the hatchery after 219 and 285 days at liberty respectively. Based on changes in geolocations, vertical movements, and water temperatures, the migrations of steelhead J and M were divided into five and six phases respectively. Steelhead J moved into coastal California waters while steelhead M remained in freshwater for the majority of its time at liberty. Large increases in temperature and opacity were recorded before and after the ocean phases, likely as the steelhead moved through the Sacramento Delta. Both steelhead kelts remained relatively close to the surface throughout their migration but there were diurnal differences in the vertical movements. In freshwater, the steelhead tended to be deeper during the day (e.g., steelhead J: 3.08?±?1.50 vs 1.65?±?1.15 m, day vs night) but in the ocean, the steelhead were typically deeper during the night (1.32?±?1.61 vs 5.63?±?6.11 m). Both steelhead kelts appeared to be less oceanic than a previous study in Scott Creek, a small coastal stream approximately 100 km south of the mouth of San Francisco Bay. Although this study has a small sample size, the results suggest that steelhead kelts from a large river and estuary system, like the Sacramento River, may be relatively less oceanic than steelhead kelts from small coastal streams.     

An electromagnetic tracking system for studying the movements of rock (spiny) lobsters

A simple and inexpensive method for examining the movements of rock (spiny) lobsters is described. Electromagnetic tags emitting a pulsed signal were detected in loop antennae placed on the bottom in a grid pattern within a shallow coastal reef area. Signals were received in a tuned receiver and then interpreted using an oscilloscope. Results showing examples of the nocturnal movements of Panuliruscygnus George the western rock lobster in Western Australia are presented.