When Giants Turn Up: Sighting Trends,Environmental Influences and Habitat Use of the Manta Ray Manta alfredi at a Coral Reef

Manta rays Manta alfredi are present all year round at Lady Elliot Island (LEI) in the southern Great Barrier Reef, Australia, with peaks in abundance during autumn and winter. Drivers influencing these fluctuations in abundance of M. alfredi at the site remain uncertain. Based on daily count, behavioural, weather and oceanographic data collected over a three-year period, this study examined the link between the relative number of sightings of manta rays at LEI, the biophysical environment, and the habitat use of individuals around the LEI reef using generalised additive models. The response variable in each of the three generalised additive models was number of sightings (per trip at sea) of cruising, cleaning or foraging M. alfredi. We used a set of eleven temporal, meteorological, biological, oceanographic and lunar predictor variables. Results for cruising, cleaning and foraging M. alfredi explained 27.5%, 32.8% and 36.3% of the deviance observed in the respective models and highlighted five predictors (year, day of year, wind speed, chlorophyll-a concentration and fraction of moon illuminated) as common influences to the three models. There were more manta rays at LEI in autumn and winter, slower wind speeds, higher productivity, and around the new and full moon. The winter peak in sightings of foraging M. alfredi was found to precede peaks in cleaning and cruising activity around the LEI reef, which suggests that enhanced food availability may be a principal driver for this seasonal aggregation. A spatial analysis of behavioural observations highlighted several sites around the LEI reef as ‘multi-purpose’ areas where cleaning and foraging activities commonly occur, while the southern end of the reef is primarily a foraging area. The use of extensive citizen science datasets, such as those collected by dive operators in this study, is encouraged as they can provide valuable insights into a species'' ecology.     

Genome-wide SNPs detect no evidence of genetic population structure for reef manta rays (Mobula alfredi) in southern Mozambique

Little is known about the extent of genetic connectivity along continuous coastlines in manta rays, or whether site visitation is influenced by relatedness. Such information is pertinent to defining population boundaries and understanding localized dispersal patterns and behaviour. Here, we use 3057 genome-wide single-nucleotide polymorphisms (SNPs) to evaluate population genetic structure and assess the levels of relatedness at aggregation sites of reef manta rays (Mobula alfredi) in southern Mozambique (n = 114). Contrary to indications of limited dispersal along the southern Mozambican coastline inferred from photo-identification and telemetry studies, our results show no evidence of population structure (non-significant F_ST < 0.001) for M. alfredi along this coast. We also found no evidence that individuals sampled at the same site were more related than expected by chance for males, females or across both sexes, suggesting that kinship may not influence visitation patterns at these sites. We estimated the effective population size (N_e) of this population to be 375 (95% CI = 369–380). Comparison to a distant eastern Indian Ocean site (Western Australia, n = 15) revealed strong genetic differentiation between Mozambique and Western Australia (F_ST = 0.377), identifying the Indian Ocean basin as a barrier to dispersal. Our findings show that genetic connectivity in M. alfredi extends for several hundred kilometres along continuous coastlines. We therefore recommend that the population in Mozambique be considered a discrete management unit, and future conservation plans should prioritize integrated strategies along the entire southern coastline.Subject terms: Conservation genomics, Structural variation     

The geographic distribution of reef and oceanic manta rays (Mobula alfredi and Mobula birostris) in Australian coastal waters

The known distribution of manta rays in Australian waters is patchy, with records primarily centred around tourism hotspots. We collated 11,614 records of Mobula alfredi from photo-ID databases (n = 10,715), aerial surveys (n = 378) and online reports (n = 521). The study confirms an uninterrupted coastal distribution from north of 26°S and 31°S on the west and east coasts, respectively. More southerly M. alfredi records relate to warm-water events with a southernmost extent at 34°S. Coastal sightings of Mobula birostris were rare (n = 32), likely reflecting a preference for offshore waters, but encompass a wider latitudinal extent than M. alfredi of 10–40°S.     

Fine‐scale oceanographic drivers of reef manta ray (Mobula alfredi) visitation patterns at a feeding aggregation site

Globally, reef manta rays (Mobula alfredi) are in decline and are particularly vulnerable to exploitation and disturbance at aggregation sites. Here, passive acoustic telemetry and a suite of advanced oceanographic technologies were used for the first time to investigate the fine‐scale (5‐min) influence of oceanographic drivers on the visitation patterns of 19 tagged M. alfredi to a feeding aggregation site at Egmont Atoll in the Chagos Archipelago. Boosted regression trees indicate that tag detection probability increased with the intrusion of cold‐water bores propagating up the atoll slope through the narrow lagoon inlet during flood tide, potentially transporting zooplankton from the thermocline. Tag detection probability also increased with warmer near‐surface temperature close to low tide, with near‐surface currents flowing offshore, and with high levels of backscatter (a proxy of zooplankton biomass). These combinations of processes support the proposition that zooplankton carried from the thermocline into the lagoon during the flood may be pumped back out through the narrow inlet during an ebb tide. These conditions provide temporally limited feeding opportunities for M. alfredi, which are tied on the tides. Results also provide some evidence of the presence of Langmuir Circulation, which transports and concentrates zooplankton, and may partly explain why M. alfredi occasionally remained at the feeding location for longer than that two hours. Identification of these correlations provides unique insight into the dynamic synthesis of fine‐scale oceanographic processes which are likely to influence the foraging ecology of M. alfredi at Egmont Atoll, and elsewhere throughout their range.     

DISTRIBUTION PATTERNS OF INDIVIDUALLY IDENTIFIABLE WEST INDIAN MANATEES (TRICHECHUS MANATUS) IN FLORIDA

Photographs of distinctively scarred manatees ( Trichechus manatus ) were taken at aggregation sites throughout Florida and assembled into a catalog for identifying individuals. Resightings of known manatees in different years or at different locations enabled us to document site fidelity and long-distance movements. Of the 891 individuals included in the catalog as of August 1986, 470 (53%) were resighted at least once. We documented 219 cases of seasonal returns to specific aggregation sites and 98 instances of movements between areas. Movements in excess of 820 km were documented for individuals on the east coast of Florida. Rapid movements south during early winter, as well as northerly spring movements, verify a seasonal migration pattern for many individuals along Florida's east coast. The wide-ranging migratory habits of manatees in Florida must be considered if meaningful management strategies are to be developed and implemented.     

Reproductive ecology of the reef manta ray Manta alfredi in southern Mozambique

The application of a photographic identification methodology using the unique ventral surface markings (natural spot patterns) of an observed population in southern Mozambique enabled many aspects of the reproductive ecology of reef manta rays Manta alfredi to be examined. The region encompassing the study site was identified as a mating ground for M. alfredi based on observations of mating events and fresh mating scars on females. The distribution of these pectoral fin scars was highly biased and indicated a strong lateralized behavioural trait, with 99% of these scars occurring only on the left pectoral fin. No other elasmobranch has been reported to display behavioural lateralization. The study region also acts as a birthing ground, with individuals typically giving birth in the austral summer period after a gestation of c. 1 year. Reproductive periodicity in M. alfredi was most commonly biennial, but a few individuals were pregnant in consecutive years, confirming an annual ovulatory cycle. The production of a single pup appears to be the normal situation, although observations in the wild as well as during opportunistic dissections of individuals killed by fisheries revealed that two pups are conceived on occasion. Many aspects of the study have contributed to the limited baseline data currently available for this species and have highlighted the potential need for more conservative conservation strategies.     

Occurrence of Indo-Pacific bottlenose dolphins (Tursiops aduncus) off the Wild Coast of South Africa using photographic identification

The present study represents the first reported boat-based photographic identification study of Indo-Pacific bottlenose dolphins (Tursiops aduncus) off the Wild Coast of southeast South Africa. This area is known for the annual sardine run, which attracts apex predators to the region during the austral winter. Dedicated photo-identification surveys were conducted along this coast at three different study sites in February, June, and November of each year from 2014 to 2016. During 47 surveys, 136 bottlenose dolphin groups were encountered, an estimated 4,474 dolphins observed, and 2,149 individuals were identified. Although most individuals (N = 1,770, 82.4%) were only observed once, some were resighted 2–7 times (N = 379, 17.6%), with an average of 305 days (range: 88–705 days) between resightings. The majority of bottlenose dolphins were resighted within the same study site (N = 192), indicating some degree of residency. However, 65 individuals were observed at two different study sites, indicating individual movements along the coast. Our findings contrast earlier suggestions that bottlenose dolphins only use the Wild Coast during the sardine run, as we found large number of animals year-round with some level of site fidelity. This highlights the importance of the Wild Coast to bottlenose dolphins and provides further information on their status off southeastern South Africa.     

Trends in sightings and population structure of white sharks, <Emphasis Type="Italic">Carcharodon carcharias,</Emphasis> at Seal Island,False Bay,South Africa,and the emigration of subadult female sharks approaching maturity

A long-term sightings and photographic identification (photo-ID) database documented the inter-annual and monthly trends in white shark (Carcharodon carcharias) sightings, sex ratios and life-history stages at Seal Island, False Bay, South Africa, over the period 2004–2012. A total of 1105 sightings were recorded during 171 scientific surveys incorporating 577 h of observation (annual mean 64.10 h, range 23.71–178.69 h). The mean annual sighting rate was 1.87 (range 0.90–3.19) sharks per hour and sighting rates declined over the nine-year study period. It is unknown whether the decline resulted from an actual population decline, or was due to changes in shark distribution or environmental conditions, but it highlights the need for ongoing monitoring at this site. The overall sex ratio was 1.0: 1.0: 2.3 for male: female: unsexed sharks, respectively. Of the 1105 sightings, 39% (n = 433) were photo-ID’d, representing 303 individual sharks (112: 111: 80, male: female: unsexed) and 130 resightings. Of the 303 photo-ID’d sharks, 71% were sighted in a single year only, indicative of transient behaviour. Of the 29% of sharks that were resighted in more than one year, 65% were resighted in the following year, indicating site fidelity to Seal Island, whereas 35% skipped one or more years between encounters, indicative of a temporary absence. The majority (60%) of the photo-ID’d sharks were immature, 32% were subadult and only 8% were mature. No young-of-the-year sharks and few adults were recorded, indicating that Seal Island is not an adult aggregation site, nor a pupping or nursery area, but rather is best described as a seasonal feeding ground. Large females were rarely resighted again after they approached length-at-maturity (≥450 cm TL), whereas some adult males were recorded consistently across years. The emigration of subadult female sharks approaching maturity from Seal Island, combined with the small number of mature sharks of both sexes reported from any South African location, indicate that adult aggregation sites, and thus areas of reproductive importance, still remain unknown.     

Notes on the feeding behaviour and swimming pattern of Manta alfredi (Chondrichthyes,Mobulidae) in the Red Sea

The feeding behaviour and the swimming pattern of the reef manta (Manta alfredi) are described based on video recordings conducted in Sudan, Red Sea, Indian Ocean, during June 2001. Three circular swimming patterns are described: horizontal circle pattern, inclined circle pattern and double circle pattern, related to feedings on zooplankton distributed in the water column. The observation of circle-like swimming patterns for reef manta rays in shallow waters expands our knowledge about M. alfredi and suggests that the distribution and abundance of zooplankton influence the occurrence and group size of this species in the studied area, as reported from other localities for Manta species.     

Site affinity of whitespotted eagle rays Aetobatus narinari assessed using photographic identification

Photographic identification was used to track the movements of the whitespotted eagle ray Aetobatus narinari around South Caicos, Turks and Caicos Islands. A total of 165 individuals were identified, aided by the computer program I³S Spot. The sex ratio across all study sites in 2015 was not significantly different from 1:1 (χ² = 2·8, P > 0·05). 33·9% of all individual rays were resighted at least once and the maximum number of days between the first and last sighting was 1640 (median 165, interquartile range, IQR = 698). Sightings of individuals occurred at locations differing from the original sighting location 24·6% of the time (0·7–20 km away). The entire population around South Caicos has yet to be sampled and these rays exhibited site affinity during the study period; they are either resident to South Caicos or are using the area for parts of the year before making movements elsewhere and then returning. Given these results, A. narinari is suited to local‐scale management and conservation efforts.     

Integrating multiple data sources to assess the distribution and abundance of bottlenose dolphins Tursiops truncatus in Scottish waters

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Occurrence of false killer whales (Pseudorca crassidens) and their association with common bottlenose dolphins (Tursiops truncatus) off northeastern New Zealand

On a global scale, false killer whales (Pseudorca crassidens) remain one of the lesser‐known delphinids. The occurrence, site fidelity, association patterns, and presence/absence of foraging in waters off northeastern New Zealand are examined from records collected between 1995 and 2012. The species was rarely encountered; however, of the 61 distinctive, photo‐identified individuals, 88.5% were resighted, with resightings up to 7 yr after initial identification, and movements as far as 650 km documented. Group sizes ranged from 20 to ca. 150. Results indicate that all individuals are linked in a single social network. Most observations were recorded in shallow (<100 m) nearshore waters. Occurrence in these continental shelf waters is likely seasonal, coinciding with the shoreward flooding of a warm current. During 91.5% of encounters, close interspecific associations with common bottlenose dolphins (Tursiops truncatus) were observed. Photo‐identification reveals repeat inter‐ and intraspecific associations among individuals with 34.2% of common bottlenose dolphins resighted together with false killer whales over 1,832 d. While foraging was observed during 39.5% of mixed‐species encounters, results suggest that social and antipredatory factors may also play a role in the formation of these mixed‐species groups.     

Courtship and mating behaviour of manta rays Mobula alfredi and M. birostris in the Maldives

The aim of this 14 year study was to elucidate the entire courtship and mating behaviour of manta rays Mobula alfredi and M. birostris using behavioural observations, video and photographic records. From 2003 to 2016, over 11,000 surveys were undertaken at known manta ray aggregation sites in the Maldives to record any observed manta rays reproductive activity. From 47,591 photo‐ID sightings, 4,247 individual M. alfredi were identified and 226 individual M. birostris from 229 photo‐ID sightings, all recorded at 22 atolls across 265 different sites. Courtship activity was observed on 206 surveys at 30 different sites. A total of 229 courtship events were recorded, with 90% (n = 205) of them occurring at cleaning sites. The observed courtship activity was categorised into seven distinct stages which are described in detail: initiation, endurance, evasion, pre‐copulatory positioning, copulation, post‐copulatory holding and separation. Photographs provide the first scientific record of the entirety of manta rays courtship and mating. Both M. alfredi and M. birostris appear to engage in the same elaborate courtship rituals, exhibiting the same behaviours during all stages of the courtship and mating process.     

Batesian mimicry in the nonrewarding saprophytic orchid Danxiaorchis yangii

Batesian mimicry, a type of deceptive pollination, is a complicated strategy used by nonrewarding plants to attract pollinators, but some hypotheses concerning this have not been systematically verified. In order to show in detail a case of Batesian mimicry on saprophytic orchid Danxiaorchis yangii, the ecological relationship between Danxiaorchis yangii, Lysimachia alfredi and Dufourea spp. was explored. Lysimachia alfredi could provide a reward to Dufourea sp., whereas Danxiaorchis yangii not. The floral morphology and geographical distribution of these two plants were highly overlapping, and the fruit set rate of Danxiaorchis yangii was significantly positively correlated with the number of nearby L. alfredi individuals. In a glass cylinder experiment, Danxiaorchis yangii and L. alfredi attracted Dufourea spp. through visual signals, but the insect could not distinguish between flowers of the two plants before landing on flowers. The ultraviolet reflection spectra of flowers between the two plant species were highly similar. In the hexagonal color models constructed according to the visual characteristics of bees, the flower color signals of these two plant species highly overlap, indicating that the visual signals of the flowers of the two plants to the pollinator were greatly similar. All of these results provided evidence that Danxiaorchis yangii simulated the visual signals of L. alfredi through Batesian mimicry, thereby deceptively attracting Dufourea spp.     

Long-term variation in the winter resident bird community of Guánica Forest, Puerto Rico: lessons for measuring and monitoring species richness

ABSTRACT.   Because the winter season is potentially limiting for migratory birds, understanding their nonbreeding distributional patterns is essential. At a given site, patterns of species occurrence and abundance may vary over time and, within a species, wintering strategies may vary with regard to the degree that individuals are site-faithful both within and between winters. We examined long-term patterns in the composition of a winter resident bird community to determine how long a site must be studied to understand the wintering community. Over a 34-yr period of constant-effort mist netting at a site in Guánica, Puerto Rico, we captured 21 species of winter resident birds, with mean total captures varying from 8.3 to 18.9 individuals per net line and 6–14 species captured per year. Species richness capture/recapture models generated numbers similar to actual capture rates. Capture and recapture data allowed us to categorize winter residents into three groups: sporadic winter residents (14 species), regular winter residents (four species captured nearly every year), and dominant winter residents (three species captured each year with high rates of recapture). Our results suggest that sampling for at least three consecutive winters is needed to accurately characterize the bird community at a site. However, sampling for 5 yr is better, and 10-yr samples generate patterns similar to those based on our entire 34-yr sample. A 1-yr sample provides minimal information about the composition and characteristics of a winter resident bird community.     

Homing and Daytime Tidal Movements of Juvenile Snappers (Lutjanidae) between Shallow-Water Nursery Habitats in Zanzibar, Western Indian Ocean

We studied daily tidal movements of tagged juvenile Lutjanus fulviflamma and Lutjanus ehrenbergii between two adjacent habitats, a subtidal channel and shallow tidal notches in the fossil reef terrace, in a shallow marine bay on Zanzibar Island (Tanzania). Due to a large tidal range, the notches were dry at low-tide and were only accessible to the snappers at high-tide. Of the resighted individuals, 48% showed clear movement between the two habitats, orientated in a direction perpendicular to the tidal currents. Individuals resighted more than once showed site fidelity, indicating homing in both the channel and the notches. We suggest that a significant part of this population of juvenile snappers may move from a low-tide resting habitat to a high-tide resting habitat during the daytime, perhaps to avoid predation by larger predators that may enter the channel at high-tide.     

Population estimates of Alfred mantas (<Emphasis Type="Italic">Manta alfredi</Emphasis>) in central Maldives atolls: North Male,Ari and Baa

The Alfred manta, Manta alfredi, is economically important in the Maldives due to the large number of tourists attracted to SCUBA dive and snorkel with them. It is important to understand the size of populations around main diving points and throughout the Maldives so that manta based tourism activities can be developed more successfully and the impact of tourism can be monitored. Using natural markings to identify individual manta rays, Petersen and Jolly-Seber probabilistic estimators were used to assess population size for main dive sites and the central atolls of North Male, Ari and Baa. Between 2000 and 2009, 1835 individual mantas were identified from 3373 sightings. There was a significant gender bias with a male: female ratio of 1:1.8. Only 24.7% of manta rays were re-sighted between 1 and 34 times with a maximum gap of 7 years and 11 months between sightings. Estimates for populations around main dive sites were 181–562. The estimated population of M. alfredi for North Male atoll is 537 (SE ± 53.2). Estimated atoll populations were extrapolated to obtain an estimate of Manta alfredi population for the entire Maldives of 9,677 individuals.     

The annual gametogenic cycle of the sea anemone Metridium senile from the Gulf of Maine

A year-long study was conducted to quantify the reproductive cycle of the clonal sea anemone, Metridium senile, an important member of the benthic community in the Gulf of Maine (GOM). M.senile is an abundant sessile invertebrate that forms identifiable clonal aggregations that are maintained by pedal laceration but individuals can also reproduce sexually. Specimens were collected monthly from a shallow (< 10 m) subtidal site and the reproductive cycle of male and female anemones was described on a seasonal basis using histological preparation of the gonads and light microscopy. Gametogenesis was determined for both sexes and showed that females undergo spawning during the seasonal transition from summer to fall, while males have mature sperm within testicular cysts in the winter, spring, and summer. Environmental data recorded at the study site shows that the spawning period occurs during a period of peak water temperature (August-September), and rapid gametogenesis occurs during periods of high food availability (April). The low percentage of reproductively mature anemones and the dominance of asexual reproduction in this population are believed to be the result of the high flow regime at this site.     

Running the Gauntlet: Regional Movement Patterns of Manta alfredi through a Complex of Parks and Fisheries

Manta rays (Genus Manta) are economically important for fisheries and tourism in Indonesia. These species have been listed by the International Union for the Conservation of Nature Red List as Vulnerable to extinction; therefore, human exploitation of manta rays must be regulated. A better understanding of the habitat use and movement patterns of manta rays in Indonesia is needed in order to employ effective conservation measures. To gain better insight into the movements of Manta alfredi we used ‘Manta Matcher’, an online database with an integrated automated matching algorithm, to compare photographs from 2,604 encounters of M. alfredi collected by recreational divers and dive operators throughout Indonesia over a nine-year period. This photographic comparison revealed that manta rays migrated between regional sanctuaries such as Nusa Penida, the Gili Islands, and the Komodo National Park (up to 450 km straight-line distance). The areas between these sanctuaries are heavily fished and trafficked by ships, and when manta rays travel through these regions they risk being fished and injured by ship strikes. These long-range manta ray movements suggest connectivity between M. alfredi populations in neighboring islands and raise concerns about the future management of regional populations. It is recommended that a national conservation strategy be developed to protect the remaining populations in the country.     

Distribution of the reef manta ray Mobula alfredi and the oceanic manta ray Mobula birostris in the Philippines: a collaborative effort for conservation

Little is known about manta ray population size, structure and connectivity in the Philippines. In collaboration with dive operators, non-governmental organizations and authorities, sightings of manta rays were collated into a single national database. Using in-water photographs and videos gathered through citizen science and dedicated research efforts, this study compiled sightings between 2004 and 2020, showing 22 separate sites throughout the archipelago with manta rays present. A total of 392 individual reef manta rays (Mobula alfredi) and 107 oceanic manta rays (Mobula birostris) were identified from the collected footage. Four specific sites in the provinces of Masbate and Palawan together hosted 89% of all identified individuals and accounted for 95% of sightings, highlighting these areas are key aggregation sites. This study also reports the movements of M. birostris within the Philippines, based on photo-identification of three individuals moving 150 km between Cebu and Masbate. Despite the growing number of recreational divers in Daanbantayan and San Jacinto, an 80% decline in M. birostris sightings was observed at these sites. To ensure effective future conservation, it is recommended that efforts focus on the identification and protection of manta ray hotspots and migratory corridors, the creation of a sustainable tourism framework and, most important, the implementation of mitigation strategies to reduce fisheries interactions.