Atlantic humpback dolphins Sousa teuszii in the Saloum Delta (Senegal): distribution,relative abundance and photo-identification

During October and November 2015, the first systematic survey of Sousa teuszii was carried out in the Saloum Delta (Senegal, West Africa), comprising 1 617.5 km of boat-based survey coverage. Thirty sightings were recorded in the Saloum and Diomboss rivers, and along the southern coastline. Dolphins were also observed entering the Bandiala and Djinack channels, and travelling across the border into Gambia. The initial sighting locations were 0.043–1.192 km from shore, and tracked dolphins did not move more than 2.082 km from shore. Groups comprised 1–29 animals (mean 9.3 animals), and at least three neonate calves were observed during November. The overall relative abundance was 0.018 sight. km^?1 and 0.175 ind. km^?1. Sightings were concentrated in the Diomboss where relative abundance reached 0.037 sight. km^?1 and 0.331 ind. km^?1. Non-intensive photo-identification produced a minimum population size of 103 animals, the highest recorded for S. teuszii anywhere in its range. Photo-identification also confirmed a movement of individuals between different parts of the Saloum Delta. Combined travel–forage dominated the behaviour. Dolphins were photographed capturing mullet (Mugil sp.) on three occasions. The distribution, population size and movements of S. teuszii are discussed in relation to management.     

Photo-identification and habitat use of Atlantic humpback dolphins Sousa teuszii around the Río Nuñez Estuary in Guinea,West Africa

Ecological data for the Atlantic humpback dolphin Sousa teuszii are scant. Six on-effort Sousa teuszii sightings were recorded during 817.6?km of boat-based effort in the Río Nuñez region of Guinea during October and November 2013. Two incidental sightings were also reported. Groups comprised 1–25 animals. Photo-identification produced a minimum population estimate of 47 animals. Most sightings (n = 5) were located close (<1?km) to shore along a 5.7?km stretch of coast on the west side of Île de Taïdi, primarily over shallow, sand–mud habitat. Two very distinctive individuals were present in all four Taïdi photo-identification encounters, suggesting high site fidelity and stable associations. Two sightings occurred in the outer Río Nuñez Estuary much farther from the coast (5–12?km) but in relatively shallow water (≤15?m) over sand–mud sediment. Focal follows (n = 5: 0.2–3.8?h duration) produced 9.02?h of behavioural data. Travel (51%), foraging (39%) and feeding (9.2%) dominated, with Taïdi dolphins spending more time foraging and feeding than the outer estuary groups. Three individuals had linear-severed dorsal fins consistent with injuries from fishing line. Some management implications of variation in habitat, site fidelity and movements of Sousa teuszii groups are discussed.     

Multi-stakeholder collaboration yields valuable data for cetacean conservation in Gamba,Gabon

Private industry, the Government of Gabon and two international NGOs collaborated to conduct marine surveys off the coast of Gabon, Central Africa. Surveys addressed multiple objectives of surveillance and monitoring, the documentation of the distribution of and threats to the marine megafauna, and capacity-building among government agents and local early-career scientists. During 22 days of survey effort over a two-year period, observers documented humpback whales Megaptera novaeangliae, bottlenose dolphins Tursiops truncatus, Atlantic humpback dolphins Sousa teuszii and common dolphins Delphinus delphis. Humpback whale presence was limited to the months of July to November. Bottlenose dolphins were present year-round and photo-identification of individuals indicated a closed, resident population, with an abundance estimate of 118 (CV = 21.6%, 95% CI 78–180). Small open-decked fishing vessels with gillnets were observed concentrated around river mouths within 2 km of shore, while commercial trawlers were at least 10 km offshore; all were confirmed to be registered and legal. Observations of marine turtles, flocks of marine birds, and floating logs and other debris were sparse. This multi-stakeholder collaboration to conduct a marine survey can serve as an effective model by which funding and logistic support from private industry paired with technical expertise from NGOs and academic institutions can benefit marine and coastal conservation.     

Abundance and degree of residency of humpback dolphins Sousa plumbea in Mossel Bay,South Africa

Indian Ocean humpback dolphins Sousa plumbea inhabit nearshore waters from South Africa to eastern India. Humpback dolphins are vulnerable to conservation threats due to their naturally small population sizes and use of nearshore habitats, where human activities are highest. We investigated the abundance and residency of this species inhabiting Mossel Bay, South Africa, using photographic mark-recapture. Data were collected during 81 surveys in Mossel Bay between 2011 and 2013. Open population modelling using the POPAN parameterisation produced a ‘super-population’ estimate of 125 individuals (95% CI: 61–260) and within-year estimates of between 33 and 86 individuals (2011: 71 [95% CI: 30–168]; 2012: 33 [15–73], 32 [15–70]; 2013: 46 [20–108]). Although less appropriate, closed capture models were also run for comparison with previous studies in the region and generated similar, but slightly smaller, population estimates within each year. We compared our catalogue with opportunistic data collected from East London, Plettenberg Bay, De Hoop and Gansbaai. The only catalogue matches attained were between Plettenberg Bay (n = 44 identified) and Mossel Bay (n = 67 identified), separated by 140?km. Population exchange was moderate, with nine individuals resighted in multiple years between these two areas. This study supports previous findings of long-range movements for this species and provides a baseline from which to assess future impacts on the population.     

Spawning patterns of four species of predominantly temperate pelagic fishes in the sub-tropical waters of southern Queensland

The diverse pelagic fish assemblage of sub-tropical southern Queensland includes fishes with predominantly temperate distributions, such as tailor Pomatomus saltatrix, sardine Sardinops sagax, round herring Etrumeus teres, and Australian anchovy Engraulis australis. The peak spawning seasons of P. saltatrix, S. sagax and E. teres occur during late winter and early spring (June–October). Eggs and larvae of these three species are widely distributed in shelf waters and comprise >50% of the ichthyoplankton assemblage during this period. Mean monthly sea surface temperatures (SSTs) during late winter and early spring range from 21 to 23 °C, and are thus similar to those recorded in southern Australia during summer and autumn, which is the spawning season of these three species in those temperate waters. E. australis eggs occur mainly in inshore waters, and comprise >50% of fish eggs collected during summer and autumn when mean monthly SSTs in southern Queensland exceed 27 °C. E. australis also spawns mainly during summer and autumn in temperate Australia. Hence, water temperature may be less important as a determinant of the spawning season of E. australis than it is for the other three species. The suitability of southern Queensland for spawning by predominantly temperate species during late winter and early spring may contribute to the high diversity of the region's pelagic fish assemblage. Adult P. saltatrix, S. sagax and E. teres appear to migrate northwards into southern Queensland during early winter to spawn, and larvae may be transported southwards into temperate waters by the East Australian Current. This dispersal-migration pattern is similar to those observed for several species, including P. saltatrix, in the western boundary current systems off the east coasts of North America and Africa. Hence, pelagic fishes in ecosystems off the east coast of three continents migrate into sub-tropical waters to spawn, and larvae are transported back into temperate nursery areas by the prevailing current.     

Establishment and closure of the directed Florida whiting (Menticirrhus americanus) fishery and implications for North Atlantic right whale (Eubalaena glacialis) conservation and management

Today, ocean and coastal marine resource management is extremely complex. Marine resource managers are charged with conserving and managing many diverse species. Southern kingfish (Menticirrhus americanus), commonly known as whiting, are found from southern New England to Florida. During the fall through winter, western North Atlantic right whales (Eubalaena glacialis) are primarily found in the coastal nearshore waters off South Carolina, Georgia, and Florida, overlapping the whiting's range and habitat.     

Cetacean research in the southern African subregion: a review of previous studies and current knowledge

Cetacean research, in terms of the number of papers, and areas for which data are available, has expanded considerably in the southern African subregion in the past decade, especially in the South-West Indian Ocean. We review cetacean research within this subregion from the 1800s to the present to provide an overview of findings, investigate trends and identify knowledge gaps. Data are presented separately for large whales (those subject to commercial whaling) and smaller cetaceans, and are separated by era and ocean basin. Over 550 peer-reviewed papers and books were identified relating to research on cetaceans within the subregion. More than half (284) have been produced since 1990 and 193 relate specifically to South African waters. The most-studied species are those that are most accessible due to their coastal distributions (southern right whale Eubalaena australis: 45 papers, humpback whale Megaptera novaeangliae: 31 papers, killer whales Orcinus orca: 27 papers, Indo-Pacific bottlenose dolphin Tursiops aduncus: 30 papers, Indo-Pacific humpback dolphin Sousa chinensis (plumbea form): 25 papers) and/or were hunted commercially (sperm whale Physeter macrocephalus: 25 papers). Identified conservation concerns vary throughout the subregion, but include bycatch and directed hunts, oil and gas development, ecotourism activities, shifts in prey resources, and noise and chemical pollution. The inshore stocks of Bryde's whales Balaenoptera edeni, the Indo-Pacific humpback dolphin and the Atlantic humpback dolphin S. teuszii were identified as the populations of highest conservation concern, although there are considerable knowledge gaps relating to deep-water species and almost no data (even on species occurrence) are available for several areas and countries.     

Spawning strategies and transport of early stages of the two Cape hake species,Merluccius paradoxus and M. capensis,in the southern Benguela upwelling system

Seasonal and short-term variability of environmental parameters influence the spawning strategies of fish species. In this study, the spawning strategies and the transport of early stages of the two Cape hake species off South Africa were investigated. Distribution of eggs and larvae of Merluccius paradoxus and M. capensis was analysed in order to derive more detailed and species-specific information on spawning season, spawning location, and transport of early stages. Samples were collected during three pilot surveys between January and October 2007 and during an extensive survey in September/October 2008 in the southern Benguela upwelling system off South Africa. Eggs and larvae of M. paradoxus were found in greater numbers than those of M. capensis during all surveys. Highest abundances were found from September to October, indicating one spawning peak for M. paradoxus during late austral winter to spring. The western Agulhas Bank was identified as the primary spawning ground, and smaller spawning events occurred on the West Coast. Larvae of both species were mainly distributed in subsurface waters between 25 and 100 m. More than 50% of all larvae caught had a total length between 3 and 4 mm and size increased significantly with decreasing latitude. Merluccius capensis were found closer inshore than M. paradoxus, indicating that early stages of the two species followed separate drift routes. We assume that this distribution pattern most likely evolved from differences in spawning location and phenology. The spawning strategies of M. paradoxus and M. capensis are well adapted to a time-frame of optimal transport conditions favourable for larval survival in the highly variable environment of the southern Benguela upwelling system, but the peak spawning of the two species is separated in time and space.     

Dugong abundance and distribution in the Bazaruto Archipelago,Mozambique

Despite the distribution of dugongs Dugong dugon ranging across nearshore waters of the tropical and subtropical regions of the Indian Ocean and western Pacific Ocean, their distribution in the western Indian Ocean is highly fragmented and appears to be declining. The population of the Bazaruto Archipelago is believed to comprise the only viable population in the region. In all, 27 surveys were flown over the Bazaruto Bay area to define the distribution and estimate the abundance of the species in the area. A total of 9 052 nautical miles of survey effort was flown during the surveys, from which there were 355 sightings of 760 dugongs. Two core areas of distribution were apparent within the surveyed area; a northern core area spread within the 10 m isobath between the Save River mouth and Ponta Bartolomeu Dias (21°24′S), and a southern core area aligned with the shallow sandbanks to the north and south of Santa Carolina Island. Group sizes recorded in the Bazaruto Archipelago were comparable to group sizes recorded in other regions where dugongs occur, although few large (>20) groups of dugongs were seen in this study. Line transect analyses of each survey showed dugong densities were considerably lower than densities recorded in surveys in Australian waters or in the Arabian Gulf, with a population estimate of 247 dugongs (CV = 34.1) when all surveys were considered, and 359 dugongs (CV = 38.2) when only the surveys that were carried out under adequate sighting conditions were included.     

Video observations on the habitat association of demersal nekton in the midshelf benthic environment off the Orange River,Namibia

A semi-quantitative assessment is made of the animals observed in archived videotapes taken from the research submersible Jago, during diamond mining and exploratory surveys off the mouth of the Orange River on the west coast of southern Africa (28°15′S, 29°11′S) in November 1996. The seabed environment is described and nekton associations with substratum features are identified. The area is characterized by heterogeneity to its physical and biological struture. The variety of observed nekton is low, and communities are dominated by goby Sufflogobius bibarbatus, juvenile hake Merluccius spp. and cuttlefish Sepia spp. (on soft substrata), as well as false jacopever Sebastes capensis and kingklip Genypterus capensis (on rocky substrata).     

Feeding and vertical migration of the chaetognath Sagitta friderici (Ritter-Zahony, 1911) in the southern Benguela during spring 1987, with notes on seasonal variability of feeding ecology

The feeding biology and the vertical migration of Sagitta friderici were examined over 24 h at two stations in the southern Benguela during spring (October) 1987. Together with studies conducted during summer (February 1991) and winter (May 1984), they serve to allow valuable generalizations of the biology and ecology of this abundant chaetognath. Populations migrate vertically and feed nocturnally, although the timing and the extent of migration vary between studies. S. friderici exhibits ontogenetic layering and the cross-shelf distribution of maturity stages differs, suggesting that it is able to take advantage of cross-shelf water movement in order to maintain populations in the nearshore waters of the West Coast. S. friderici prey almost exclusively on copepods (cannibalism is rare), and there is a positive relationship between the lengths of predator and prey that is influenced by the size structure of the prey environment. This casts doubt on the validity of a chaetognath species-specific relationship between predator and prey size. S. friderici selects its prey on the basis of size, and not species. Daily ration is related linearly to prey density, so reflecting the low density of prey and providing support for theoretical predictions regarding ingestion rates under oceanic conditions. The impact of S. friderici predation on the copepod assemblage is generally less than 3% of the standing stock, although it could be much higher under conditions of low copepod biomass and poor secondary production.     

Shelf spawning habitat of Emmelichthys nitidus in south-eastern Australia – Implications and suitability for egg-based biomass estimation

The spawning habitat of Emmelichthys nitidus (Emmelichthyidae) in south-eastern Australia is described from vertical ichthyoplankton samples collected along the shelf region off eastern through to south-western Tasmania during peak spawning in October 2005–06. Surveys covered eastern waters in 2005 (38.8–43.5°S), and both eastern and southern waters in 2006 (40.5°S around to 43.5°S off the south-west). Eggs (n = 10,393) and larvae (n = 378) occurred along eastern Tasmania in both years but were rare along southern waters south and westwards of 43.5°S in 2006. Peak egg abundances (1950–2640 per m⁻²) were obtained off north-eastern Tasmania (40.5–41.5°S) between the shelf break and 2.5 nm inshore from the break. Eggs were up to 5-days old, while nearly 95% of larvae were at the early preflexion stage, i.e. close to newly emerged. Average abundances of aged eggs pooled across each survey declined steadily from day-1 to day-5 eggs both in 2005 (97-18) and 2006 (175-34). Moreover, day-1 egg abundances were significantly greater 2.5 nm at either side of the break, including at the break, than in waters ≥5 nm both inshore and offshore from the break. These results, complemented with egg and larval data obtained in shelf waters off New South Wales (NSW; 35.0–37.7°S) in October 2002–03, indicate that the main spawning area of E. nitidus in south-eastern Australia lies between 35.5°S off southern NSW and 43.5°S off south-eastern Tasmania, and that spawning activity declines abruptly south and westwards of 43.5°S around to the south-west coast. In addition, quotient analyses of day-1 egg abundances point to a preferred spawning habitat contained predominantly within a 5 nm corridor along the shelf break, where waters are 125–325 m deep and median temperatures 13.5–14.0 °C. Spawning off eastern Tasmania is timed with the productivity outburst typical of the region during the austral spring, and the temperature increase from the mixing between the southwards advancing, warm East Australian Current and cooler subantarctic water over the shelf. Overall, ichthyoplankton data, coupled with reproductive information from adults trawled off Tasmania, indicate that E. nitidus constitutes a suitable species for the application of the daily egg production method (DEPM) to estimate spawning biomass. This finding, together with evidence in support of a discrete eastern spawning stock extending from southern NSW to southern Tasmania, strengthens the need for DEPM-based biomass estimates of E. nitidus prior to further fishery expansion.     

Structure of a toothed cetacean community around a tropical island (Mayotte,Mozambique Channel)

We describe the structure of a toothed cetacean community around the island of Mayotte (South-West Indian Ocean, 45°10′ E, 12°50′ S), using data collected from small boat-based surveys conducted between July 2004 and June 2006. In all, 16 odontocete species were recorded. Diversity (Shannon-Weaver index) was particularly high along the outer slope of the barrier reef. Patterns of spatial distribution underscore the existence of three main cetacean habitat types: the inner lagoon (Indo-Pacific bottlenosed dolphin Tursiops aduncus and humpback dolphin Sousa chinensis), the outer-reef slope (spinner dolphin Stenella longirostris, pantropical spotted dolphin S. attenuate and melon-headed whale Peponocephala electra) and oceanic waters deeper than 500 m (e.g. Blainville's beaked whale Mesoplodon densirostris). Group characteristics were highly variable among species, with oceanic small delphinids characterised by larger group sizes than strictly coastal and non-delphinid oceanic species. The outer slope of the barrier reef appears to be of primary importance in terms of density and diversity of odontocetes around Mayotte. Results support the hypothesis that a number of cetacean species, particularly several delphinid species, are dependent on coral reef complexes.     

Impact of the 1997–1998 El Niño and 1999 La Niña on nutrient supply in the Gulf of Alaska

Nutrient surveys of the Gulf of Alaska, from 1997 through 1999, show that coastal waters of British Columbia and southern Alaska experienced nitrate depletion each spring and summer. Through the 1997–1998 El Niño, waters with less than 1 μM NO₃ covered 250,000 km² area greater than 1999. Silicate levels as low as 0.2 μM were observed in coastal waters, suggesting that diatom growth may have been nutrient limited both in 1998 and 1999. Detailed sampling off the southern coast of British Columbia revealed that 1998 nitrate levels were only half the average of that during the 1970s winter, were depleted 1 month earlier in spring and remained low throughout the summer. Satellite images show that, compared to 1997 and 1999, chlorophyll levels were much lower in the spring of 1998 throughout the coastal waters of the Gulf of Alaska. Conditions changed dramatically during the 1999 La Niña, with ocean-mixed layer depths increasing by 20 m in winter and 40 m in spring when compared to that during 1997–1998 El Niño. Winter nutrient levels increased and summer upwelling returned. Over the past several decades, a trend towards greater stratification of coastal waters appears to be affecting the supply of nutrients to the mixed layer. The effects of stratification were especially obvious during the 1998 El Niño.     

Macroparasites of angelfish Brama brama (Bonnaterre, 1788) in the southern Benguela Current ecosystem

The angelfish Brama brama is a mesopelagic species distributed circumglobally in temperate to warm-temperate waters, including continental-shelf-edge and upper-slope waters of the Benguela Current ecosystem. Little is known about the parasite assemblage of Benguela B. brama, with only three parasite taxa having previously been documented from this species in the southern Benguela. This study describes the macroparasites recorded from 35 B. brama collected during research surveys off the west coast of South Africa in 2015 and 2016. A total of six macroparasite taxa were documented, including the nematode Anisakis pegreffii, the copepod Hatschekia conifera, the cestode Hepatoxylon trichiuri, an acanthocephalan from the genus Rhadinorhynchus, a monogenean from the family Diclidophoridae, and an unidentified species. Three of these (He. trichiuri, Rhadinorhynchus sp. and the unidentified species) had not previously been found to infect B. brama. The most prevalent macroparasite taxa were A. pegreffii (94%), the unidentified species (71%) and Ha. conifera (60%). Two of the parasites, Ha. conifera and He. trichiuri, showed seasonal variation in infection, and infection with the latter was positively correlated with host length. These findings increase our knowledge of B. brama biology and contribute to our understanding of the biodiversity of the southern Benguela ecosystem.     

山东半岛近岸海域蟹类群落结构特征的研究

根据2014年8月和10月、2015年2月和5月使用单船底拖网进行的4个航次调查数据,对山东半岛近岸海区的莱州湾及渤海南部、山东半岛北部和山东半岛南部等3个海域蟹类种类组成、时空分布、优势种、群落结构稳定性等群落结构特征进行了研究。结果显示,4次调查共捕获蟹类20种,隶属于10科17属,其中日本蟳（Charybdis japonica）和三疣梭子蟹（Portunus trituberculatus）是主要经济种类,其余均为小型饵料型蟹类;海区内主要的优势种为双斑蟳（Charybdis bimaculata）、日本蟳（Charybdis japonica）和三疣梭子蟹（Portunus trituberculatus）,3个海域优势种季节更替均十分明显;莱州湾及渤海南部和山东半岛南部2个海域生物量季节变化明显,春季较低,在夏季达到峰值,夏季到冬季呈降低趋势,山东半岛北部海域生物量季节变化不明显。3个海域蟹类群落物种多样性指数（H'）、丰富度指数（D）及均匀度指数（J'）均处于较低水平,并有一定的季节变化;水深、水温和盐度是影响蟹类群落结构的重要因子,渤莱沿岸流、黄海暖流和黄海冷水团通过影响海域内水温与盐度影响蟹类群落结构。研究表明山东半岛近岸海区蟹类以小型饵料型蟹类为主,蟹类多样性水平较低,优势种更替明显,群落季节更替指数较高,季节间迁移频繁,蟹类群落不稳定。     

Spatial distribution of Illex argentinus in San Matias Gulf (Northern Patagonia,Argentina) in relation to environmental variables: A contribution to the new interpretation of the population structuring

Traditionally, it was assumed that major spawning activity of Illex argentinus occurs in discrete pulses along the outer-shelf/slope off Argentina/southern Brazil during late-fall/winter and that early life stages develop near the Brazil–Malvinas Confluence (BMC). However, a novel hypothesis of the population structuring of the species was proposed that states that coastal waters may be important as spawning and feeding grounds. Here, we analyzed the spatial distribution of Illex argentinus inside San Matias Gulf based on the position of the CPUE of jiggers in order to improve the knowledge of the population structuring in coastal regions. Squids were mainly concentrated on the northern region of the gulf where favorable oceanographic conditions (e.g. water stratification, chlorophyll-a concentration peaks) to feeding and spawning are present. These results provided empirical evidences that individuals of I. argentinus use Argentinean coastal waters, particularly San Matias Gulf, as permanent feeding and spawning grounds which supports the new hypothesis.     

Distribution of and predation on saury Scomberesox saurus scombroides in continental shelf waters off the Cape Province,South Africa

Although the size distribution of larvae and early juveniles of the saury Scomberesox saurus scombroides in continental shelf waters off the Cape Province, South Africa, is consistent with a south-north passive dispersal by known currents, the size of late juveniles and adults increases from north-west to south-east. Occurrence of these stages is highly seasonal: they are found in summer off the Western Cape and mainly from late summer to winter in Algoa Bay in the Eastern Cape, possibly reflecting longitudinal migration. Large numbers of late juveniles and adults are often found downstream of upwelling plumes off the Western Cape when warmer waters lie close inshore. They are sometimes taken in purse-seines in association with adult round herring Etrumeus whiteheadi and pilchard Sardinops ocellatus. Important predators of late juvenile and adult saury are fast-moving, surface-feeding species: yellowfin tuna Thunnus albacares, Cape gannet Morus capensis and Cory's shearwater Calonectris diomedea. Cape gannets consume an estimated 2 277–6 044 tons of saury annually in South African waters. Predator diets provide important time-series of occurrence, abundance and length-frequency distributions of late juvenile and adult saury.     

Relationship between measurements of hake biomass and sea surface temperature off southern Namibia

A good relationship between the variability in measurements of Cape hake Merluccius capensis biomass during winter and summer swept-area research surveys and sea surface temperature off southern Namibia for the years 1983–1990 is presented. The results call into question the hypothesis that hake density is related to biomass and imply that abnormally warm summers could support higher catches, regardless of the state of the stock. A hypothesis that anomalous warm conditions could induce hake to concentrate closer to the sea bed, making them more susceptible to bottom trawling, is presented.     

Natal site fidelity by breeding female southern elephant seals in relation to their history of participation in the winter haulout

Of the four types of terrestrial haulout periods undertaken by southern elephant seals Mirounga leonina, only the purpose of the winter haulout is unknown. Returning to a haulout site from distant pelagic foraging grounds bears significant costs in terms of increased energy expenditure, reduced foraging time and increased exposure to predation; therefore, each haulout must serve a purpose. We examined the hypothesis that the winter haulout serves to maintain familiarity with the natal site, thereby increasing site fidelity. To this end, we analysed a long-term mark-recapture dataset for female southern elephant seals at Marion Island, Southern Ocean. Results indicate that, whereas greater natal site fidelity as primiparous females was associated with recorded presence ashore at the study site during the winter haulout as immatures, this was not the case for multiparous females. Furthermore, recorded presence ashore during both the moult haulouts as immatures, and all haulouts as immatures, irrespective of haulout type, was also associated with increased site fidelity. This suggests that any haulout at the natal island as an immature seal, whether for the moult or winter haulout, assists in maintaining site fidelity. Therefore, while the winter haulout facilitates greater natal site fidelity, whether this is the sole reason for this terrestrial period remains uncertain.