The Biology of Tiger Sharks, Galeocerdo Cuvier, in Shark Bay, Western Australia: Sex Ratio, Size Distribution, Diet, and Seasonal Changes in Catch Rates

Tiger sharks, Galeocerdo cuvier, are apex predators in a variety of nearshore ecosystems throughout the world. This study investigates the biology of tiger sharks in the shallow seagrass ecosystem of Shark Bay, Western Australia. Tiger sharks (n = 252) were the most commonly caught species (94%) compared to other large sharks. Tiger sharks ranged from 148–407cm TL. The overall sex ratio was biased towards females (1.8:1), but the sex ratio of mature animals (> 300cm TL) did not differ from 1:1. Contrary to previous accounts, tiger sharks were caught more often in all habitats during daylight hours than at night. Tiger shark catch rates were highly correlated with water temperature and were highest when water temperatures were above 19°C. The seasonal abundance of tiger sharks is correlated to both water temperature and the occurrence of their main prey: sea snakes and dugongs, Dugong dugon. Stomach contents analysis indicated that sea turtles and smaller elasmobranchs were also common prey. The importance of major seagrass grazers (dugongs and green sea turtles, Chelonia mydas) in the diet of tiger sharks suggests the possibility that these sharks are keystone predators in this ecosystem.     

Ontogenetic dietary shifts and feeding behavior of the tiger shark, Galeocerdo cuvier, in Hawaiian waters

Synopsis Stomach content data from 281 tiger sharks caught during shark control programs in Hawaii between 1967 and 1969, and during 1976 were analyzed to examine feeding habits and ontogenetic shifts in diet. As sharks increased in size, prey diversity and frequency of occurrence of large prey items increased. The percent occurrence of teleosts and cephalopods in stomachs decreased as sharks increased in length, while occurrence of elasmobranchs, turtles, land mammals, crustaceans, and undigestible items increased. Comparisons between the diets of tiger sharks from Hawaii and other locations indicate that ontogenetic shifts are universal in this species and that tiger sharks may be opportunistic feeders that prey heavily on abundant, easy to capture prey. Small tiger sharks may be spatially segregated from medium and large sharks and appear to be primarily nocturnal, bottom feeders. Large tiger sharks feed near the bottom at night, but also feed at the surface during the day. Prey, similar in size to humans, begin to occur in the diet of tiger sharks approximately 230 cm TL, and therefore sharks of this size and larger may pose the greatest threat to humans. Ontogenetic shifts in diet may be attributed to increased size of sharks, expanded range and exploitation of habitats of larger sharks, and/or improved hunting skill of larger sharks.Deceased 1974     

The Last Frontier: Catch Records of White Sharks (Carcharodon carcharias) in the Northwest Pacific Ocean

White sharks are highly migratory apex predators, globally distributed in temperate, sub-tropical, and tropical waters. Knowledge of white shark biology and ecology has increased recently based on research at known aggregation sites in the Indian, Atlantic, and Northeast Pacific Oceans; however, few data are available for the Northwest Pacific Ocean. This study provides a meta-analysis of 240 observations of white sharks from the Northwest Pacific Ocean between 1951 and 2012. Records comprise reports of bycatch in commercial fisheries, media accounts, personal communications, and documentation of shark-human interactions from Russia (n = 8), Republic of Korea (22), Japan (129), China (32), Taiwan (45), Philippines (1) and Vietnam (3). Observations occurred in all months, excluding October-January in the north (Russia and Republic of Korea) and July-August in the south (China, Taiwan, Philippines, and Vietnam). Population trend analysis indicated that the relative abundance of white sharks in the region has remained relatively stable, but parameterization of a 75% increase in observer effort found evidence of a minor decline since 2002. Reliably measured sharks ranged from 126–602 cm total length (TL) and 16–2530 kg total weight. The largest shark in this study (602 cm TL) represents the largest measured shark on record worldwide. For all countries combined the sex ratio was non-significantly biased towards females (1∶1.1; n = 113). Of 60 females examined, 11 were confirmed pregnant ranging from the beginning stages of pregnancy (egg cases) to near term (140 cm TL embryos). On average, 6.0±2.2 embryos were found per litter (maximum of 10) and gestation period was estimated to be 20 months. These observations confirm that white sharks are present in the Northwest Pacific Ocean year-round. While acknowledging the difficulties of studying little known populations of a naturally low abundance species, these results highlight the need for dedicated research to inform regional conservation and management planning.     

A novel mode of embryonic nutrition in the tiger shark,Galeocerdo cuvier

How are tiger shark embryos nourished to large size without a placental connection? Tiger sharks belong to the family Carcharhinidae, and all carcharhinid sharks are placental with the exception of the tiger shark. The aim of this study was to test the hypothesis that tiger shark embryos are nourished to large size by imbibing a clear uterine fluid found in their egg cases. Based on weights of fertilized eggs and of term embryos, the tiger shark is a matrotrophic species, and its embryos appear to reach gains of 2119% in wet weight and 1092% in dry weight during gestation. By measuring the total energy content of the fluid in the egg case by chemical oxygen demand (COD), the authors demonstrate that clear liquid in the tiger shark egg case is an energy-rich embryotrophe that nourishes the embryos to large size. We suggest that the process be termed ‘embryotrophy'. The process appears to be an adaptation for producing large broods of large embryos in a species lacking a placental connection.     

Vertical Movement Patterns and Ontogenetic Niche Expansion in the Tiger Shark,Galeocerdo cuvier

Sharks are top predators in many marine ecosystems and can impact community dynamics, yet many shark populations are undergoing severe declines primarily due to overfishing. Obtaining species-specific knowledge on shark spatial ecology is important to implement adequate management strategies for the effective conservation of these taxa. This is particularly relevant concerning highly-mobile species that use wide home ranges comprising coastal and oceanic habitats, such as tiger sharks, Galeocerdo cuvier. We deployed satellite tags in 20 juvenile tiger sharks off northeastern Brazil to assess the effect of intrinsic and extrinsic factors on depth and temperature usage. Sharks were tracked for a total of 1184 d and used waters up to 1112 m in depth. The minimum temperature recorded equaled 4°C. All sharks had a clear preference for surface (< 5 m) waters but variability in depth usage was observed as some sharks used mostly shallow (< 60 m) waters whereas others made frequent incursions into greater depths. A diel behavioral shift was detected, with sharks spending considerably more time in surface (< 10 m) waters during the night. Moreover, a clear ontogenetic expansion in the vertical range of tiger shark habitat was observed, with generalized linear models estimating a ~4-fold increase in maximum diving depth from 150- to 300-cm size-classes. The time spent in the upper 5 m of the water column did not vary ontogenetically but shark size was the most important factor explaining the utilization of deeper water layers. Young-of-the-year tiger sharks seem to associate with shallow, neritic habitats but they progressively move into deeper oceanic habitats as they grow larger. Such an early plasticity in habitat use could endow tiger sharks with access to previously unavailable prey, thus contributing to a wider ecological niche.     

Structure and Dynamics of the Shark Assemblage off Recife,Northeastern Brazil

Understanding the ecological factors that regulate elasmobranch abundance in nearshore waters is essential to effectively manage coastal ecosystems and promote conservation. However, little is known about elasmobranch populations in the western South Atlantic Ocean. An 8-year, standardized longline and drumline survey conducted in nearshore waters off Recife, northeastern Brazil, allowed us to describe the shark assemblage and to monitor abundance dynamics using zero-inflated generalized additive models. This region is mostly used by several carcharhinids and one ginglymostomid, but sphyrnids are also present. Blacknose sharks, Carcharhinus acronotus, were mostly mature individuals and declined in abundance throughout the survey, contrasting with nurse sharks, Ginglymostoma cirratum, which proliferated possibly due to this species being prohibited from all harvest since 2004 in this region. Tiger sharks, Galeocerdo cuvier, were mostly juveniles smaller than 200 cm and seem to use nearshore waters off Recife between January and September. No long-term trend in tiger shark abundance was discernible. Spatial distribution was similar in true coastal species (i.e. blacknose and nurse sharks) whereas tiger sharks were most abundant at the middle continental shelf. The sea surface temperature, tidal amplitude, wind direction, water turbidity, and pluviosity were all selected to predict shark abundance off Recife. Interspecific variability in abundance dynamics across spatiotemporal and environmental gradients suggest that the ecological processes regulating shark abundance are generally independent between species, which could add complexity to multi-species fisheries management frameworks. Yet, further research is warranted to ascertain trends at population levels in the South Atlantic Ocean.     

Crossing Latitudes—Long-Distance Tracking of an Apex Predator

Tiger sharks (Galeocerdo cuvier) are apex predators occurring in most tropical and warm temperate marine ecosystems, but we know relatively little of their patterns of residency and movement over large spatial and temporal scales. We deployed satellite tags on eleven tiger sharks off the north-western coast of Western Australia and used the Brownian Bridge kernel method to calculate home ranges and analyse movement behaviour. One individual recorded one of the largest geographical ranges of movement ever reported for the species, travelling over 4000 km during 517 days of monitoring. Tags on the remainder of the sharks reported for shorter periods (7-191 days). Most of these sharks had restricted movements and long-term (30-188 days) residency in coastal waters in the vicinity of the area where they were tagged. Core home range areas of sharks varied greatly from 1166.9 to 634,944 km². Tiger sharks spent most of their time in water temperatures between 23°-26°C but experienced temperatures ranging from 6°C to 33°C. One shark displayed seasonal movements among three distinct home range cores spread along most of the coast of Western Australia and generalized linear models showed that this individual had different patterns of temperature and depth occupancy in each region of the coast, with the highest probability of residency occurring in the shallowest areas of the coast with water temperatures above 23°C. These results suggest that tiger sharks can migrate over very large distances and across latitudes ranging from tropical to the cool temperate waters. Such extensive long-term movements may be a key element influencing the connectivity of populations within and among ocean basins.     

Residency Patterns and Migration Dynamics of Adult Bull Sharks (Carcharhinus leucas) on the East Coast of Southern Africa

Bull sharks (Carcharhinus leucas) are globally distributed top predators that play an important ecological role within coastal marine communities. However, little is known about the spatial and temporal scales of their habitat use and associated ecological role. In this study, we employed passive acoustic telemetry to investigate the residency patterns and migration dynamics of 18 adult bull sharks (195–283 cm total length) tagged in southern Mozambique for a period of between 10 and 22 months. The majority of sharks (n = 16) exhibited temporally and spatially variable residency patterns interspersed with migration events. Ten individuals undertook coastal migrations that ranged between 433 and 709 km (mean  = 533 km) with eight of these sharks returning to the study site. During migration, individuals exhibited rates of movement between 2 and 59 km.d⁻¹ (mean  = 17.58 km.d⁻¹) and were recorded travelling annual distances of between 450 and 3760 km (mean  = 1163 km). Migration towards lower latitudes primarily took place in austral spring and winter and there was a significant negative correlation between residency and mean monthly sea temperature at the study site. This suggested that seasonal change is the primary driver behind migration events but further investigation is required to assess how foraging and reproductive activity may influence residency patterns and migration. Results from this study highlight the need for further understanding of bull shark migration dynamics and suggest that effective conservation strategies for this vulnerable species necessitate the incorporation of congruent trans-boundary policies over large spatial scales.     

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.     

Segregation and foraging ecology of whale sharks, Rhincodon typus, in the southwestern Gulf of California

Sharks segregate by sex and size, but few studies have attempted to explain such behaviors. To address this, we examined aggregations and the foraging ecology of whale sharks in Bahía de La Paz (BLP) with aerial and ship surveys and direct observation. Zooplankton abundance and composition, and hydrographic conditions were analyzed in relation to whale shark occurrence to explore underlying factors causing segregations. We observed large aggregations of juveniles (<9 m total length, TL) inshore, comprised by 60 % male individuals, and small aggregations of adults (>9 m TL) offshore, composed of 84 % females. Juvenile sharks were associated to turbid shallow waters in BLP, where they performed stationary and dynamic suction feeding on dense copepod swarms. Adults occurred in oceanic waters and fed by ram-filtering on diffuse patches of euphausiids, with no association to oceanographic conditions. Such segregation may be advantageous to juvenile R. typus utilizing shallow coastal waters to find abundant preferred prey needed for their fast growth rates. Our studies suggest that the main driving forces of whale shark segregation by sex and size in BLP may be diet preference for juveniles and habitat preference for adult sharks.     

Intraspecific variation in vertical habitat use by tiger sharks (Galeocerdo cuvier) in the western North Atlantic

Tiger sharks (Galeocerdo cuvier) are a wide ranging, potentially keystone predator species that display a variety of horizontal movement patterns, making use of coastal and pelagic waters. Far less, however, is known about their vertical movements and use of the water column. We used pop‐up satellite archival tags with two data sampling rates (high rate and standard rate tags) to investigate the vertical habitat use and diving behavior of tiger sharks tagged on the Puerto Rico–Virgin Islands platform and off Bermuda between 2008 and 2009. Useable data were received from nine of 14 sharks tagged, tracked over a total of 529 days. Sharks spent the majority of their time making yo‐yo dives within the upper 50 m of the water column and considerable time within the upper 5 m of the water column. As a result, sharks typically occupied a narrow daily temperature range (~2°C). Dives to greater than 200 m were common, and all sharks made dives to at least 250 m, with one shark reaching a depth of 828 m. Despite some similarities among individuals, a great deal of intraspecific variability in vertical habit use was observed. Four distinct depth distributions that were not related to tagging location, horizontal movements, sex, or size were detected. In addition, similar depth distributions did not necessitate similar dive patterns among sharks. Recognition of intraspecific variability in habitat use of top predators can be crucial for effective management of these species and for understanding their influence on ecosystem dynamics.     

Age and growth of the bonnethead shark, Sphyrna tiburo, from northwest Florida, with comments on clinal variation

Age and growth rates of the bonnethead shark, Sphyrna tiburo, from northwest Florida were estimated from vertebrae collected between October 1992 and October 1995. The von Bertalanffy growth equation was fit to male and female vertebral age data. Initial growth was rapid (≈ 200 mm TL) for both sexes from age 0–1. At age 2 growth slowed for males but continued for females. Similar to many species of sharks, females grew slower than males (K = 0.28 and K = 0.69, respectively) but attained a larger maximum size (L_∞=1226 and L_∞=897). Maximum age was estimated in males and females to be 8+ and 12+ years, respectively. Growth of young-of-year sharks was 21 to 30 mm TL per month determined by three different methods. A comparison of age and growth estimates from populations at more southerly latitudes suggest that clinal variation in total length may be evident among bonnethead sharks in the Gulf of Mexico with females reaching larger sizes in northern areas as compared to south Florida. This revised version was published online in July 2006 with corrections to the Cover Date.     

Characterization of a Subtropical Hawksbill Sea Turtle (Eretmocheyles imbricata) Assemblage Utilizing Shallow Water Natural and Artificial Habitats in the Florida Keys

In order to provide information to better inform management decisions and direct further research, vessel-based visual transects, snorkel transects, and in-water capture techniques were used to characterize hawksbill sea turtles in the shallow marine habitats of a Marine Protected Area (MPA), the Key West National Wildlife Refuge in the Florida Keys. Hawksbills were found in hardbottom and seagrass dominated habitats throughout the Refuge, and on man-made rubble structures in the Northwest Channel near Cottrell Key. Hawksbills captured (N = 82) were exclusively juveniles and subadults with a straight standard carapace length (SSCL) ranging from 21.4 to 69.0cm with a mean of 44.1 cm (SD = 10.8). Somatic growth rates were calculated from 15 recaptured turtles with periods at large ranging from 51 to 1188 days. Mean SSCL growth rate was 7.7 cm/year (SD = 4.6). Juvenile hawksbills (<50 cm SSCL) showed a significantly higher growth rate (9.2 cm/year, SD = 4.5, N = 11) than subadult hawksbills (50–70 cm SSCL, 3.6 cm/year, SD = 0.9, N = 4). Analysis of 740 base pair mitochondrial control region sequences from 50 sampled turtles yielded 12 haplotypes. Haplotype frequencies were significantly different compared to four other Caribbean juvenile foraging aggregations, including one off the Atlantic coast of Florida. Many-to-one mixed stock analysis indicated Mexico as the primary source of juveniles in the region and also suggested that the Refuge may serve as important developmental habitat for the Cuban nesting aggregation. Serum testosterone radioimmunoassay results from 33 individuals indicated a female biased sex ratio of 3.3 females: 1 male for hawksbills in the Refuge. This assemblage of hawksbills is near the northern limit of the species range, and is one of only two such assemblages described in the waters of the continental United States. Since this assemblage resides in an MPA with intensive human use, basic information on the assemblage is vital to resource managers charged with conservation and species protection in the MPA.     

Occurrence and biological characteristics of broadnose sevengill sharks (Notorynchus cepedianus) in Pacific Northwest coastal estuaries

The broadnose sevengill shark (Notorynchus cepedianus) is a high-order marine predator distributed worldwide in shallow coastal waters of temperate seas. Recent reports have suggested it may be a prevalent component of Pacific Northwest coastal estuarine communities, although biological characteristics of the shark population remain undocumented despite growing interest in recreational harvest of the species. Longline sampling was conducted in Willapa Bay and Grays Harbor, Washington, USA seasonally during 2003–2006 to collect sevengill shark size, maturity, and sex ratio data, and establish some baseline catch rate information. Sevengill sharks were collected on 65% of longline sets and catches were composed of subadult and mature individuals (122–283 cm TL) of both sexes. Most male sevengill sharks were large sexually mature adults, based on external clasper calcification levels, whereas most comparably sized females were considered subadults, based on literature-based size-at–maturity estimates. Neonates and young sharks <120 cm were not collected, nor have they been reported in other historic estuary sampling efforts. Sex ratios were skewed toward males in Willapa Bay and suggest some degree of sexual segregation for the species, as has been shown for populations elsewhere. We suggest sevengill sharks are a largely ignored but potentially important predator in Pacific Northwest estuaries. This study therefore provides some of the first, basic information for guiding management decisions associated with a late-maturing, slow-growing shark species in these coastal habitats.     

Food habits and ontogenetic changes in the diet of the sandbar shark, Carcharhinus plumbeus, in Hawaii

The sandbar shark, Carcharhinus plumbeus, is a wide-ranging coastal species in tropical and temperate regions, and it is the most common species of shark in Hawaii, as in many locations where it occurs. Information on the diet and feeding habits of this species in the Pacific Ocean are extremely limited. For this study we quantified the diet of sandbar sharks in Hawaii based on records collected during the Hawaii Cooperative Shark Research and Control Program from 1967 to 1969. During this program a total of 565 stomachs were examined, of which 265 contained food. Sharks ranged in size from 59 to 190 cm total length. Teleosts were the most common prey group, but both cephalopods and crustaceans also occurred frequently. Ontogenetic changes in diet of sandbar sharks were apparent, with crustaceans forming a greater proportion of the diet of smaller sharks. Both cephalopods and elasmobranchs increased in importance with increasing shark size. Prey diversity also increased with size, with large, mobile, and reef prey species found more commonly in the diet of larger sharks. Mature male and female sharks appeared to segregate by depth, though major differences in the diet between the sexes were not apparent. However, there was some evidence of dietary differences between sharks caught in different depths and seasons. The results of this study suggest that sandbar sharks in Hawaii and throughout the world, are primarily piscivores, but also consume a variety of invertebrate prey, and that their diet varies with geographical location and stage of development.     

Vertebral Bomb Radiocarbon Suggests Extreme Longevity in White Sharks

Conservation and management efforts for white sharks (Carcharodon carcharias) remain hampered by a lack of basic demographic information including age and growth rates. Sharks are typically aged by counting growth bands sequentially deposited in their vertebrae, but the assumption of annual deposition of these band pairs requires testing. We compared radiocarbon (Δ¹⁴C) values in vertebrae from four female and four male white sharks from the northwestern Atlantic Ocean (NWA) with reference chronologies documenting the marine uptake of ¹⁴C produced by atmospheric testing of thermonuclear devices to generate the first radiocarbon age estimates for adult white sharks. Age estimates were up to 40 years old for the largest female (fork length [FL]: 526 cm) and 73 years old for the largest male (FL: 493 cm). Our results dramatically extend the maximum age and longevity of white sharks compared to earlier studies, hint at possible sexual dimorphism in growth rates, and raise concerns that white shark populations are considerably more sensitive to human-induced mortality than previously thought.     

Ontogeny of head and caudal fin shape of an apex marine predator: The tiger shark (Galeocerdo cuvier)

How morphology changes with size can have profound effects on the life history and ecology of an animal. For apex predators that can impact higher level ecosystem processes, such changes may have consequences for other species. Tiger sharks (Galeocerdo cuvier) are an apex predator in tropical seas, and, as adults, are highly migratory. However, little is known about ontogenetic changes in their body form, especially in relation to two aspects of shape that influence locomotion (caudal fin) and feeding (head shape). We captured digital images of the heads and caudal fins of live tiger sharks from Southern Florida and the Bahamas ranging in body size (hence age), and quantified shape of each using elliptical Fourier analysis. This revealed changes in the shape of the head and caudal fin of tiger sharks across ontogeny. Smaller juvenile tiger sharks show an asymmetrical tail with the dorsal (upper) lobe being substantially larger than the ventral (lower) lobe, and transition to more symmetrical tail in larger adults, although the upper lobe remains relatively larger in adults. The heads of juvenile tiger sharks are more conical, which transition to relatively broader heads over ontogeny. We interpret these changes as a result of two ecological transitions. First, adult tiger sharks can undertake extensive migrations and a more symmetrical tail could be more efficient for swimming longer distances, although we did not test this possibility. Second, adult tiger sharks expand their diet to consume larger and more diverse prey with age (turtles, mammals, and elasmobranchs), which requires substantially greater bite area and force to process. In contrast, juvenile tiger sharks consume smaller prey, such as fishes, crustaceans, and invertebrates. Our data reveal significant morphological shifts in an apex predator, which could have effects for other species that tiger sharks consume and interact with. J. Morphol. 277:556–564, 2016. © 2016 Wiley Periodicals, Inc.     

Biology of the Galapagos shark,Carcharhinus galapagensis, in Hawai'i

Synopsis Catch records from the Hawai'i Cooperative Shark Research and Control Program, which operated in Hawai'i from 1967–1969, were examined and data on the Galapagos shark,Carcharhinus galapagensis were analyzed. A total of 304 Galapagos sharks was caught, predominantly with longlines. More female sharks were caught than males, and the catch was skewed geographically. On the island of O'ahu the highest catch rates occurred along the north and south coasts. High catch rates also occurred near points of land, where longshore currents converge. Average depth of capture was greater for juveniles (45.1 m) and mature males (60.2 m), than for subadults (38.8 m) and mature female sharks (34.2 m). Males appear to reach maturity between 205 and 239 cm total length, and females between 215 and 245 cm. Litter size ranged from 4 to 16 pups, with an average of 8.7. In Hawaiian waters Galapagos sharks are born at just over 80 cm total length. Mating and parturition apparently occur early in the year, and gestation is estimated to be about 12 months. Stomach contents consisted mainly of teleosts and benthic prey, and ontogenetic changes in diet occurred as sharks increased in size. Sharks consumed a smaller proportion of teleosts and more elasmobranchs with increasing size. Dietary diversity also increased with increasing size of shark.     

Whale Shark (Rhincodon typus) Seasonal Presence,Residence Time and Habitat Use at Darwin Island,Galapagos Marine Reserve

The life history of the whale shark (Rhincodon typus), including its reproductive ecology, still remains largely unknown. Here, we present results from the first whale shark population study around Darwin Island, Galapagos Marine Reserve. Following a diversified approach we characterized seasonal occurrence, population structure and size, and described habitat use of whale sharks based on fine scale movements around the island. Whale shark presence at Darwin Island was negatively correlated with Sea Surface Temperature (SST), with highest abundance corresponding to a cool season between July and December over six years of monitoring. From 2011 to 2013 we photo-identified 82 whale sharks ranging from 4 to 13.1 m Total Length (TL). Size distribution was bimodal, with a great majority (91.5%) of adult female individuals averaging 11.35 m±0.12 m (TL±SE), all but one showing signs of a potential pregnancy. Population dynamics models for apparently pregnant sharks estimated the presence of 3.76±0.90 (mean ± SE) sharks in the study area per day with an individual residence time of 2.09±0.51 (mean ± SE) days. Movement patterns analysis of four apparently pregnant individuals tracked with acoustic tags at Darwin Island revealed an intense use of Darwin''s Arch, where no feeding or specific behavior has been recorded, together with periodic excursions around the island''s vicinity. Sharks showed a preference for intermediate depths (20–30 m) with occasional dives mostly to mid-water, remaining the majority of their time at water temperatures between 24–25°C. All of our results point to Darwin Island as an important stopover in a migration, possibly with reproductive purposes, rather than an aggregation site. Current studies carried out in this area to investigate regional scale movement patterns may provide essential information about possible pupping grounds for this enigmatic species.