Reference section 2 - Whale shark Biology and Ecology Introduction The whale shark (Rhincodon typus) is a filter feeding shark and the largest living fish. Despite its large size (up to 18 m), seasonal occurrence in coastal waters in some areas, and popularity with tourists most aspects of its biology and ecology are poorly known. This course aims to provide tour operators with enough of the available information about whale sharks to field questions from guests. This section therefore focuses on the most asked about aspects of the whale shark - where the whale shark fits into the animal kingdom, where it lives, what are the names and functions of its body parts, what it eats and what eats it. Such is the extent of our lack of knowledge of the whale shark that most questions about its life could be legitimately answered: ‘’we don’t know’’! As such this section also outlines inferences or suggestions based on what might be known from studies of other closely related species. These suggestions should not be presented by the tour operator as fact but rather used to introduce their guests to reasonable possibilities about this majestic shark and hopefully adding to a sense of wonder and inviting further discussion. Included in this section you will find: 1. 2. 3. 4. 5. 6. Taxonomy and Classification Distribution Physiology and Anatomy Diet and Feeding Reproduction and life cycle Predators KEY POINTS The whale shark is the largest fish in the world but despite its size and popularity very little is known about its biology This section outlines the facts and some of the theories regarding whale shark biology and ecology 1. Taxonomy and Classification The common name of the whale shark (a fish) can cause confusion. It refers to the fact it shares some characteristics with baleen whales (warm blooded mammals), namely its large size and use of filters to catch tiny prey. The key feature that makes sharks and rays different from all other animals including mammals, reptiles, birds and even other fish is that their internal skeleton is made of cartilage instead of bone. There are a number of other structures and adaptations that are exclusive to sharks: Cartilage skeleton – This is lighter and more flexible than bone A covering of dermal denticles – These protect the skin from damage and parasites and improve hydro-dynamics so the shark can move faster and more quietly. They have several sets of replaceable teeth – teeth are continuously being replaced and therefore are always sharp 5 to 7 gills used to extract oxygen from water Did you know? Humans have cartilage in our bodies too. That’s what the flexible part of our nose and ears are made from. These features have made sharks very successful animals. They have been dominant species on Earth, pre-dating dinosaurs by 220 million years. Today there are 440 different species of sharks ranging from smallest, the lantern shark (a deep seas species which is about 17cm long) to the largest, the whale shark. Sharks live in many different geographic and vertical habitats and some like the bull shark and river shark can survive in fresh, brackish, and salt water. The physical appearances of the 440 shark species vary substantially. Key distinguishing features like the number of dorsal fins or number of gills and the presence of an anal fin have been used to group species together into different taxonomic families and orders (i.e., closely related families). For example the mackerel sharks (order Lamniformes) include seven families including Laminae (i.e., great white, mako, salmon and porbeagle sharks). The whale shark is in the order Orectolobiformes (carpet sharks) and is the only species in the family Rhincodontidae. It doesn’t have any very close relatives but it does share some features with other carpet sharks like the nurse shark and the zebra shark. For example they all have two dorsal fins and their mouths are positioned at the very front of their bodies – in front of the eyes. The zebra shark also has clear ridges along its back like the whale shark and the nurse shark has similar nostrils. There are two other large filter feeding sharks ,the basking shark and megamouth shark, but they are in the mackerel shark order and are not closely related to the whale shark. KEY POINTS The whale shark is a true shark. It has a skeleton made of cartilage, rows of replaceable teeth, and skin covered in denticles instead of scales. Sharks are very successful animals and appeared about 220 million years before dinosaurs did. There are over 440 different species that live in diverse aquatic habitats Sharks have been split into eight orders. Whale sharks are in the order of carpet sharks (Orectolobiformes). Whale sharks are not closely related to any sharks, though they do share some features with other carpet sharks including a mouth at the front of the head, prominent lateral ridges, two dorsal fins and nostril barbells. 2. Distribution Whale sharks live in all tropic and warm-temperate seas except evidently the Mediterranean. It is thought to be primarily pelagic (prefer an open-ocean habitat) but seasonal feeding aggregations do occur at several coastal sites throughout the tropics. The most well documented sites are in the Gulf of Mexico, Gulf of California, Belize, Honduras, Western Australia, the Galapagos, New Zealand, Philippines, Indonesia, Madagascar, Mozambique, Kenya, India, Pakistan, Maldives, Seychelles, Indonesia, and Thailand. Whale sharks have been found offshore and close to land, entering lagoons or coral atolls, and near the mouths of estuaries and rivers. Its range is generally restricted to about ±30° latitude although whale sharks have been seen at higher latitudes on occasion off the coast of South Africa and eastern North America. Although whale sharks exist mainly in areas where the sea surface temperature is between 25˚C and 30˚C tagging studies have shown they are capable of diving very deep in excess of 1600 metres where they tolerate temperatures as low as 3˚C. Whale sharks are highly mobile animals, capable of swimming thousands of miles. Despite often being described as such, it is yet to be seen whether whale sharks are truly migratory. Migratory animals move between particular areas on regular seasonal or annual schedules as opposed to simple movements within more defined home ranges, or immigration or emigration. KEY POINTS Whale sharks are found throughout all tropic and warm temperate seas. There are about 18 to 20 countries where whale sharks are known to occur at particular seasons. Whale sharks spend time in offshore and near shore marine habitats. Whale sharks can dive deeply (up to 1600m or more) and tolerate very low water temperature (down to 3˚C) Although not necessarily migratory the whale shark is a highly mobile animal capable of traveling thousands of miles annually. 3. Physiology and Anatomy The largest whale shark that has been directly measured was reported to be 12.65 metres (41.50 ft) long and a weighed more than 21.5 tonnes (47,000 lb), though few other reliable measurements have been made Some claims of of sharks up to 18m long have been made though none validated. The body surface coloration is mostly grey dorsally (above) and white ventrally (below). Three prominent ridges run along the sides and the skin is marked with a checkerboard of spots and stripes which may function as camouflage. The patterns of spots are used in some areas to identify sharks from photographs, amethods that assumes that they are unique to an individual and that they do not change with age. The skin of whale sharks is relatively thin, a couple of millimeters, and is covered in tiny teeth-like scales called denticles. These denticles provide a tough, hydrodynamic surface layer. Below the skin is a layer of fatty tissue that can be 10-15cm thick and is probably important for insulating the shark’s muscles and vital organs and perhaps a nutrient and energy store. Whale sharks have two dorsal fins and two lateral pectoral fins. The upper lobe of the tail (i.e., caudal) fin of juvenile sharks is larger than the lower lobe compared with adults which have a more crescent-shaped tail. Spiracles Spiracles are the small circular openings just behind the whale shark’s eyes. These openings appear in many species of sharks and are actually the remains of a pair of gills that grown apart from the other five pairs of gills. The function of the spiracles in whale sharks is not known. In some species of bottom-dwelling sharks (e.g. nurse sharks) spiracles are used in respiration, creating a flow of water that delivers oxygen to the gills when the animal is at rest. Large pelagic or open ocean sharks like the great white shark do not have spiracles and have to keep moving to create this flow of oxygenated water, over the gills. The function of the spiracles of whale sharks might be intermediate as whale sharks are more closely related to bottom-dwelling sharks though they also spend substantial time swimming throughout the water column. Consequently, the spiracles might function when whale sharks are occasionally stationary but not feeding (when water is passing over the gills). Mouth The mouth of this large filter feeder is also relatively large and can be up to 1.5 metres wide. It contains between 300 and 350 rows of tiny (3mm) scale-like teeth which look like a rasp close-up and account for the species original latin name Rhinodonte, which means rasp-tooth. These teeth do not appear to have function in feeding. Gills Whale sharks have five large pairs of gills which serve two functions They extract oxygen from the water to support metabolism and also filter out small prey from seawater. The gills are modified with 20 filter pads (two for each gill). These filters have tiny 1.5 millimeter gaps. Water entering the mouth is forced past the holes in the filters and out through the gill slits. Any organisms larger than 0.5 millimeters is sieved out by the filters and swallowed by the shark. Eyes There are two small eyes located near the front of the wide, flat head. The forward, lateral placement of the eyes allows the sharks to see forward, backward and above and below. The eyes appear quite well evolved though vision is likely rather poor and limited to detecting relatively close (i.e., < 3 m) movements. The eyes can roll and retract into the sockets when something approaches very closely if the shark is exposed to a flash from a camera strobe. Nostrils The nostrils are relatively small and not very well developed. However, their position at the front of the head above the mouth and far apart suggests that they may have an important function in detection of substances in the water. The large, well developed internal sensory structures (olfactory capsules) connected to the nostrils support this suggestion. Similar findings of an acute sense of smell in its close relative the nurse shark suggests the detection of chemicals dissolved in seawater may be important in locating concentrated prey. Other Whale Shark Senses The hearing of whale sharks has not been studied. Anatomically, their inner ear is the largest of any animals, perhaps simply because of the species large physical size or perhaps related to adaptations for low frequency hearing. Similar to some other sharks, whale sharks have anatomical structures that function to detect vibrations (lateral line) in water and small electrical currents (ampullae of Lorenzini) produced by other animals. These structures may have similar function in whale sharks though they have not been investigated yet. KEY POINTS Largest whale shark measured was 12.85m although they might grow as long as 18m. Whale shark skin is thin, though tough, with a thick fatty layer below it. It is covered by teeth-like scales called denticles Whale sharks’ bodies are covered by spots and stripes that are being used to identify individuals, conditioned on assumptions that patterns are unique to individual sharks and that they do not change with age or size. The whale shark has 300 rows of very tiny rasp-like teeth that apparently have no function The gills of whale sharks function to extract oxygen and to filter prey from seawater. Spiracles are the small holes behind the whale sharks eyes which in some closely related sharks are connected by channels to their gills and provide some ventilation when the sharks are stationary. The small eyes are on the side of the head, apparently to maximize range of vision. Visual detection is probably limited to movements very nearby (<3m). The two nostrils are at the very front of the shark above the mouth and might function to detect (i.e., smell) chemicals dissolved in seawater. How important each different sense is to the whale shark is unknown but they probably use a combination of vision, smell, hearing and detection of electrical currents. 4. Diet and Feeding Whale sharks mostly eat tiny planktonic animals (e.g. euphausiids, copepods, fish and squid eggs or larvae) though they have also been observed to ingest small fishes like sardines and anchovies. Whale sharks are able to feed in two different ways: ram filter feeding,which involves swimming along with their mouths open and feeding on prey that happen to be caught as water passes through the gill rakers. suction filter feeding,where the whale shark can actually remain stationary in the water and by opening and closing its mouth can actively suck large quantities of water and prey in to its mouth. When the mouth closes the water is forced out through the filter pads and any prey is sieved out. Suction feeding allows the whale shark to feed on more mobile prey like small fish which would usually be able to escape from a ram filter feeder. It has been suggested that the whale sharks gills are more suited to suction filter feeding than they are to ram filter feeding because the filter pads on their gills have smaller gaps and work best when water is being physically forced through rather than just flowing through them. The structure of the gills in other filter feeding sharks (e.g. basking sharks) appear to have contrasting structures with larger gaps that appear to be only suitable for ram filter feeding. Whale sharks evidently only use the suction method when they are feeding on very high concentrations of prey. Consequently, some think that the sharks might actively search for specific areas or seasonal events where high densities of food are more likely and that this might explain why whale shark aggregations are seasonal in most areas. In areas where there is no clear whale shark season (e.g., the Maldives) other factors like the lunar cycle might have stronger influences on the number of sharks observed. These are also natural triggers for spawning and plankton blooming events. Information collected from tagging studies has shown that whale sharks are constantly diving and ascending spending only short periods of time near the surface and diving as deep as 1600 metres, evidently to search for the areas where there are high densities of prey in the water column. KEY POINTS Whale sharks feed on tiny prey including plankton, fish and squid eggs and larvae, and small shoaling baitfish. They use two different feeding methods: ram and suction filter feeding Suction filter feeding evidently enables them to feed on more mobile prey There are two other filter feeding sharks, the basking shark and the megamouth shark Whale sharks apparently seek out high densities of prey and as a result they are often seasonal visitors to specific locations Whale sharks constantly dive and ascend, perhaps to actively search for areas of high prey density 5. Reproduction and life cycle No one has ever witnessed whale shark courtship, mating, or birth. It is now thought that whale sharks give birth to relatively large litters of live young (i.e., ovoviviparity), though this is based on observations of one pregnant female. During mating it is thought that male whale sharks inseminate females with their claspers (sexual organs ( located between the ventral pelvic fins). The presence of claspers is the easiest way to determine the sex of a whale shark, though they can be hard to identify in very young males. Females are thought to be capable of storing a male’s sperm and fertilizing many different eggs, retaining the egg cases until they hatch inside their uteri until they hatch. It is not known how long this process takes (gestation period) no one knows nor how often it occurs. One female whale shark captured off Taiwan had 300 embryos in her uterus in various stages of development. Whale shark offspring or ‘pups’ are thought to be about 55–60 cm in length at birth and weigh about 1 kg and grow to about 1.4 m long and 20 kg within the next four months. Whale sharks like most sharks mature very slowly, perhaps not maturing sexually until around 8 to 9 metres long and 25 to 35 years old and may live for up to 70 to 100 years, though there is very little evidence to support these suggestions. Did you know? Scientists are attempting to estimate whale shark age by counting the number of growth rings in the vertebrae (back bones) of dead whale sharks - much like one would count the rings on a tree stump. The problem is currently they can only make a good guess as to what one growth ring represents in units of time. Another mystery is that groups or aggregations of whale sharks in most hot-spots are composed primarily of adolescents with very few females and virtually no adults. In the Indian Ocean for example (e.g., the Seychelles, Maldives, Mozambique and Western Australia) over 60% of the sharks observed are adolescent males. The Galapagos Island archipelago in the Pacific Ocean is the only site where large mature female sharks are regularly seen. Remember! We must keep in mind that humans can only encounter whale sharks with any regularity near the surface and near the coast. It may simply be that females prefer a slightly different habitat to the males - one that is out of reach of humans - deeper down or further out to sea. KEY POINTS No one has ever reported whale shark reproductive behaviour. Males inserts one of its claspers into the female, females store sperm and fertilize many eggs. Whale sharks give birth to live young – they are Ovoviviparous Pups are about 60cm when they are born WS do not reach sexual maturity until approximately 9m or 30 years old and possibly live until 70 -100 years old The vast majority of whale sharks encountered by humans around the world are juvenile males 6. Predators The whale sharks best defence is its large size and as such adult whale sharks do not have many natural enemies. There are however reports of great white sharks and killer whales attacking adult whale sharks. New born whale sharks are much more vulnerable and specimens have been found in the stomachs of blue sharks and swordfish. KEY POINTS Whale sharks do not have many natural enemies due to their size Great white shark and killer whale attacks on adult whale sharks have been reported Juvenile sharks are more vulnerable. Young sharks found in stomachs of smaller sharks and swordfish