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(Birgus latro) A Comprehensive Account Of The Biology & Conservation Issues


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INTRODUCTION
The immense Coconut Crab (or Robber Crab) is the largest terrestrial arthropod on earth, (Grubb 1971). They have been found to attain weights in excess of 5kg (Fletcher & Amos 1994) and can span over 50cm from claw to claw, (Reyne 1939). They are very powerful creatures and have vastly strong pincers. These chelae can easily cut through an ordinary broom handle, (Johnson 1965). The Coconut Crab comes from the same family as the hermit crab. The family Coenobitatae contains only two genera Coenobita, the land-hermit-crabs, and Birgus, containing only the Coconut Crab, Birgus latro, (Johnson 1965). Although the two genera appear very different in adulthood, they share extremely similar juvenile lives. Like adult land hermit crabs, the Coconut Crab occupies an old snail shell when it is young. Eventually, it gets so big that it cannot find any suitable shells. It then undergoes metamorphosis into the adult form, (Harms 1932). As an adult, Birgus look very peculiar. They are easily recognised by their spider like abdomen, which is hard and dull coloured above but soft and brilliantly coloured below, (Johnson 1965). The crab occurs in two different colour phases, a blue and a red (Grubb Harms 1971). From the front, the Coconut Crab still appears very similar to a hermit crab. However hermit crabs have special hind-legs adapted for gripping onto their adopted shell. In adult Coconut Crabs, this last pair of legs has become greatly reduced, (Johnson 1965). The next pair has become readapted for walking and is used to help the crab climb trees. The Coconut Crab lives almost exclusively on small tropical islands. The wide distribution across the tropical regions of the Indo-Pacific covers a region of 180 longitude, (Reyne 1939). The crab is also present outside the tropics on islands washed with tropical warm currents. An example is the Ryu Kyu Islands of Japan, which are washed by the warm Japan Stream, (Reyne 1939). Birgus is also found in the Indian Ocean on islands such as Aldabra and Christmas Island, (Reyne 1939). The crabs have become so widespread due to the fact

The Coconut Crab

William Owen Murdoch The University Of Nottingham. (2004).

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that they spawn in the sea. The currents can then carry the larvae many miles and propagate new populations on new islands, (Harms 1932). Despite this wide distribution, the Coconut Crab is threatened with extinction. In remote regions, the crab has become a valuable cash crop. It is considered a delicacy and is eaten privately or sold in restaurants. Intensive harvesting in combination with the ease of capture has resulted in its red-listed status and in some countries the crab is already virtually extinct, (Fletcher & Amos 1994). The aim of this site is to promote interest in the Coconut Crab and aid its recovery from the brink of extinction. Click on the links to find out more.

BIOLOGY
ECOLOGY
HABITAT
The Coconut Crab (Birgus latro) is most commonly found on island habitats and generally within 4km from the sea, (Fletcher & Amos 1994). They mainly inhabit dense forest regions but can be abundant in sandy coconut groves, (Grubb 1971). Although rarely observed in person, tracks can be seen on the dunes and the upper shore along the coastline, (Grubb 1971). Many of the Indo-Pacific islands they inhabit are formed from raised coral beds. These limestone structures provide numerous crack, crevices and potholes in which the crabs can hide, (Fletcher & Amos 1994). These potholes are also a good source of fresh rainwater, (Grubb 1971). In regions where rock crevices are not available, the crabs may improvise by using hollowed out tree stumps as a place to hide, (Grub 1971). Alternatively they may dig a burrow or occupy a burrow excavated by a different species of crab, (Grubb 1971). These hides are mainly used during the day or during moulting. Then at night, the crabs become more active and begin to forage. As an important conservation note, Birgus are not normally found in areas cleared for agriculture or livestock, (Fletcher & Amos 1994).

FOOD
Coconut Crabs are omnivorous and show considerable flexibility and complexity in feeding behaviour, (Harms 1932). As their name implies, they are particularly fond of coconuts as a source of food. The crabs have commonly been observed feeding on broken coconuts, (Johnson 1965) and the oils are probably an important source of dietary fat (Harms 1932). However there is much controversy over whether the crabs actually climb the trees and cut the nuts down. Similarly, there is controversy over whether a Coconut Crab can open an undamaged nut. These issues are discussed in the behaviour section. Although an important dietary component, coconuts are not an essential part of the Coconut Crabs diet. The crab is commonly found on islands without any coconut trees at all, (e.g. Christmas Island), (Reyne 1939). In such situations, Birgus must rely more heavily on other sources of food. Pandanus fruits of the screw-pine (Pandanus sp.) appear to be a favourite substitute, (Reyne 1939, Johnson 1965). Once again there is debate over whether the crabs climb up the trees to feed on the fruit but there are many eye witness reports, (Reyne 1939). The crabs have also been reported feeding on a great many other varieties of fruits and nuts. Gibson-Hill (1947) reported that the crabs were, very fond of the fruit Arenga listeri. He noted that if one crab was at the top of the tree feeding, a number of others soon collected underneath for the berries that fell down. Gibson-Hill (1947) also reported that the crabs

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would eat the pith of the same tree as well as the fruit of Terminalia catappa (Tropical Almond), Barringtonia racemosa (Wild Guava) and Artocarpus integrifolia, (Jack Fruit). Harms (1932) reported the nuts of Canarium as an alternative food source. Andrews (1900) stated that the Coconut Crab is particularly fond of the fruits of Arenga listeri, (the Sago Palm). Other observed foods include, Candle Nuts (Aleurites sp.), nutmeg (Myristica fragans) and figs (Ficus sp.), (Reyne 1939). In areas populated by man, there have been reports of the crabs feeding on crops. These include sweet potato, peanuts, melons and papaws, (Reyne 1939). The Coconut Crab is also a scavenger and will eat a variety of dead animal matter. They use their keen sense of smell to detect food sources from a distance, (Johnson 1965). Examples of carrion include washed up fishes (Gibson-Hill 1947) and dead rats (Reyne 1939). They will also eat dead crabs, even of their own kind, (Gibson-Hill 1947). It is also not unknown for the Coconut Crab to take the role of an active predator. Harms (1932) reported one particular hunting method. The crab would seek out a burrow of a landcrab (genus Cardisoma). When the land-crab sticks its large claw out in defence, the Coconut Crab seizes it and hangs on until the Cardisoma claw is cast off. It can then devour the claw at its leisure. Another case of predation has been observed by Andrews (1900). He suggested that the Coconut Crab would occasionally attack injured animals. Reyne (1939) reported a case where chickens were released on an island densely populated by Coconut Crabs. The chickens were reportedly chased and attacked by the resident crabs. In experimental conditions, the crabs have also been seen to kill and eat smaller crab species as well as beetles, (Reyne 1939). Finally, the Coconut Crab has been reported to indulge in active cannibalism if the conditions promote it. Harms (1932) noted that if other sources of animal food were lacking, then the crabs would turn to cannibalism. Reyne (1939) reported that crabs kept together in a small terrarium would tear each other apart. The smaller ones were commonly torn in two and devoured. In a spacious cage with plenty of food, the crabs will not attack each other, (Harms 1932).

PREDATORS
Coconut Crabs evolved in an island environment in which they were the dominant species. However, the local bird populations have always predated small young individuals. In the juvenile stage of a Coconut Crabs life, it undergoes a high frequency of moults, (Harms 1932). These often leave the crab with a temporarily soft shell and such individuals make easy pickings. In recent years, vulnerability has increased due to the introduction of foreign species onto these islands, (Johnson 1965). Rats are thought to attack young individuals. However the two species co-exist in high numbers so the significance is probably quite low, (Johnson 1965). Pigs are also thought to prey on the Coconut Crab and have the potential to attack fairly large individuals, (Johnson 1965). Unfortunately, the most significant predator is man. The crabs are a valuable cash crop and are commonly overexploited. It is this predation pressure that threatens to wipe out the Coconut Crab species.

ALDABRA CASE STUDY


An Ecological Study of the Terrestrial Decapod Crustaceans of Aldabra In the years of 1971 and 1972 Henry G. L. Alexander (a PhD student from the University of London) wrote a thesis on his study of the terrestrial decapods of Aldabra. Aldabra is a sizeable atoll located in the South Western Indian Ocean and is unusual in that it has largely escaped the influences of man. As a result it possesses rich fauna contained within areas of mixed scrub and non-managed coconut plantations. The island is thought to be around 80,000 years old and Coconut Crab colonisation would have been instant due to the nature of their reproductive cycle. Today, Birgus is widely spread throughout the island. Alexander studied a range of different crab species during his study but for the purposes of this website I will make reference mainly to Birgus. Alexander made his observations using a combination of fixed route walks and a comparison between five fixed habitat plots. He found that the Coconut Crab was especially common in the areas with the most coconut trees. In the coconut plantations, they were found to grow to above average size. Burrowing: Crabs that lived in the rocky inland areas tended to take shelter in the deep crevices and potholes. The crabs that lived in sandier areas tended to dig themselves large shallow burrows. These were measured at around 20 to 40cm deep and up to 1.5m long. The burrows were lined with coconut husks fibres and these were often heaped at the entrance. No substitute bedding was used in areas where coconuts were not available. From this, Alexander concluded that the husks were not used as bedding but just the residues of feeding. The burrows are thought to be used for hibernation during ecdysis (Harms 1932) but also as a moist environment to prevent desiccation during the dry season. Alexander noticed a marked increase in burrowing during May. This was a month when food is plentiful and so perhaps the crabs were gorging themselves prior to ecdysis. When the crabs emerged from their burrows, they were found to have enlarged abdomens indicating ecdysis had indeed occurred. Predation: Once in adult form, the crabs had no major predators. The young ones were vulnerable to rats and birds but the adults outgrow these threats. Only 8.6% of the sample population were found to have injuries. These were an indication of intraspecific fighting. Their walking legs were the most frequently damaged body part since they are vulnerable during stamping displays. The carapaces of the injured were often found to have stab marks from the sharpended legs of aggressors. Feeding: The Coconut Crab does not feed exclusively on coconuts. The coconut plantations were very localised on Aldabra while the Coconut Crabs were spread throughout. Alexander observed the crabs regularly eating Pandanus but it seemed to be eaten only when there was nothing superior around. Pandanus, also called screw pine, constitutes a group of tropical shrubs of the family Pandanaceae. They sometimes produce large conelike fruits and it is these on which the crabs were feeding. The crabs have also been observed carrying these fruits back to their burrows for consumption, (Grub 1971).

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Other foods include: Fungal Fruiting Bodies: e.g.

Fimbristylis obtusifolia Eragrostis subaequiglumis Fish on the stand line. Dead Coenobite rugosa (A hermit crab). Cast Birgus limbs from intraspecific fights.

Alexander also observed the Coconut Crab predating Icerya seychellarum westwood and Geograpsus grayi. Coconuts are thought to be the Coconut Crabs favourite food and Alexander spent some time investigating the relationship. Alexander noted that a considerable number of nuts exhibited greater attention to the soft eye end. In view of the fact that this end is often powerfully scented, he suggested that olfaction plays a significant role in locating the best angle of attack. Alexander described the opening of coconuts in some detail:

The soft eye is located by the chelae fingers being arranged over the nut and a firm grip taken on the edge of the eye depressions, the soft eye giving in under such pressure. The tip of the dactylus (usually of the smaller, right chela) is inserted. Pieces of the nut are then broken off until a hole is made large enough to admit the whole cheliped. The crab can now scrape away the coconut flesh using the spoon shaped ridges on the finger tips. Alexander (1976) Pg. 85
Alexander also carried out an experiment in which he enclosed a Coconut Crab in a pen with just water and either, (i) a whole coconut or (ii) a dehusked coconut. He found that no tested crab could open a whole coconut sufficiently. The dehusked ones appeared far easier for them to open. In the wild, the crabs must preferentially seek out damaged coconuts or ones that have slowly been dehusked by crabs over time. Another interesting discovery was that the crabs from coconut tree areas were markedly larger than crabs from areas without coconuts. Alexander used his five habitat plots to test this observation. The results are laid out below: SITE Plot 1 Plot 2 Plot 3 Ile Michel Plot 5 Middle Camp COCONUTS ABUNDANT ? Y Y Y Y N N No. OF BIRGUS WEIGHED 218 702 41 189 322 77 MEAN WEIGHT (g) 592 611 596 895 393 380

The mean weight of the crabs in the coconut abundant plots is 673.5g. The mean weight in the plots where coconuts were not abundant is 386.5g. Therefore, the unavailability of coconuts appears to half the mean weight of the local population. Ile Michel has such a large mean weight because it is inaccessible to local people and so these prize catches are never caught.

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The Ecological Effects of Birgus on the Aldabra Ecosystem: Alexander established that the ecological effects of Birgus could be divided into two interlinked partitions: (1) Transfer of material from the marine environment to the terrestrial environment. (2) Transfer of material from the terrestrial environment to the marine environment. These are summarised on the following diagrams:

Birgus latro may play a number of other ecological roles: Scavenging rotting material may reduce the number of carrion breeding flies. By eating the early forms of leaf litter, they are effectively acting as decomposers. Dragging coconuts about help distribute the coconut seed. Their predation of certain crabs will influence the population size of those prey species and subsequent ecological implications may be present. Young Birgus latro are a food source for birds like the Flightless Rail. The burrowing action of the crabs may help aerate the soil but will also contribute to soil erosion.
Population The most populated habitat plot was plot two. The plot was estimated as 47,600m and contained 350 individuals. This was calculated using capture and release data with the aid of Manly & Parrs (1968) methods of estimating population size. The results equated to a population density of one crab every 136m. Breeding Finally, Alexander carried out a study of the breeding habits of the Coconut Crab. Ovigerous females were observed climbing down the sea cliffs at night until their body was just submerged by each wave wash. It only took two to three waves to wash all the eggs away from under their abdomens. Breeding appeared to occur in accordance with the lunar rhythm. During the months of January to April, there were three egg-laying events. Alexander concluded that these events corresponded to the new moon high tides, but I would beg to differ. Looking at the data it is clear that the main breeding event in February was six days after the new moon. This is relevant my own hypotheses laid out in the My Fieldwork section. In relation to my fieldwork, it is important to note here that Alexander observed Cardisoma (Land Crabs), orientating to the moonlight (the lighter horizon) when on egg shedding migrations. They were attracted to any bright light source during that receptive reproductive period. Interestingly, a similar event was noted in a Coconut Crab undergoing its egg shedding migration. The event was witnessed an hour and a half after dark. The moon was at approximately 20 elevation. The subject was on a coastal rock cliff, about 1m away from the edge where it met the sea. Instead of going straight to the water, the crab continued 12 meters across the cliff top toward the moon. Eventually it met the sea and shed its eggs.

REPRODUCTION & LIFE CYCLE


MATING & SPAWNING
Copulation in Birgus latro takes place exclusively on the land. Sexual maturity is thought to commence at around 5 years of age when the individual is around 25mm in thoracic length (Fletcher 1988b). The females never mate shortly after moulting (Helfman 1977b), probably because their shell is still soft and vulnerable to damage. Schiller et al. (1991) identified the summer period as the season of spawning. However I suspect the spawning season is very specific to an islands individual climate. The fertilisation itself takes place deep in the jungle and it is only the females that migrate to the sea, (Gibson-Hill 1947). Eggs are released into the sea after dark at around 25 to 45 days of gestation, (Shiller et al. 1991). The release date generally coincides with the high tides of that location, (Alexander 1976).

PELAGIC PHASE
Once released into the water, the Coconut Crabs eggs hatch immediately. The larvae that hatch develop through four phases called the Zoeal Larval Stages, (Harms 1932). In total, the transition takes around 2 to 3 weeks, (Fletcher & Amos 1994). The larvae are known to reside in the surface layers of the ocean, (Lavery & Fielder 1995). The position within the water column is maintained by both geotactic and phototactic responses, (Fletcher & Amos 1994). The final larval stage eventually undergoes metamorphosis into a phase denoted as the Amphibious Glaucothoe Stage, (Harms 1932). The glaucothoe finds a gastropod shell and then attempts to climb to shore. The above four diagrams illustrate the morphological developments of the four zoeal larval stages, (in chronological order). 1. 2. 3. 4. 1st Zoeasta Stage (Ventral View) 2nd Zoeasta Stage (Lateral View) 3rd Zoeasta Stage (Lateral View) 4th Zoeasta Stage (Dorsal Swim Position)

Above is a diagram to represent the Amphibious Glaucothoe Stage.

MATURATION
Once the glaucothoe hauls its new shell out of the water, it migrates up the beach to the inter-tidal area. Over time it will migrate further inland, but initially they spend most of their time around the high tidemark. This band of coastline gives them good physical protection within the rubble as well as a ready supply of food from the washed up organic material. This region is also the preferred habitat for numerous hermit crab species. They are easily seen creeping around at dusk. Shell competition is high but there is generally a ready supply of shells from competitor mortality and from the sea. The glaucothoe form inhabits a shell for the first nine months of life (Harms 1932) and then undergoes metamorphosis into a juvenile Coconut Crab. By the time the metamorphosis has finished, they emerge with a cephalothoracic length of around 5mm (Fletcher & Amos 1991). At this stage in their development, the crabs spend most of their time buried underground, only surfacing from time to time, to forage briefly (Held 1963). During this metamorphosis, their abdomens expand and their tergal plates develop. Below is a diagram illustrating the midway stage: When the crabs emerge from their burrow the abdomen is still enlarged and somewhat translucent. The tergal plates have not hardened and are not particularly substantial. I only

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found one such individual during my studies. I suspect that they must undergo a series of initial moults before they can assume the true adult morphology.

MOULTING
Once they reach their adult form, Coconut Crabs become very slow growing indeed. An increase in size occurs with each moult. Moulting decreases in frequency with age. Similarly, the percentage post-moulting size increase, decreases with age. Small crabs of less than 20mm in thoracic length moult around 2 to 3 time per year, (Fletcher & Amos 1991). Those greater than 25mm moult around once per year, (Fletcher & Amos 1991). Very large crabs may moult less than once a year, (Fletcher & Amos 1991). These large crabs may only increase up to 3% per moult, so it takes up to 10 years for them to reach the legal catchable size in Vanuatu, (Fletcher & Amos 1991). Males probably take around 40 years to reach the maximum size of around 4kg and females can grow even slower, (Helfman 1973).

BEHAVIOUR
GENERAL BEHAVIOUR
On inhabited islands the Coconut Crab, Birgus latro, is very shy and rarely observed in its natural surroundings. When lots of people are around, the population become strictly nocturnal, (Reyne 1939). It is only on uninhabited islands that observations are easily made during the day. Between the years of 1897 and 1898, Gibson-Hill observed the behaviour of the coconut crab on Christmas Island. He noted that on Christmas Island, Birgus spends most of its time in the open, and individuals seem to move over fairly wide areas. However, they reside in cracks and crevices when it is hot and dry. Alternatively, they evict other crabs like Gecardoidea from their burrows, (Gibson-Hill 1947). On islands with large population densities, (like Christmas Island), the crabs can be seen collecting together in groups of a dozen or more, (Gibson-Hill 1947). These collections gather around a source of food that they have detected. Gibson-Hill (1947) reported that the crabs were equally as active during the day as during the night. However, it is commonly reported that the crabs are inclined to seek shelter from the heat during the day and are only truly diurnal following rainfall, (Reyne 1939). Andrews, (1900) suggested that a reversal could be initiated if the island became infested with rats. If a large number of rats are active during the night then the Coconut Crabs become preferentially active during the day. Gibson-Hill (1947) also reported that the crabs are very sensitive to the observers presence in the jungle. The observer has to be extremely stealthy and this makes them very difficult to study. On densely populated islands, observation is easier. Andrews (1900) found that simply sitting down in a clear area within the jungle resulted in a sighting within 15 minutes. The undisturbed movements of Birgus were recorded by Grub (1971). He reported that Birgus moves very slowly when not alarmed and emits a continuous clicking sound. In the relaxed state they walk forward, but to flee they reverse this motion and run backwards at considerable speed. Gibson-Hill (1947) described the retreat as an awkward series of jerks. A startled crab attempts to keep running until it can jam its abdomen into a crevice. The next line of defence is to wave the massive chelae, (pincers) to deter the attacker. These chelae are extremely powerful and are lined with little teeth like structures called denticles. They are also lined with little hairs and when these hairs are disturbed, an instant closure results, (Grub 1971). Grub (1971) reported them as being powerful enough to pierce tins but despite all this equipment, they are not very aggressive, (Johnson 1965). Gibson-Hill (1947) suggested that this defence mechanism was more a show of strength. It is most effective

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during sparring matches with other Coconut Crabs. He reported that, two hungry specimens may spar for hours until one elects, and is able, to retreat. Gibson-Hill (1947) also reported the habit of climbing trees. He suggested that when startled, a Coconut Crab could be induced to climb, some four or five feet up the trunk of a tree. Grub (1971) also reported this behaviour and noted that it uses, its long pereiopods with their curved dactyls to grip the tree. On narrow, horizontal branches it reportedly travelled upside down in the manner of a sloth, (Grub 1971). Gibson-Hill (1947) noted that they appear to prefer to ascend the tree headfirst and then at the crown, turn around and descend headfirst. It has often been suggested that this arboreal nature is to obtain food such as coconuts or Pandanus fruits but this is the subject of much debate. Another interesting behaviour is the hoarding of food. Birgus have been commonly reported raiding camps for food and then carrying it back to their holes. This has been reported by Andrews (1900), Alexander (1976), Grub (1971). The observations of Grubb (1971) suggest a degree of intelligence with respect to feeding. He noted that the scent of a Pandanus fruit could attract around ten individuals but often only one large Birgus would have access to the food. Dominant individuals are prone to monopolizing a food source and defend it vigorously. If approached by a competitor, they will run at the aggressor with chelipeds fully raised and extended. The dominance is decided by a brief pushing contest. Grubb (1971) also observed the crabs exhibiting a level of cunning. He noted two individuals attracted to the scent of Pandanus on which a group of tortoises were feeding. The crabs attempted to steal the fruit by creeping around or under the tortoises and then dragging segments away with them. All in all, I find the behaviour of Birgus a fascinating subject and my view is shared by a great many naturalists. Johnson (1965) is particularly keen and states that, On present evidence these creatures show the nearest approach to intelligent behaviour to be found amongst non-social arthropods, or amongst invertebrates in general with the exception of the cephalopod molluscs. To find out more click one of the behavioural sub-topics.

FEEDING
The Coconut Debate It is generally agreed that coconuts are the favourite food of the Coconut Crab. It has even been demonstrated that individuals from islands without coconut trees exhibit a smaller average size, (Alexander 1976). The crabs are commonly seen eating broken coconuts in the coconut groves, (Johnson 1965), but there is great debate over how these coconuts are obtained and opened. Generally, the local people of the Indo-Pacific believe that the crab climbs up the trees in order to cut coconuts down. Then it descends, de-husks and opens intact coconuts. Johnson (1965) interviewed a great many islanders who work on coconut plantations. However, no reliable worker claimed to have actually witnessed the behaviour. Both Gibson-Hill (1947) and Johnson (1965) carried out experiments to test the crabs abilities and they both concluded that the crabs were unable to open a fresh coconut. Johnson (1965) also observed a Coconut Crab attempting to pull open a damaged coconut. The coconut was cracked and the crab was pulling the two halves in opposite directions. Unfortunately, the crabs attempt ended in failure. In his review of the Coconut Crab literature, Reyne (1939) drew attention to the observations of Rock (1916). Rock (1916) carried out a series of experiments, which showed that Birgus could break open a coconut when an initial hole is made in it. He concluded that the crabs

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could only feed on coconuts that were severely damaged by the fall from their tree. I concur that this is probably the most likely explanation. Cannibalism Cannibalism is common in land crabs and it has been suggested that it is just as common in herbivorous species as it is in carnivorous species, (Burggren and McMahon 1988). The Coconut Crab is particularly prone to the behaviour. Helfman (1979) observed a large Birgus feeding on a living smaller individual, which had presumably been caught and dismembered. Helfman (1979) also noted that free ranging Birgus would attack and devour tethered individuals. As a result, captive individuals must be kept separate. Helfman (1979) suggested that this cannibalistic behaviour has induced smaller Birgus to concentrate their activities when the larger Birgus are resting. However, on islands with large population densities, (like Christmas Island), the crabs can be seen collecting together in groups of a dozen or more, (Gibson-Hill 1947). These aggregations are centred on a source of food and if the source is plentiful then brawls are avoided. The theories on cannibalism are not backed up with strong evidence. However one is particularly endearing. Nitrogen sources in diet are considered to be a limiting factor, particularly in herbivorous crabs, (Burggren and McMahon 1988). When population density is high or food availability is low, then low nitrogen availability may stimulate the behaviour of cannibalism. The hypothesis states that, cannibalism of juveniles presents an appropriate feedback loop for population regulation in terrestrial animals with pelagic larvae, (Burggren and McMahon 1988). In a situation where food is scarce, the level of recruitment can still be high because the recruitment may come from a neighbouring island where reproduction is flourishing. To resolve the onslaught of juveniles, the adult crabs on the food-scarce island have to resort to cannibalism. This not only regulates the population at the appropriate level but also provides a source of nitrogen. Cannibalistic adults will not jeopardise their own offspring because their larvae are probably recruited elsewhere. This intraspecific competition is thought to have driven the crabs to occupy habitats further inland, (Burggren and McMahon 1988). This in tern may have driven the evolution of the Coconut Crabs high degree of terrestriality. Avoidance Of Recently Eaten Food Avoidance of recently eaten foods is a common characteristic of generalist feeders. The Coconut Crab is essentially a generalist feeder and is likely to follow the same pattern of preference as its cousins, the Coenobita. These generalist foragers are often confronted by nutritional constraint because their diet is founded on an assortment of nutritionally incomplete food. Such animals have been found to obtain a more nutritionally balanced diet by avoiding food they have recently eaten. By alternating their food preference among different food types, these scavengers receive a more balanced diet. There are two hypotheses to explain how the preference comes about: 1) Nutritional Wisdom Hypothesis The hypothesis essentially states that the negative preference induction is a product of the individuals innate nutritional wisdom. This allows the animal to sense which nutrients its body is lacking. The food containing these nutrients, compose the food it desires, (Waldbauer & Friedman 1991). 2) Malaise Hypothesis The hypothesis infers that an individual will consume a nutrient deficient food type until the consumption of that food induces a metabolic disturbance. It is this disturbance that induces the negative preference and the individual then searches for a new source of food, (Waldbauer & Friedman 1991).

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Although not proven in Birgus latro, this negative preference induction has been illustrated in Coenobita compressus, (Thacker 1996 and Thacker 1998). Thacker (1998) set out to determine which of the two hypotheses were correct in C. compressus. To do so, Thacker investigated the possibility of induced food odour preference. The test subjects were fed a specific food type for a length of time. They were then put in an odour choice chamber to see if they would avoid the odour of that food. This method induced the response effectively. The alternative treatment was to starve the individuals and expose the crabs only to the odours. With no actual consumption of food, a negative preference was not induced. The evidence suggested that food odour preferences are not based on the nutritional content of the food. Instead, the crabs avoid the odour of food they have recently consumed and then randomly seek out new odours. This evidence is in strong support of the Malaise Hypothesis. Finally, by monitoring the growth rates of the individuals, Thacker (1998) found that animals that fed on a single food type grew slower than individuals with a mixed diet. This evidence suggests that the avoidance of recently eaten food in the wild helps scavengers like Birgus, obtain increased growth rates and therefore better survivorship and fecundity.

DRINKING BEHAVIOUR
Reyne (1939) reviewed the literature on the Coconut Crabs habit of drinking. Reyne (1939) determined that drinking was first described by Lister (1888). He quotes Lister (1888) as saying:

I watched one of them at a tide-pool on the shore, apparently replenishing the store of water in the branchial cavities. The movement was similar to that of eating. The end of one of the large chelae was dipped in the water and lifted toward the mouth, where it was touched by one of the palps and the moisture passed on, whilst the chela descend for a further supply. Lister (1888) pg. 512 - 531
Reyne (1939) explained that the same behaviour had been observed by Alcock (1905). Except Alcock (1905) witnessed the crabs drinking fresh water from puddles of collected rainfall. Alexander (1976) observed five Coconut Crabs in a coconut tree drinking from the rainwater that had collected in the leaf axils. Harms (1932) kept a number of Birgus captive and observed their drinking behaviour. He noted that they drunk every 3 to 4 days and the events lasted 5 to 10 minuets. This behaviour was most prevalent shortly before dark. Harms (1932) also noted that as the air became dryer, the frequency of drinking increased. Finally, Gross (1955) described the process of drinking in detail. He noted that the Coconut Crab is able to drink from small puddles without immersing itself. Liquid is picked up by the chelipeds and transferred to the maxillipeds, which in turn pass it to the mouth, (Gross 1955). It can also use the last pair of thoracic appendages to take water from the maxillipeds and pass it to the branchial chamber, (Gross 1955). This is thought to be a means of moistening the respiratory membranes.

MOULTING BEHAVIOUR
Ecdysis is defined by the Oxford Dictionary as, the action or process of shedding an outer skin or integument. This process, otherwise known as moulting is a requirement for growth in animals with an exoskeleton. Before an investigation by Held (1963), the moulting behaviour of Birgus latro was poorly understood. The process had never before been described and freshly moulted samples were hard to come by. Harms (1932) managed to get a photograph of such an individual but Held (1963) failed to do so after several years of study.

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In 1961, Held set out to solve the problem. Held collected a young Birgus from an atoll in the Marshall Islands. The captured individual had a cephalothorax length of 20.8mm. It was carried in a jar to a terrarium in the University of Washington. The crab was given ample food, warmth and humidity. During its first month it remained hidden under a coconut husk during the day. Only at night did it venture out to forage. On the 32nd day it dug a burrow ending in a spherical chamber and sealed the entrance with soil. It continued to forage on a nightly basis for a further two months until on the 86th night the crab did not leave the burrow and remained there for 30 days. When it emerged, the cephalothorax had increased by around 9%. These hidden periods persisted again and the increase in size at each interval was a clear indication of ecdysis. Unfortunately it had not been possible to view the crab during these hidden periods, so Held (1963) put the crab in a new specially designed chamber. As a result, it was possible to view the process during the fourth moult. Held (1963) reported that the ecdysis took place on the 28th day of that hidden period. He described the freshly moulted crab as being pale blue with the colour fading to near white in the proximal areas. In the following days, the colour became much darker and the crab began to devour its shed carapace. It is clear to see that during the inter-moult period, the crab gorges itself on food and puts on a great deal of weight. In contrast, the cephalothorax size does not change during the intermoult periods since it can only change size via ecdysis. The specimen was kept for a total of 463 days, during which the weight increased by 197% and the cephalothorax length increased by 44%. It is important to note that the crab was given a luxurious quantity of food and an array of varieties. Therefore, this rate of moulting may not represent the rate found under natural conditions. Similarly, Held (1963) suggests the possibility that this growth rate is continuous throughout the ageing process and that a crab of 100mm in cephalothorax length is around 5 years old. This estimate is not in agreement with Fletcher & Amos (1994) since they suggest that moulting frequency decreases with age. Even so, the study by Held (1963) was a fascinating insight into the secretive world of Coconut Crab moulting behaviour.

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PHYSIOLOGY
GENERAL MORPHOLOGY & PHYSIOLOGY
Land dwelling decapods are innately different to their marine relatives. The adaptation to life on land has driven them to evolve unique morphological and physiological properties. These properties have been largely driven by the need to adapt feeding mechanisms, locomotion, blood salt regulation, water conservation, respiration, reproduction and sensory equipment.

ANATOMY
Defining Characteristics:
Fatty abdomen tucked under the cephalothorax, (Fletcher & Amos 1991). Abdomen armoured dorsally with calcified Tergal Plates, (Fletcher & Amos 1991). Broad abdomen compared to the morphology of hermit crabs, (Fletcher & Amos 1991). No twist in the abdomen as seen in hermit crabs, (Fletcher & Amos 1991). Branchiostegites are greatly extended with a pronounced droop, (Cameron 1981). Carapace is dorsally flattened, (Alexander 1976). Eyestalks are sub-cylindrical, (Alexander 1976). Massive chelae lined with denticles, (Alexander 1976). Left chelipeds are slightly larger, (Alexander 1976). Abdomen is broad and straight, (Alexander 1976). Acute rosterus, (Alexander 1976). Considerable sexual dimorphism (females being smaller), (Alexander 1976). Colour varies from red brown to a maroon blue. (Blue only occurs when the pigment astaxanthin links to a particular protein. The link denatures on death and colour reverts to red/brown), (Alexander 1976).

AREAS OF INTEREST
The greatest areas of interest in the Coconut Crabs physiology are those that relate to its adaptation to the terrestrial environment. The fundamental adaptations are the crabs ability to breathe air and to regulate their hydro-mineral concentrations. These and other areas of interest are explored in more detail within the listed categories.

VENTILATORY FLOW
Birgus latro has been described as having a true lung and a true circulatory system which pumps oxygenated blood to the systemic regions and returns deoxygenated blood (haemolymph) to the lungs, (Semper 1878).
The branchial chambers are enclosed within lateral extensions of the carapace called branchiostegites (Cameron 1981). In ordinary crabs, these branchial chambers house the gills. In Birgus however, the inner lining has instead been elaborated into a lung, (Cameron 1981). Complex spongy layers of arboreal tufts replace the conventional flat epithelial lining, (Cameron 1981). These tufts protrude a few millimetres into the air space of the modified gill chamber. These tufts are under high blood pressure and are used as an instrument of gaseous exchange in much the same manner as alveoli in mammals. These tree-like structures almost completely fill the branchial cavity. The respiratory membrane is extremely efficient at gaseous exchange since it is very thin (less than 1m thick) and has such a large surface area, (Burggren and McMahon 1988). Electron micrography of this branchiostegal membrane has shown it to be reduced in chitin and cytoplasmic layers, (Storch and Welsch 1984). This reduction reduces the diffusion barrier between the hemolymph (blood) and the air.

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The gills themselves are very small and do not protrude into the lung chamber, (Cameron 1981). It has been noted that there is a general decrease in gill size with increasing terrestriality and this is clearly the case in Birgus, (Harms 1932). These gills have been removed in the past, with no ill effect, (Harms 1932), so it would seem that they no longer contribute to respiration during adulthood. Since the Coconut Crabs lungs are located internally, there is a need for a ventilatory mechanism to move atmospheric air in over the respiratory surface. This is carried out by a pair of blade like structures which are an adaptation of the expodites of the second maxilla, (Burggren and McMahon 1988). These blades are called scaphognathites. They are located at a narrow opening to the anterior end of each branchial chamber. These ventilatory pumps draw air in via a series of oscillations, (Burggren and McMahon 1988). During rest, this scaphognathite pumping is often intermittent, (McDonald 1977). The method is an adaptation of how water is conducted over the gills in marine crabs and the technique is just as effective in air, (Burggren and McMahon 1988). The scaphognathites of Birgus are able to pump almost a litre of gas per kg of body mass per minute, (Cameron and Mecklenburg, 1975) which is equivalent in volume to the water flow in the marine crab, Cancer magister, (McMahon, McDonald and Wood, 1979). Cameron and Mecklenburg, (1973) recorded ventilatory pressure fluctuations in Birgus that could not be solely accounted for by the scaphognathite pump. Harms (1932) suggested that certain sub-carapace muscles had a ventilatory function by creating sudden large changes in branchial volume. This may explain the findings of Cameron and Mecklenburg, (1973).

ACID-BASE REGULATION
Following strenuous activity, it has been shown that a Coconut Crab experiences a pronounced change its acid-base balance, (Smatresk and Cameron 1981). Birgus is not well suited to sprinting and can only achieve high speeds for a short period of time, (Smatresk and Cameron 1981). A physiological study by Smatresk and Cameron (1981) found that the post exercise hemolymph was high in lactate levels. This indicates a high level of anaerobic metabolism during exercise. The resulting acidosis can result in a severe drop in hemolymph pH, which is reflected in the vigour of the pulmonary and cardiac amelioration once at rest. In their experiments, Smatresk and Cameron (1981) subjected the crabs to vigorous exercise and then examined the effects. They found that respiratory acidosis persisted for around fifteen minutes following exercise. By comparison, metabolic acidosis lasted two hours. During this time, the ventilation rate was seven times that of the resting rate. This hyperventilation continued for the entire two hours. Interestingly, Smatresk and Cameron (1981) found that ablation of the gills resulted in an increased ventilatory response to acidosis. In such individuals, the period of respiratory acidosis was extended from fifteen minutes to one hour. The metabolic acidosis was also more severe. It is therefore possible that the apparently redundant gills in Birgus latro do still have a function. It would seem that the gills make at least some contribution to acid-base regulation. In a complimentary publication, Cameron (1981) concludes that the gills could only contribute a very small proportion of acid-base regulation since they were not in contact with any appreciable volume of water, (with which to exchange ions). Instead, he suggests the possibility that the acid levels are reduced via acidification of the carbonates in the shell. This is supported by the fact that a post-exercise rise in blood carbonates has been witnessed in other crab species, (Wood and Randall 1981). Acid + A Carbonate A Salt + Water + Carbon Dioxide

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OXYGEN TRANSPORT
In all decapod crustaceans, carrier molecules known as respiratory pigments facilitate oxygen uptake and transport. In crabs, oxygen is carried on a molecule called hemocyanin, (Burggren and McMahon 1988). In mammalian haemoglobin, the haem groups contain a ferrous iron atom, which is capable of carrying a single oxygen molecule, (Toole & Toole 1995). In crabs, hemocyanin utilises copper instead of iron. The oxygen affinity is affected by how the subunits of the molecule are arranged and this differs from genus to genus and sometimes species to species, (Burggren and McMahon 1988). The oxygen capacity of land crab hemolymph (e.g. Birgus latro), tends to be higher than the capacity of their aquatic counterparts, (Burggren and McMahon 1988). This increased capacity allows the land crabs to pick up more oxygen and transport more oxygen per unit flow of hemolymph. This enables them to reduce the hemolymph flow past the gas exchange sites and thus help reduce water loss experienced during respiration, (Burggren and McMahon 1988).

OSMOTIC REGULATION
As the Coconut Crab became more and more terrestrial, its methods of osmoregulation and ion regulation had to adapt. Osmoregulation in aquatic crabs is driven by the need to balance internal osmotic potential with external (environmental) osmotic potential. Birgus latro does not have this exact problem because it is never fully immersed in water as an adult. However, the aerial environment poses inevitable problems. The osmotic properties of the body fluids are likely to be subject to change depending on the stresses imposed and appropriate responses are required to maintain a tolerable blood concentration. Gross, (1955) investigated this by keeping a test sample of Coconut Crabs in a controlled environment for a number of weeks. Some of the crabs were only allowed access to seawater. Others were only allowed access to fresh water. The rest were allowed access to both. At the end of the test period, the blood concentration was measured. The blood of the specimens exposed only to seawater became more concentrated. The blood of the specimens exposed only to fresh water became more dilute. The blood of the specimens with access to both had a concentration intermediate to the two previous. The blood concentration of the Coconut Crab therefore varies in accordance with the salinity of the water available but when both are readily available, the blood is moderately hypotonic to seawater. Gross (1955) also showed that individuals conditioned to have a low blood concentrations visited the seawater more often than individuals with a normal blood concentration. This suggested that the crabs adjust their body fluid concentration by controlling how much water they drink and how saline it is. The Coconut Crab is well adapted to use this mechanism as it is able to drink from small puddles without immersing itself, (Gross 1955). Liquid is picked up by the chelipeds and transferred to the maxillipeds, which in turn pass it to the mouth, (Gross 1955). It can also use the last pair of thoracic appendages to take water from the maxillipeds and pass it to the branchial chamber, (Gross 1955). This is thought to be a means of moistening the respiratory membranes. The Coconut Crab also has methods of preventing water loss. A reduction in integumental permeability, (Herreid 1969) and behaviour such as burrowing helps them maintain a tolerable blood concentration during times of drought. Gross (1955) also proved that the tissue of Birgus exhibited a tolerance to a wide range of blood concentrations. He noted that individuals with extreme body fluid concentrations suffered no adverse effects in terms of activity. This tolerance is probably very useful in times when either salt or fresh water are unavailable.

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This primitive form of osmoregulation is a clear step towards homeostasis and is in agreement with the argument that Birgus is the most terrestrial crab on earth. However all terrestrial decapods are unable to produce urine which is hyper osmotic to blood, therefore they are prone to losing water in extreme situations, (Harris and Kormanik 1981). For this reason, their ability to osmoregulate is not nearly as well as their mammalian counterparts.

LIPID CONTENT
The Coconut Crab, Birgus latro has been found to have high lipid levels around its organs and this is unusual in decapods of high temperature environments, (Lawrence 1970). An investigation by J. M. Lawrence (1970) discovered that the highest concentration of lipids was contained within the hepatopancreas. Lipids contributed 58% of the organs dry weight. 93% of the lipid component was made up of neutral lipids. These neutral lipids are a good indication that the hepatopancreas acts as a nutrient storage organ. Lawrence also found that the testis of Birgus as well as the thoracic muscle and claw muscle had high levels of nutrient lipids. He did not propose a full explanation for why Birgus in particular, had such high lipid levels. However I suspect that these lipid reserves are relied upon as a source of metabolic water during drought and as a source of nutrition during ecdysis. Decapods from high temperature marine environments do not possess such large lipid deposits, probably because water is never scarce.

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GENETICS
INTRODUCTION
Over the last hundred years, there has been a massive decline in Coconut Crabs numbers throughout their range. In order to provide a long-term solution, it is obvious that a management scheme must be implemented as soon as possible. However, this concept is not without problems itself. The scale of such a scheme cannot be determined without primary information on the population structure. This is where the science of genetics comes into play. The degree of dispersal is the major factor that controls the form of population structure that develops. This has interesting repercussions with the Coconut Crab since its larval stage involves a two to four week marine planktonic period. This is the only way in which the genes can migrate from island to island.

ALTERNATIVE POPULATION STRUCTURES


The population structure of the crabs could fall anywhere within a continuum between two extremes: (i) The entire world population exist as one large homogeneous population. (ii) Distinct sub-populations may exist on each island or small island groups. In the first extreme, there would have to be a free mixing of larvae from all over the world and genetic variation would be very low indeed. The great distances over which ocean currents can carry larvae makes this a plausible argument. In the second extreme, the populations would exist in reproductive isolation. It would only occur if the larvae could only travel short distances and ultimately end up back where they started. Genetic variation between these sub-populations would therefore be quite high. The geographic discontinuity of Birgus makes this a plausible argument.

THE IMPLICATIONS OF POPULATION STRUCTURE


When implementing a management scheme, it is vital to understand the population structure. In the case of the Coconut Crab, there would be two very different management strategies for the two extremes of structure type. If the world population exists as one homogeneous group, then a substantial degree of larval dispersal and mixing will be occurring. Each island receives recruits from other islands. It is possible that an island will get most of its recruits from the population of a neighbouring island if it is downstream of that island, (in terms of ocean currents). This could have serious implications, since if an upstream population becomes extinct, it would result in a severe reduction of recruitment at the downstream island. To sum up this structure scenario, all the populations would be interdependent, each relying on the success of their neighbours. Such a scenario would require a management scheme that incorporated the word population as a whole. Alternatively, if the Coconut Crab exists as distinctive sub-populations, then each population will depend on the successful recruitment of from its own stock. Recruitment would be directly proportional to the reproductive populations in that area. As a result, any management schemes would have to treat these populations on a more personal basis to promote a maintainable ratio between local spawning and recruitment. It is fundamental to maintain each population since it will not be possible to rely on other populations to bolster a declining one.

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GENETIC RESEARCH
(A) Allozyme Electrophoresis: Until recently, there has not been any research carried out to illuminate the correct state of affairs. The scientists leading this pursuit are S. Lavery, D. R. Fielder and C. Mortiz. In 1995, Lavery & Fielder published their research for the Australian Centre For International Agricultural Research. The research was to use allozyme electrophoresis analysis to determine the genetic differentiation between Birgus from different locations. In this technique, the variation is represented by different forms of any one enzyme. These different forms are called allozymes. These allozymes can be distinguished by their slight differences in electrical charge. The variation in charge mirrors the variation in DNA sequencing that codes for the enzyme. Even though previous studies have shown marine decapods to exhibit low genetic variation, Lavery & Fielder hypothesised that Birgus would differ due to its short larval period. Lavery & Fielder aimed to investigate the differences between Birgus groups on: (i) adjacent islands. (ii) different islands within an island group. (iii) different island groups. (iv) islands in the Pacific Ocean and islands in the Indian Ocean. Each of these comparisons was meant to represent one order of magnitude in distance. The primary islands of investigation were those of the Vanuatu Archipelago. Samples were taken from four islands within Vanuatu; Espirito Santo, Hui, Tegua and Loh. The rest of the samples came from the Solomon Islands, Christmas Island, the Cook Islands and Niue. Three hundred specimens were collected and eleven tissues types were examined. The allele frequencies were calculated for crabs from each island and these were then compared to the location data using statistical analysis.

Seventy-six enzyme systems were initially screened and eighty-three different enzyme loci were identified. However only seven of these were polymorphic. The analysis using these

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seven showed that the islands of the Vanuatu Archipelago had genetically homogenous populations of Coconut Crabs. Likewise, there was no significant difference between the Solomon Islands and Vanuatu. However Lavery & Fielder found significant heterogeneity between the Solomon - Vanuatu group and the Cook Islands and Niue. Most significantly, there was a high degree of differentiation between the Pacific and Indian Ocean groupings. It appeared that the enzyme loci for peptidase had two forms, of which one was exclusive to the Indian Ocean and the other to the Pacific. The genetic divergence between the different populations is summarised in the dendrogram below, which illustrates the genetic distances between each of the sites:

The uniformity of the Vanuatu-Solomons group most probably represents a large homogenous population with a great deal of genetic mixing. However, Lavery & Fielder (1995) point out that the there may be significant differences between those populations that were simply not represented by the portions of the genome that was studied. Secondly, the sample size used was too small to say anything for certain. Allozyme investigations are renowned for only illuminating large genetic differences. This is why the difference between the Pacific and Indian Ocean populations was detected. Secondly, it is possible that geographical patterns of allozyme variation are affected by selection, resulting in ambiguous assumptions regarding population structure. To illuminate the finer genetic differences and to provide more solid evidence, there was a need to examine a different type of genetic marker. (B) Mitochondrial DNA: Allozyme research had suggested the possibility of significant genetic variation between Birgus populations (Lavery & Fielder 1995). However it was far from conclusive evidence. In 1996, Lavery et. al. released a paper in the Journal Of Molecular Ecology. Lavery et. al. (1996) set out to evaluate gene flow and identify population units via the analysis of mitochondrial DNA, (mtDNA), polymorphism. The method of sampling was similar to the previous investigation with Birgus taken from, the Solomon Islands, Vanuatu, Niue, the Cook Islands, Papua New Guinea, the Philippines and Japan. In total, 160 individuals were sampled for mtDNA analysis.

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Substantial mtDNA variation was discovered in Birgus using a suite of restriction enzymes. The study provided clear evidence of genetic differentiation between the populations of Birgus in the Pacific Ocean and Indian Ocean. The net divergence between these two populations is only 1.4%. However this far exceeds the divergence between populations of the Pacific, which are separated by an equivalent distance. The only statistically significant differences between populations within the Pacific were between the most peripheral sampling locations of the Cook Islands, Japan and Niue. The central populations of the West Pacific provide no evidence for divergence. Below is a representation of the genetic relationships between the sample populations. This multidimensional scaling plot illustrates just how remote the Indian Ocean, (Christmas Island) population is in terms of genetic variation.

Lavery et. al. (1996) explained these findings as a direct result of larval current interactions. They noted that the larvae have a planktonic stage of 3 to 4 weeks. Since they only have a weak swimming ability, it is certain that their journey is almost entirely current driven. The average current goes at around 11cm per second so on average a Birgus larvae could only travel 350km. This restricts the populations in that they can only be transferred from the Papua New Guinea, to the Solomons, to Vanuatu in a stepping stone manner, (since each is around 350km from the next). However, there are not enough stepping stones to get to the Indian Ocean and thus, that population is separate.

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The mtDNA findings are in close agreement with the previous allozyme findings and therefore these conclusions can be viewed with a fair degree of confidence. As an extra point of interest, it is thought that when allozyme and mtDNA research are in agreement, there is generally an equal dispersal of male and female individuals throughout the populations.

THEORIES OF EVOLUTIONARY HISTORY


Lavery et. al. (1996) used their findings to establish a theory on the evolutionary history of the Coconut Crab. They suggest that Birgus has recently colonised the Pacific islands. This occurred during a previous ice age, during which the sea level dropped. This drop exposed more islands, allowing a stepping-stone spread of Birgus from the Indian Ocean to the Pacific. Since the colonisation of the Pacific, there has been a large increase in population over a prolonged period of time. The rise in sea level has resulted in fewer and fewer connecting islands and thus an increase in genetic diversity. Therefore this strong historical barrier between the Pacific and Indian Ocean populations would appear to be to blame for the extensive diversification between the two.

THE IMPLICATIONS FOR CONSERVATION:


The Vanuatu and Solomon Islands appear to support a genetically uniform stock of Birgus and should therefore require an overall management plan if they are to survive. This can either be viewed as a blessing or a problem. It could be a good thing in that it could be possible to rejuvenate depleted islands with larval stock from neighbouring populations. A pessimistic view is that the total population numbers in the area have already fallen below the essential number required to prevail over normal larval and juvenile mortality and thus keep the region populated. Similarly, it may be that well populated islands depend solely on a neighbour for recruitment and therefore a chain reaction of localised extinctions could begin. Lavery et. al. (1996) have already noted evidence of dwindling recruitment on the well populated island of Santo and fear it may be simply an isolated remnant population which loses its own larval stock to the sea. An obvious solution to this threat may be the initiation of artificial rearing and stocking in these depleted areas. The discovery of these distinct populations has significant implications for such a program. A captive breeding programme could have two possible objectives: (i) To maintain all existing stocks as each population. (ii) To concentrate efforts on only selected islands, resulting in homogenisations. The first objective is ideal since it will maintain the genetic diversity. However, it could be expensive and difficult to co-ordinate. Due to such limitations, the process of artificial rearing will ultimately lead to the loss of genetic diversity, which in turn may lead to the loss of survival traits. It would be wise to identify these survival traits and use them to improve the genes of the stock via selective breeding. Each population may be adapted for its own specific environment, so when they are moved to their captive breeding site, they may fare less well than if they had been left alone. Introducing these alien populations may introduce genes into the local population, which actually lower the general fitness of the population. This may cause the population to rely on being restocked with new individuals. The best compromise is to introduce the breeding concept to the indigenous people who live near each of the populations. Most of the problems associated with captive breeding could be overcome with a well-designed programme. This would enable the local people to harvest

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some of their stock as food, but also ensure the survival of the genetic diversity throughout the two oceans.

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MY FIELDWORK
INTRODUCTION
In the summer of 2003, I travelled to a remote island in Indonesia with Operation Wallacea. The charity organisation runs a series of biological and social science expedition projects designed to underpin the achievement of specific wildlife conservation aims, (Website 13). The expeditions have been running for the last 8 years in a remote corner of the island of Sulawesi. I travelled to Hoga Island, which is part of the Tukangbesi Archipelago in South East Sulawasi. In 1996, the area was declared as the Wakatobi Marine National Park. At 1.39 million hectares it is the second largest marine protected area in Indonesia. It contains 80,000 people, many of whom rely on the environment for their livelihood, (Website 13). It is this conflict of interest that has become the main threat to the local Coconut Crab population. My work with the Coconut Crabs of Hoga Island encompassed both behavioural and ecological studies, which are detailed herewith.

BEHAVIOURAL STUDY
Introduction

Birgus latro is often nicknamed the Robber Crab by indigenous people. The name arises from its habit of carrying off almost any portable article, (Burggren and McMahon 1988). During Alexanders (1976) visit to Aldabra, he noted the wide range of objects removed from camp. These included cooking utensils, cutlery, machetes and a whisky bottle. Gibson-Hill (1947) also noted this behaviour, quoting the loss of cooking-tins, knives and forks. He stated that he once found a pair of crabs fighting over a silver wrist-watch taken from a pile of clothes fifteen yards away.
Gibson-Hill (1947) suggested that this behaviour was related to feeding. He suggested that the crabs could detect food residues on even the cleanest of items and this encouraged them to steal with intent to consume. Local people have suggested that this behaviour is an attempt to steal shiny objects, which are simply to the crabs liking. This notion is also repeated in informal literature, (Website 9) and I hypothesised it may be a loosely viable explanation. Attraction to shiny objects is common in nature. Short-legged Munchkin cats commonly pursue household jewellery and small metal objects, (Website 10). The Colorado Woodrat was hunted in the gold rush era because it was rumoured to hoard gold nuggets, (Website 11). The behaviour is nicknamed miserly behaviour and encompasses the subsequent hoarding of the stolen materials. This common behaviour is not well understood but hypotheses generally orientate towards either feeding or navigation. Blue tits in the UK are renowned for pecking at milk bottle tops and magpies collect similar objects. In the case of the blue tit, the behavior is learnt. A blue tit lands on a bottle top and pecks due to an innate behaviour. When this behavior is rewarded with milk, it is reinforced. Observation by other blue tits continues the trend. A similar case is the behavior exhibited by barracuda, Sphyraena barracuda. Barracuda are known to attack divers wearing shiny metal trinkets because they trigger the same innate feeding response as the shiny scales of prey, (Website 12). An opposing theory is that the attraction to shiny objects is related to navigation. Birds are thought to use the stars and the moon to navigate at night and perhaps their attraction to

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shiny objects is simply the result of confused navigation. This theory was potentially applicable to the Coconut Crab and my project aimed to provide evidence of this. Hypotheses

The Main Hypothesis:


The attraction to shiny objects in Birgus latro is a super normal response to a stimulus that represents a natural navigation queue. The hypothesis suggests that Birgus are attracted to shiny objects since they represent the reflection of the moon on the sea. It is this reflection, which the crabs use to locate the sea during spawning. This conclusion arose form two known observations. Firstly, the female crabs are known to carry their eggs down to the sea when the tide is at its highest, (Whitten, Mustafa and Henderson 1987). This high tide coincides with the full moon which is obviously when the reflection is brightest, (Whitten, Mustafa and Henderson 1987). The second observation was made by Alexander (1976). He noted an interesting event centred on a Coconut Crab undergoing its egg shedding migration. The event was witnessed an hour and a half after dark. The moon was at approximately 20 elevation. The subject was on a coastal rock cliff, about 1m away from the edge where it met the sea. Instead of going straight to the water, the crab continued 12 meters across the cliff top toward the moon. Eventually it met the sea and shed its eggs. This observation suggests that spawning females are indeed attracted to the moon or its reflection. Similarly, Alexander (1976) observed Cardisoma (Land Crabs), orientating to the moonlight (the lighter horizon) when on egg shedding migrations. They were attracted to any bright light source during that receptive reproductive period.

Sub-Hypotheses :
If the main hypothesis is the case, then a number of so-called sub-hypotheses should follow suit: (1) The crabs should only be receptive to shiny objects at the full moon of the lunar cycle. (2) The intensity of attraction over a year should relate to the intensity of spawning in the natural environment (3) Only sexually mature adults should be receptive to shiny objects. (4) Shiny objects should repel sexually immature individuals. Method 30 coconut crabs were captured. The sample included a male and a female of each size category. These were kept in individual hutches over an entire lunar period. Each individual was tested each evening for its response to a shiny object. This was carried out using a Y-chamber. A Y-chamber is a simple choice chamber shaped like the letter Y. The individual is placed at one end of the Y and the lid is sealed. As it walks forward, its path becomes forked and it is forced to choose one of two directions. At the end of one of these, the shiny object was place. The object used was silver tinsel, lightly illuminated by a dull bulb. Three different sized Y-chambers were built. The smallest was half the size of medium sized chamber, which was in turn, half the size of the largest. The size used depended on the size of the crab being tested.

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Problems With This Study Although based on sound observations, the hypotheses were extremely hard to prove. The main problems were related to practicality. The crabs took a long time to acclimatise once placed in the choice chamber. They were very aware of a persons presence and were often simply reluctant to move at all. This made the project time consuming and lacking in sufficient repeats. If left alone, the crabs attention often turned towards escape which generally involved damaging the lid of the chamber. The variations in rain, humidity and temperature made it hard to rule out these influences in activity levels. The crabs seemed to have a highly developed sense of gradient. They preferred to travel up hill so the chamber had to be perfectly flat. Flattening the test area became a time consuming chore. The final problem was the surprising lack of available test subjects. It proved difficult to capture all the desired size and sex cohorts. The absence of large individuals and ovigerous females appeared to be an obvious indication of over hunting by man. This concerning fact, coupled with the problems of the behavioural study prompted me to pursue an assessment of the Birgus population on Hoga Island.

ECOLOGICAL STUDY
The ecological study was prompted by the severe conservation issues that came to light during the behavioural study. The population of Coconut Crabs on Hoga Island was clearly rather low. This study set out to establish just how severe the situation was. An estimation of population density was established using the baited trail method, (see manual for future study). A grid of 26 bait sites was cut into the forest in the La Babas region of Hoga Island. The grid covered an area of approximately 0.017 km2. The trail was baited every day at 3pm and revisited at 7pm to capture feeding crabs. Over a 15-day period, only 27 crabs were caught. This information can be used to calculate catch per unit effort as explained in Fletcher & Amos (1994):

The CPUE on Hoga was 0.07. This level can be considered extremely low, (given that a CPUE can get as high as 5). Most regions of the world have a CPUE lower than 1 since the species is so endangered. Fletcher & Amos (1994) suggest that such sparse populations should not be subjected to harvesting at all. The CPUE can also be used to gain an approximate population density. Fletcher & Amos (1994) specify the following formula:

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The Catchability Coefficient (q) was calculated by Fletcher et al. (1990b). The figure was calculated by obtaining the actual number of crabs on an island and then relating this to the CPUE and the island surface area. From this they obtained a q value of 12,000. Since then, the above formula has been used to produce a reliable estimate of population density on other islands. When applied to my study, the formula suggests that there are 14.3 Coconut Crabs in the 0.017km2 grid. This equates to 841 crabs per km2. This low density is most probably a direct result of over harvesting by the local population. Although few would admit to actually eating the crabs, (since it is illegal), almost all the local people were strangely familiar with how to handle them. The extent of the problem is evident in the graph below. This graph encompasses all the coconut crabs I collected, (including those from the behavioural study). Although some of the decline is due to natural juvenile mortality, it seems likely that the crabs are harvested as soon as they become an edible size. Only a small proportion of the population are within the weight range equivalent to a reproductive adult, (100g +). Unless hunting is reduced dramatically, it would seem that this population will soon cease to contribute to the larvae of the Tukangbesi Archipelago.

IMPROVEMENTS FOR MY PROJECT


The behaviour hypothesis can only be proven on an island with a high density of Coconut Crabs, (CPUE of 3+). I propose that their shy but devious nature requires the enlistment of technological innovation. I would suggest that captive choice chamber tests are not feasible since the crabs tend to destroy their hutches and apparatus. Instead, the observations must be made in the field. I would recommend establishing 40 observation sites along two transects which would dissect a small island right across the north and south axis: Each of the four limbs of the transect would have 10 equally spaced observation points. Each of these points would be monitored with an infrared motion sensitive camera. Each of these is wired to a database onto which movements are recorded.

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The hit zone at each observation point would have a radius of two meters. This would encapsulate a centrally suspended shiny object, which is gently illuminated by a dim bulb. The frequency of crab visitors would be recorded on each day and the system could be left to run for an entire year. The frequency of visits should correlate to the degree of attraction and should therefore vary with lunar cycle and time of the year. For best results, an attempt should be made to capture and release all of the crabs on the island. Marking these crabs will enable the observer to establish whether it is males or females that are visiting the site. Although it would be a costly operation it would certainly find some interesting results. It would be likely to illuminate a variety of secretive Coconut Crab behaviours as well as any attraction to shiny objects that may exist.

POSSIBLE FUTURE PROJECTS


Could the availability of large unnatural (man-made) shells enable Coconut Crabs to delay their metamorphosis into the adult form? Washed up tins and bottle tops are often used by the juveniles instead of gastropod shells. Do these delay the onset of maturity and thus the reproductive rate of these animals? There have been no studies into the effects of water pollution on the larvae of Coconut Crabs. Could the toxicology of anthropogenic leachate and sewage result in a lowered recruitment of these animals? How do the effects of Coconut Lethal Disease influence Coconut Crab populations? Could this disease help deplete local Coconut Crab populations?

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A MANUAL FOR FUTURE STUDY


HANDLING COCONUT CRABS
At the first encounter, handling a Coconut Crab may seem like a daunting task. It is however not half as dangerous as it looks providing the handler adheres to a set of rules. The simplest way to pick up a Coconut Crab is by spreading your hands across the carapace and gripping firmly either side of the cephalothorax. The crab will either attempt to attack or escape depending on its size. The solution to both is to approach it from behind. Their natural flight response is to run backwards, and they are capable of reaching high speeds for a short period of time. By approaching it with your hand behind it, it can only walk forward. However you must be quick otherwise it will turn around and run fast. Secondly, when your hand approaches from behind it will be unable to use its dangerous pincers to attack. As soon as contact is made, press the crab firmly onto the ground. This will not hurt it and will prevent it hurting you. Once the crab is under control you can arrange your fingers so you have a firm grip across the cephalothorax. Once lifted into the air, the crab will not be able to reach you with its claws but instead it will slowly attempt to poke you with its 3rd pair of sharp ended walking legs. If held correctly then they should not be able to reach you with any significant pressure. If they can reach you with a degree of force then it may be necessary to place the crab back on the ground, pressing it firmly until a better position is found. An alternative is to hold the crab upside down. This tends to calm even the most irate of individuals and they will generally just surrender to their fate. The crabs can be particularly troublesome when it comes to taking measurements. Holding them while attempting to measure the carapace is difficult when done alone because when distracted it is easy to lose sight of what the claws are up to. A confident crab handler will control the crab by pressing it to the ground and then use a free hand to hold the front walking legs in such a manner that they pin the claws in place. This then frees the other hand to make the measurement. A solution for the less confident crab handler is to use the crabstick method. The crabstick is simply a long narrow length of wood with deep notches cut into it at 5cm apart. If a crab is particularly irate, it is given the end of the crabstick to pinch with both claws. The bottom segments of the stick are then snapped off and left in the crabs grip. With something to grip hold of, the crab becomes preoccupied and allows you to take whatever measurements you need. The stick is useful because it can be used again and again (important since finding a stick with the right diameter and strength can be time consuming). This can be a particularly useful method when it comes to weighing a crab since unoccupied claws can damage the equipment. Furthermore they are never keen to let go. The crabstick segment must be retrieved in order to remove its weight from that of the crab. The crab will let go of the stick straight away as soon as its abdomen is tightly secured into a corner. This makes the crab feel safe and it will let go almost instantly. Finally, the crabstick is useful if you have captive crabs and you want to move them from one container to another without the hassle of battling to pick them up. A crabs grip on the stick will easily support its own weight so you can use it to carry them to a new location. Once in its new container, the sticks end can just be snapped off and the crab is released with a little bit of the stick still in its grip.

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CONTAINING
Containing a Coconut Crab is a surprisingly challenging experience. They are the true escape artists of the animal world. The captor should be aware that a captive crab can climb out of a wooden container that has vertical sides by pushing off the two faces either side of a corner. They will also grip onto the join between two pieces of wood. One solution is to put a lid on, however even in darkness their instinct is to travel upwards. Overnight, a large individual can push a lid off that has been held in place by three breezeblocks. With additional blocks, they will chip away at the wood over a few days. Their large claws are extremely powerful and they will squeeze through seemingly impossible openings. Finally, if the base of the container is not solid then they will dig their way out. During my study I was outsmarted time and again until I was able to use a disused concrete water tank that had smooth 3m high walls. However, this Alcatraz of containers was breached by the largest inhabitant who cunningly pushed the lid off his individual space and climbed up the thermometer (destroying it in the process). Unfortunately I dont have any simpler suggestions for containment but these criteria will no doubt be useful when it comes to developing new methods of housing.

COCONUT CRAB NECESSITIES


The most important point to note when it comes to keeping the crabs is that they must be kept in solitary confinement. They are primarily solitary animals in the wild and they will attack each other if forced into close proximity. They are also known to be actively cannibalistic. On a lighter note, the crabs obviously need food and water. It is important to provide the crabs with access to both fresh and salt water. The crabs are omnivorous but are though to benefit form the nutrition of coconuts. A balance between coconuts, vegetation and fish will suffice. Do not be surprised if they do not eat. Recently captured crabs are often full after gorging on the bait The Coconut Crab fairs best in a shaded, humid environment. It is important to recreate this in their containment. A container should be kept out of direct sunlight. The lid should be shaded by a spread of palm leaves. They should be secured so that there is a gap between the leaves and the lid of the container. A bowl of water under the leaves should be allowed to evaporate and the leaves trap the humidity. Regular spraying with a hand held plant mister could be used to increase the water content of the air. The more humid the environment, the more active the crabs will become. Even with all the necessities provided, the captured crabs sometimes appear to suffer from a form of depression. After a while, they become notably less resilient in comparison with freshly caught individuals. At the same time, their blue colour seems to become less brilliant and their general activity seems low. For this reason I would recommend that a study avoid the use of captive crabs.

FIELD SAMPLING TECHNIQUE


The field sampling method I used during my study was loosely based on same techniques as the local crab catchers in Indonesia. The technique is known as the baited trail method and is also laid out in Fletcher & Amos book, Stock Assessment of Coconut Crabs. Unfortunately this book was not known to me at the time of my study but interestingly, a great deal of our methods. This page attempts to convey my personal opinions on sampling methodology as well as Fletcher & Amos seven years of experience.

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BAITED TRAIL METHOD
The method is simple: A trail is cut through the jungle during the daytime. Along that trail, coconut baits are placed at fairly equal intervals. The trail is revisited at night and any crabs that are found at the bait sites are caught. The crabs are generally easily caught because they are quite slow moving. Once caught, their details are recorded.

It is sometimes easier to use trails that are already used by the resident crab catchers. These well-defined paths are generally a lot easier to navigate and of course avoid the hard work of trail cutting. However, in order to get fairly straight transects at a location of your choice, it is necessary to cut your own trails. This is of course restricted by the nature of the land in terms of vegetation and topology. On most islands, the Coconut Crab has a low population density. As a result, it is often necessary to target the baited trails at areas where crabs are likely to live. The characteristics of Birgus habitat are laid out below: Dense forest Cracks Crevices Caves Hollow tree stumps 4km from the ocean, (Fletcher & Amos 1994)

On Hoga Island, such habitat was common since the uplifted coral limestone reefs provided numerous potholes and caves. Areas of agriculture are sparsely populated since these holes are often filled in with compost and the crabs have nowhere to hide. In sparsely populated areas, the jungle is generally less dense and bait laying is easier. As a result, I recommend that more bait be laid in the hope of discovering any inhabitancy. A general rule of thumb is to cut trails that take an hour to cut and bait. The length will then be in proportion to the difficulty of navigation. This is scientifically acceptable because the analysis is based on the number of crabs caught per no of baits. Fletcher & Amos recommend the same method and have coined the term, Catch Per Unit Effort. The CPUE has been found to be directly related to the number of crabs in the area, (Fletcher & Amos 1994). When cutting a new trail it is a good idea to make a rough map. My method was to draw a line to represent the proposed trail. At each bait site, take a GPS reading and write these down along the line in the order they are occur. During the afternoon, use the GPS data to obtain the rough distance between each point and revise the map so that it represents these distances. This map will be useful because it is easy to miss bait sites in darkness and it will also stop you getting lost. An additional aid is to paint the bait site number on a local tree trunk. If you do get lost, it will represent a clear reference point on your map.

PREPAIRING THE BAIT


In my study, two forms of coconut bait were used, based on the advice of the local crab catchers. (i) Coconut Quarters Unhusked (green) coconuts are cut into quarters using a parang, (machete) during the morning. These are then turned husk side up and two cuts are made down the centre. This strand is prised out and is used to secure the coconut to a branch at the bait site. This prevents the large crabs removing the whole bait before one reaches the bait site. This

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method is favourable if there is an abundance of coconut trees along the trail since carrying the bait all the way is hard work. (ii) Coconut Dust The coconut flesh is prised out of the shell and chopped into a fine crumbly paste. It is a favourite with the local hunters because their wives do all the preparation and it is far easier to carry on the hunt. However I found it less effective particularly if it had dried out in the sun, (hence the name Coconut Dust)

LAYING THE BAIT


Fletcher & Amos identified two different ways methods of laying bait: (i) Random Baiting Defined as those baits put anywhere along the trail. (ii) Targeted Baiting Defined as, those put within 3m of a ledge, crevice or some other structure likely to house a coconut crab. I also experimented with both these methods but only got results with the latter. I suspected that random baiting failed because the coconut crabs were not straying far from their holes. Fletcher & Amos point out that random baiting is only profitable in areas of dense population. The island I was working on was sparsely populated and thus targeted baiting was required. Perhaps in densely populated regions, there are not enough holes to go round so crabs are more likely to occur at random locations. An additional tip is to ensure the bait is not placed too close to a crab crevice. If it is too close then the crab can retreat easily to a point out of reach. Fletcher & Amos recommend that the bait be placed greater than 30 cm away from the hole. However I would suggest a distance of 60cm, particularly if capturing crabs is a novel experience. It is also important not to place the baits too far from the holes because it may not attempt the crabs out at all.

TIMING THE BAITED TRAIL METHOD


The timing of events in this method is crucial in terms of how many crabs are caught. For this reason it is important that the timing is as consistent as possible. Preparing the bait can be time consuming and should be started in the morning. When cutting a new path, it is advisable to start at around 12.00pm since it will also take a long time. If the baiting is along an established path, then it is possible to set off as late as 3.00pm. This is early enough to let the crabs settle after the afternoons disturbances. Baits should not be put down much earlier than that since it would give the crabs a chance to consume all the bait and leave the site. I found that the most fruitful time to revisit the crabs was just after dark, (around 7.00pm on Hoga). Any later and many of the baits had been finished. With such variable results with time, it is vital to cover the trail at the same time. Fletcher & Amos recommend similar timings but also had the advantage of understanding annual variation in Coconut Crab abundance. They found that during the dry season on Vanuatu (July September), the crabs tended to be less abundant since many were moulting. The dryness also seemed to inhibit normal foraging behaviour. The best catch rates occurred in the wet season, (December February). As a result, it is important to note the time of year when conducting a study and it is probably best to only attempt population estimation during the wet season.

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THE INFLUENCE OF WEATHER


In a similar manner to the variation over the year, localised rain results in increased catches while localised dryness results in reduced catches. In addition to my own observations, Fletcher & Amos noted that too much rain during the walk can result in no crabs emerging that evening.

EQUIPMENT REQUIRED
DAY: Bait Parang (machete) GPS Unit Compass Clip Board

NIGHT: Parang (machete) GPS Unit Compass Head Torch Callipers or Ruler Weighing Scales Crabstick Clip Board The parang is necessary when cutting the trail and also to prepare bait en route. Furthermore it can be an important survival tool should you end up lost. The GPS is somewhat useless without an accurate compass so always take both. Make sure the GPS batteries are fully charged and the unit is secured in a waterproof cover. A head torch is extremely useful at night because it leaves your hands free to clamber over obstacle, make notes and catch Coconut Crabs. The callipers and weighing scale are needed to take the appropriate measurements. If the crabs are particularly heavy, they may need to be taken back to camp to a weighing scale that was not practical to carry. In this case, tie the crab with some rattan or string and take it back to camp. Make sure it is released back at the bait site that evening.

MEASUREMENTS TO RECORD
General weather conditions, (ideally to include humidity & wind since these will effect olfaction). Whether it is a new trail or not. Time the bait was set. Time the bait was revisited. How many crabs at that site. Evidence of crabs come and gone, (evidence of maceration or bait removal). How many other animals are found at the site. Any extra observations, (e.g. Crabs seen en route)

My Indonesian guide was very familiar with the markings found on macerated coconut flesh. He explained that these were often due to rats and other crabs like Hermit Crabs (coenobitids) and Land-Crabs. This was a problem to me because I could not be sure exactly which was which. Fletcher & Amos have produced a simple diagram to illustrate the differences, which should make for accurate recording: (a) Coconut crabs make large incisions in the flesh of the coconut and leave chunks lying around.

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(b) coenobitids take out smaller bites and generally do not leave large chunks on the ground. (c) rats make very neat gnawing marks on the coconut flesh.

MEASURING THE CAPTURED INDIVIDUALS


During my study, I measured the cephalothacic width while Fletcher & Amos measured the cephalothacic length. Previously, Fletcher & Amos had measured other body parts but found that they all correlated well and only one measurement was needed. In effect, it doesnt really matter what is measured as long as the method is consistent. I have come to the conclusion that the cephalothacic length is the best method since damage to the edges of the cephalothorax can cause inaccuracies when measuring the cephalothacic width. Below is a diagram of how to measure the cephalothacic length: Fletcher & Amos made a few extra observations, which I would have noted had I been aware of them. The most useful is their simple method of estimating when the crab last moulted. Fletcher & Amos noted that after a moult, the terminal segments of the walking legs become progressively worn. These dactyls have needle sharp points and numerous hairs when freshly moulted. If they havent moulted for around a year, the points become blunt and the hairs are all worn off.

CAPTURE & RELEASE EXPERIMENTS


When conducting such experiments, it is necessary to mark the carapace with a number. I did this using white paint. It is best to avoid toxic substances since the crabs are likely to eat their own exoskeleton once it is shed. Moulting is a problem for a long-term study because the mark will not last through the moult. Fletcher & Amos devised two methods to overcome this. They experimented with freeze branding and heat branding. Heat branding is believed to last up to six years while freeze branding is less long-term but also less distressing to the animal. It is worth noting however that capture and release experiments are only practical on small islands since the crabs will emigrate from the area if given enough space and time, (Fletcher et al. 1990a)

EXTRA NOTES
When I was studying the Coconut Crab there were two important factors that are often overlooked by the literature. Simple species and sex identification are often not mentioned in the literature because it so basic that it is presumed understood. Species identification is easy when studying adult Birgus because obviously they are so large and morphologically unique. However, young Coconut Crabs that are still occupying a gastropod shell can be difficult to distinguish from other coenobitids. When extremely small, (glaucothoe stage), the easiest way to distinguish the two, is to look for a predominant rostrum which indicates the individual is a Birgus, (Reese & Kinsie 1968). The rostrum is the triangular pointed section, which is predominant between the eyes of a Coconut Crab. Below is a diagram to illustrate the differences between a large adult Coenobita clupeatus and a juvenile Birgus Latro: The differences between the carapaces and abdomens are self-evident. A final point is how to distinguish between male and females. This is very simple and the local crab hunters will probably know this themselves. Females have three feather-like

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pelopods that are tucked under their ventral surface. These leg-like structures are used to hold the eggs in place during brooding.

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CONSERVATION
HISTORY OF MANS' INFLUENCE
The Coconut Crab was first recorded to science by Dampier in 1688 during his visit to Christmas Island, (Gibson-Hill 1947). However, man had influenced the Coconut Crab long before 1688. Some of the earliest human remains have been discovered in Java, Indonesia. The Java Man has been dated to give an age of 2 million years old, which is almost as old as the oldest human remains in Africa, (http://www.aiccusa.org/education/getstarted.htm). As a result, man has shared the Coconut Crabs habitat for a very long time indeed. However, it is only now that the survival of the species is coming into question. The main reason for this is a general human population boom in the last century. As a result, Indonesia is now the worlds 4th most populous nation with around 210 million people, (2001), (http://www.aiccusa.org/education/getstarted.htm). On a number of Indo-Pacific islands, the Coconut Crabs provide an important food and economic resource. It is a popular culinary treat due to its meaty claws, rich fatty abdomen and delicious taste. It is also rumoured to have aphrodisiac qualities in some cultures. The practice of harvesting the population has been sustainable until the recent human population increase. The Coconut Crab has a slow reproductive cycle because it is slow to reach sexual maturity. They are also easily captured, requiring no specialist equipment. As a result, over harvesting has become a serious problem. Increased tourism in some remote Pacific islands has encouraged some restaurants to offer the crab as an exciting addition to the menu. This over-harvesting has been worsened by severe habitat loss. When western man colonised the islands of the Indo-Pacific, they took with them a variety of alien animal species. Rats compete for the Coconut Crabs food and attack young crabs that have recently moulted. Also pigs and feral dogs are known to attack the crabs. The crabs were once top predator on their islands but now their slow reactions are no match for speedily attacking mammals.

CURRENT ISSUES
A recent addition to the population increase in the Indo-Pacific is the destabilisation of the natural balance between the indigenous people and their local wildlife. In Indonesia a large percent of the ancient human populations existed as sea-gypsies. They travelled from site to site exploiting the resources a little and then moving on to a new location allowing the old one to heal. In modern Indonesia, these peoples have been forced to settle permanently by the Indonesian Government. For example, the indigenous West Papuan people have been flooded with an annul increase of 10,000 new families from all over Indonesia. Almost 1 million settlers have overwhelmed the original 900,000 strong indigenous population, (http://www.house.gov/faleomavaega/St-papua.htm). These localised population booms have devastating effects on wildlife. One of the most recent alien animal species to damage the Coconut Crab is the invasion of the Yellow Crazy Ant. The ant is native to Africa and was accidentally transported to Christmas Island around 1915 onwards, (http://eces.org/gallery/000439.php). It is an extremely destructive species and has become widespread throughout the island. Any unsuspecting Coconut Crab that wanders near an infested area is swarmed upon. The socalled super-colony sprays formic acid, which eventually kills the crab. This has knock on effects for the whole ecosystem since the crabs are thought to help distribute the seeds from fruit.

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Another anthropogenic problem for the Coconut Crab is the problem of pollution. In Fiji, pollution from industrial and urban waste, agricultural run-off and coastal development are all helping to degrade the marine environment on which Coconut Crab's reproduction depends, (http://www.wwfpacific.org.fj/pacific_ecoregions_fime_index.htm). I have not been able to find any studies concerning the effects of pollutants on Coconut Crab larvae but I am confident that it has detrimental effects. There is the problem of atmospheric pollution which may ultimately be enhancing the green house effect and causing global warming. If global warming becomes severe, a rise in sea level would wipe out the majority of the islands on which the Coconut Crab depends. Islands that are not yet submerged will simply become more densely populated by humans as a result and the crab may ultimately face extinction. In most parts of their range, the Coconut Crab has either declined dramatically or disappeared altogether, (wells et al, see genetic sheet). The Indian Ocean has been worst hit. The distribution has been largely confined to three populations centred on Aldabra Atoll (Seychelles), Christmas Island and the Chagos Archipelago, (Lavery et al 1996). As a result the Coconut Crab, Birgus latro features on the International Union for Conservation and Natural Resources (IUCN) Red List of Threatened Species since 1996, (http://www.redlist.org/search/details.php?species=2811), (Eldredge 1996). The IUCN has also outlined all the countries the Coconut Crab is known to inhabit:

1. 2. 3. 4. 5. 6. 7. 8. 9. 10. 11. 12. 13. 14. 15. 16. 17. 18. 19. 20. 21. 22. 23. 24. 25. 26. 27. 28. 29. 30. 31. 32. 33.

American Samoa Australia British Indian Ocean Territory Christmas Island Cook Islands Fiji French Polynesia Guam India Indonesia Japan Kiribati Malaysia Marshall Islands Mauritius Micronesia, Federated States of Nauru New Caledonia Niue Northern Mariana Islands Palau Papua New Guinea Philippines Samoa Seychelles Solomon Islands Taiwan, Province of China Tanzania, United Republic of Thailand Tokelau Tonga Tuvalu Vanuatu

The IUCN also make it clear that the Coconut Crab is classed as DD. This stands for Data Deficient. The IUCN describe a taxon as data deficient when,

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there is inadequate information to make a direct, or indirect, assessment of its risk of extinction based on its distribution and/or population status. A taxon in this category may be well studied, and its biology well known, but appropriate data on abundance and/or distribution is lacking. Data Deficient is therefore not a category of threat or Lower Risk. Listing of taxa in this category indicates that more information is required and acknowledges the possibility that future research will show that threatened classification is appropriate. It is important to make positive use of whatever data are available. In many cases great care should be exercised in choosing between DD and threatened status. If the range of a taxon is suspected to be relatively circumscribed, if a considerable period of time has elapsed since the last record of the taxon, threatened status may well be justified.
(http://www.redlist.org/info/categories_criteria1994.html#categories) For this reason, it is clear that an action plan is required.

ACTION PLAN
WHAT ACTION HAS BEEN TAKEN ALREADY
Most of the significant conservation has been in the form of establishing national parks. In the Cook Islands, there are a number of national parks established from which nothing can be taken. However there is no enforcement of this ruling. Similarly in Indonesia, it is illegal to hunt Coconut Crabs but the law is poorly policed. The problem is that many of the islands that house Coconut Crabs are extremely remote. In Indonesia, such islands are so numerous that it is difficult to police them all in the traditional sense. A more local scaled form of policing is required. Some countries have more specific laws on Coconut Crab harvesting. In 1981, the Government of Vanuatu passed the Coconut Crabs Protection Act. The act controls the removal of any Coconut Crab carrying eggs or any Coconut Crab that is less than 9cm in length. In the Commonwealth of the Northern Mariana Islands, the Division of Fish & Wildlife have established a set of rules to govern crab harvesting:

Crab hunters must have a licence to hunt. Certain seasons are closed to hunting. There is a maximum allowable catch. There are limits on the sizes viable for capture.

Once again, enforcement is lacking but the rules are certainly a step in the right direction. The final example of an existing management scheme is that of Aldabra. Aldabra is an island in the Seychelles and has been a UNESCO World Heritage site since 1982. The atoll is an essential sanctuary for Coconut Crabs since it has disappeared from most other islands in the Seychelles, (http://www.unep-wcmc.org/sites/wh/aldabra.html). I consider such isolated conservation to be a problem in itself because it is probable that not all the larvae released from the Aldabra population is returning back to that island. Instead the larvae are being lost to the sea. This is a problem because there is no longer an influx of larvae from other islands. Overtime this could prove to cause population decline and a reduction in the gene pool.

WHAT ACTION STILL NEEDS TO BE TAKEN


To solve problems like the Aldabra scenario, a system of reintroduction should be undertaken as soon as possible. In fact, the Seychelles is considering this already. The present plan is to reintroduce the Coconut Crab to Cousine Island in the central Seychelles with individuals from the island of Cosmoledo in the Aldabra group. Cousine Island is privately owned and

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managed for eco-tourism, (http://www.nfi.org.za/inverts/Southern%20African%20Invertebrates%20Newsletter.htm). Boosting the population in this region is particularly important because the populations of the Indian Ocean are particularly under threat. These individual schemes are well intentioned but they never seem to work quite as well as they hope for. The problem lies with the logistics of policing the policies and the answer lies in the employment and education of local people. Both these requirements call for funding and the source is the potential for ecotourism. A good model for such a scheme is the success of a private initiative called the Chumbe Island Project off the coast of Tanzania, (http://www.unesco.org/csi/wise/wise8e.htm). The project was set up in 1991 and has been managed by a private company. Chumbe Island is a totally protected area and the protection has been backed up by private investors and income from ecotourism. After ten years it became self-sufficient by only using money from ecotourism and now the Coconut Crab began to flourish again. The key to its success was the training and employment of local people to act as park rangers. Education of the local people helped them to view their wildlife as an important source of income. As a result, they see it as a resource to be nurtured rather than exploited and the benefit is evident to all parties involved. The Chumbe Island Project should be seen as a holistic model for the rest of the Coconut Crabs world. It is important to view the conservation of these crabs as a worldwide challenge because of the nature of their reproduction. Each island population has an interdependent relationship with its neighbouring islands. Ocean currents determine the flow of larvae to and from islands and the management on each island must take this into account (see genetics section). For example, if an island depends entirely on a neighbouring island for its supply of larvae then the conservation of the supplying island must come first. Similarly if there is a tit for tat exchange of larvae between two islands, then conservation on both will help keep the gene pool wide. It is therefore obviously necessary to produce an all-encompassing scheme, which is endorsed by full international co-operation. This is of course a mammoth political task and perhaps simply not plausible. Public awareness is the first stage in such a reform and I hope that this website will help make this first stage a reality.

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GLOSSARY
ABLATION - Surgical removal of a body part or tissue. (www.thefreedictionary.com). ACIDOSIS - Abnormally high acidity (excess hydrogen-ion concentration) of the blood and other body tissues. (www.thefreedictionary.com). AGGREGATIONS - Several things grouped together or considered as a whole. (www.thefreedictionary.com). ALLELE - One of two alternate forms of a gene that can have the same locus on homologous chromosomes and are responsible for alternative traits. (www.thefreedictionary.com). ALLOZYME - An isozyme, which differs from other variants of the enzyme as result of an allelic difference. (www.thefreedictionary.com). AMELIORATION - The act of relieving ills and changing for the better. (www.thefreedictionary.com). ANAEROBIC - Living or active in the absence of free oxygen. (www.thefreedictionary.com). ANTHROPOGENIC - Of or relating to the study of the origins and development of human beings. (www.thefreedictionary.com). ARBOREAL - Inhabiting or frequenting trees. (www.hyperdictionary.com). ARTHROPOD - Invertebrate having jointed limbs and a segmented body with an exoskeleton made of chitin. (www.hyperdictionary.com). AXILS - Leaf axils are the part of a leaf that is deep and can hold water. (www.library.thinkquest.org). BRANCHIAL - Branchial cavity (chamber). A chamber limited primarily by the lateral folds of the carapace, the inner surface of which performs a respiratory function. (www.nhm.org). BRANCHIOSTEGITES - The anterior and lateral margins of the carapace extend ventrally on either side forming the walls of the cephalothorax and enclosing the branchial chambers. (www.nhm.org). CEPHALOTHARACIC - Relating to the head and the chest. (www.books.md). CHELA - A structure like a pincer on the limb of a crustacean or other arthropods. (www.hyperdictionary.com). CHELAE - Plural of Chela. CHELIPED - Either of the pair of appendages bearing a chela. (www.infoplease.com). CHITIN - A structural polysaccharide of an amino sugar found in many fungi and in the exoskeletons of all Arthropods. (www.umanitoba.ca/faculties/science/biological_sciences). COENOBITIDS Crustaceans of the family Coenobitatae. CONTINUUM - A continuous non-spatial whole or extent or succession in which no part or portion is distinct of distinguishable from adjacent parts. (www.thefreedictionary.com). CYTOPLASM - The protoplasm of a cell excluding the nucleus. (www.thefreedictionary.com). CYTOPLASMIC - Of or relating to cytoplasm. (www.thefreedictionary.com). DACTYLUS - The claw or terminal joint of a leg of an insect or crustacean. (www.books.md). DECAPOD - Crustaceans characteristically having five pairs of loco motor appendages each joined to a segment of the thorax. (www.hyperdictionary.com). DESSICATION - Dryness resulting from the removal of water. (www.hyperdictionary.com). DENATURED - A changed in nature or natural quality. (www.hyperdictionary.com). DENDROGRAM - Any branching diagram (or tree). (www.science.uts.edu). DENTICLES - Little tooth like projections. (http://www.books.md/D/dic/denticulated.php).

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DIMORPHISM - The existence of two forms of individual within the same animal species (independent of sex differences). (www.hyperdictionary.com). DIURNAL - Belonging to or active during the day. (www.hyperdictionary.com). ECDYSIS - Periodic shedding of the cuticle in arthropods or the outer skin in reptiles. (www.hyperdictionary.com). ELECTROPHORESIS - The motion of charged particles in a colloid under the influence of an electric field; particles with a positive charge go to the cathode and negative to the anode. (www.thefreedictionary.com). ENZYME LOCI - A specific location or site on a chromosome that codes for a particular enzyme. (www.thefreedictionary.com). EPITHELIAL - Of or belonging to the epithelium. (www.hyperdictionary.com). EPITHELIUM - Membranous tissue covering internal organs and other internal surfaces of the body. (www.hyperdictionary.com). EXOSKELETON - The exterior protective or supporting structure or shell of many animals. (www.hyperdictionary.com). FECUNDITY - The quality of something that causes or assists healthy growth; the state of being fertile; capable of producing offspring. (www.hyperdictionary.com). GENERA - Plural of Genus. GENERALIST - Not specialized. (www.dictionary.cambridge.org). GENUS - The second most specific taxonomic level; includes closely related species. Interbreeding between organisms within the same genus can occur. (www.hyperdictionary.com). GEOTACTIC - Moving in response to the earth's gravitational field. (www.fishbase.org). GLAUCOTHOE - The glaucothoe is a transitional stage between the larval and juvenile stages of crabs. (www.oceanexplorer.noaa.gov). HAEMOCYANIN - A substance found in the blood of the octopus, which gives to it its blue colour. (www.thefreedictionary.com). HEMOLYMPH - The blood analogue used by those animals, such as all arthropods and most molluscs that have an open circulatory system. In these animals there is no distinction between blood and interstitial fluid. The liquid fills all of the interior (hemocoel) of the body and surrounds all cells. (www.thefreedictionary.com). HOMEOSTASIS - A term which essentially means "balance" or "equilibrium". Homeostasis is the property of an open system to regulate its internal environment so as to maintain a stable condition, by means of multiple dynamic equilibrium adjustments controlled by interrelated regulation mechanisms. (www.thefreedictionary.com). HEPATO-PANCREAS - A digestive gland in Crustacea, Mollusca, etc., usually called the liver, but different from the liver of vertebrates. (www.thefreedictionary.com). HETEROGENEITY - the quality of being diverse and not comparable in kind. (www.thefreedictionary.com). HOMOGENISATIONS - The act of making something homogeneous or uniform in composition. (www.thefreedictionary.com). HOMOGENOUS - All of the same or similar kind or nature. (www.thefreedictionary.com). HYPER OSMOTIC - (Of a solution) having a higher osmotic pressure than a comparison solution. HYPOTONIC - (Of a solution) having a lower osmotic pressure than a comparison solution. (www.thefreedictionary.com). INTEGUMENT - An outer protective covering such as the skin of an animal or a cuticle or seed coat or rind or shell. (www.thefreedictionary.com).

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INTRASPECIFIC - Arising or occurring within a species; involving the members of one species. (www.hyperdictionary.com). ION - A particle that is electrically charged (positive or negative); an atom or molecule or group that has lost or gained one or more electrons. (www.thefreedictionary.com). ISOZYME - Any of a group of related enzymes that catalyze the same reaction but have different structural, chemical, or immunological characteristics. (www.thefreedictionary.com). LIPIDS - Lipids are fatty acid esters, a class of relatively water-insoluble organic molecules. (www.thefreedictionary.com). MACERATION - Softening due to soaking or steeping; extreme leanness (usually caused by starvation or disease. (www.thefreedictionary.com). MAXILLIPED - One of the mouth appendages of Crustacea, situated next behind the maxill[ae]. Crabs have three pairs, but many of the lower Crustacea have but one pair of them. (www.hyperdictionary.com). MICROGRAPHY - Examination or study by means of the microscope, as of an etched surface of metal to determine its structure. (www.thefreedictionary.com). MITOCHONDRIAL DNA - DNA which is located not in the nucleus of the cell but in the mitochondria. (www.thefreedictionary.com). MORPHOLOGICAL - The form or shape or an organism or part thereof. (www.thefreedictionary.com). MORPHOLOLGY - The branch of biology that deals with the structure of animals and plants. (www.hyperdictionary.com). OMNIVOROUS - Feeding on both plants and animals. (www.hyperdictionary.com). OSMOREGULATION - Osmoregulation is the active regulation of osmotic pressure of body fluids to keep them from becoming too dilute or too concentrated. (www.thefreedictionary.com). OVIGEROUS - Egg-bearing; applied particularly to certain receptacles, as in Crustacea, that retain the eggs after they have been excluded from the formative organs, until they are hatched. (www.books.md). PALPS - One of a set of paired, elongate or cushion-like structures arising from the sides of the head. (www.webref.org). PELEOPOD - One of the abdominal legs of a crustacean. (www.brainydictionary.com). PEREIOPODS - Thoracic appendage used in locomotion; syn., ambulatory leg, walking leg. [McLaughlin, 1980] (www.nhm.org). PEPTIDASE - Any enzyme that catalyzes the splitting of proteins into smaller peptide fractions and amino acids by a process known as proteolysis. (www.thefreedictionary.com). PHOTOTACTIC - Moving in response to light. (www.fishbase.org). PLANKTONIC - Of or relating to plankton. (www.thefreedictionary.com). POLYMORPHISM - The existence of two or more forms of individuals within the same animal species (independent of sex differences). (www.thefreedictionary.com). PROXIMAL - Situated nearest to point of attachment or origin; "the proximal end of a bone". (www.hyperdictionary.com). RESTRICTION ENZYMES - Enzymes that cuts double-stranded DNA. (www.thefreedictionary.com). ROSTERUM A beaklike projection of the anterior part of the head. (www.thefreedictionary.com). SCAPHOGNATHITES - Thin leafike appendage (the exopodite) of the second maxilla of decapod crustaceans. It serves as a pumping organ to draw the water through the gill cavity. (www.thefreedictionary.com). TAXON -Animal or plant group having natural relations. (www.thefreedictionary.com). TERGAL - Of or pertaining to back, or tergum. (www.hyperdictionary.com). TERRARIUM - An enclosure for keeping or raising and observing animals or plants indoors without standing water. (www.gardenweb.com).

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TOPOLOGY - The configuration of a communication network. (www.thefreedictionary.com). TRANSECTS - A term for a technique for estimating population densities of animals in an area. (www.thefreedictionary.com).

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REFERENCES
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Alcock, A. (1905). Catalogue of the Indian Decapod Crustacea (Pt 2, Fasc. I). Calcutta. Andrews, C. W. (1900). A monograph of Christmas Island, (Indian Ocean). British Museum of Natural History, London. Burggren, W. W. and McMahon, B. R. (1988). Biology of the Land Crabs. Cambridge University Press, Cambridge. Cameron, J. N. (1981). Acid-Base Response to Change in CO2 in Two Pacific Crabs: The Coconut Crab, Birgus latro, and the Mangrove Crab, Cardisoma carnifex. J. Exp. Zool. 218: 65-73. Eldredge, L.G. (1996). Birgus latro. In: IUCN 2003. 2003 IUCN Red List of Threatened Species. Fletcher, W. J. (1988b). Growth and recruitment of the coconut crab, Birgus latro L. in Vanuatu. Final report to ACIAR Project 8381. Fletcher, W. J., Brown, I. W. and Fielder, D. R. (1990a). Movement of coconut crabs, Birgus latro, in a rainforest habitat in Vanuatu. Pacific Science 44, 407-416. Fletcher, W. J. and Amos, M. (1994). Stock Assessment of Coconut Crabs. ACIAR, Canberra. Fletcher, W. J., Brown, I.W. and Fielder, D.R. (1999b). Use of standard and inverse Leslie experiments to estimate the density of coconut crabs (Birgus latro L.) in Vanuatu. Fisheries Research 9, 317-324. Gibson-Hill, C. A. (1947). Field notes on the terrestrial crabs. Bill. Raffles Mus. 18: 43-52. Gross, W. J. (1955). Aspects of osmotic regulation in crabs showing the terrestrial habit. Am. Nat. 89: 205-274. Grubb, P. (1971). Ecology of terrestrial decapod crustaceans on Aldabra. Phil. Trans. Roy. Soc. London, Ser. B 260: 411-416 Harms, J. W. (1932). Die realisation von Genen und die consecutive adaptation. II. Birgus latro L. als landkrebs und seine Beziekungen zu den Coenobiten. Z. wiss. Zool. 140: 167-290. Held, E. E. (1963). Moulting behaviour of Birgus latro. Nature 200: 799-800. Helfman, G. S. (1973). Ecology and behaviour of the coconut crab, Birgus latro (L.). Masters thesis, University of Hawaii, Honolulu. Helfman, G. S. (1977b). Copulatory behaviour of the coconut crab or robber crab Birgus latro (L.). Crustaceana 33, 198-202. Johnson, D. S. (1965). Land crabs. J. Malaysian Branch: Roy. Asiatic Soc. 38: 43-66. Lavery, S. and Fielder, D. R. (1995). Genetic Characteristics. In: Brown, I. W. and Fielder, D. R. (eds), the Coconut Crab: aspects of Birgus latro Biology and ecology in Vanuatu. ACIAR Monograph 8, 87-98. Lavery, S., Moritz, C. and Fielder, D. R. (1996). Indo-Pacific population structure and evolutionary history of the coconut crab Birgus latro. Molecular Ecology 5: 557-570. Lawrence, J. M. (1970). Lipid content of the organs of the coconut crab, Birgus latro (L.) (Decapoda, Paguridae). Crustaceana 19: 264-266. Lister, J. J., (1888). On the natural history of Christmas Island (Indian Ocean). Proc. Zool. Soc. London: 512-531. Manly, B. J. F. and M. J. Parr. 1968. A new method of estimating population size, survivorship and birth rate from capture-recapture data. Transactions Society British Entomology. 18: 81-89. McDonald, D. G. (1977). Respiratory physiology of the crab Cancer magister. Ph.D. thesis, University of Calgary, Alberta, Canada.

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McMahon, B. R., McDonald, D. G. and Wood, C. M. (1979). Ventilation, oxygen uptake, and haemolymph oxygen transport following enforced exhaustive activity in the Dungeness crab Cancer magister. J. Comp. Physiol. 128B: 109116. Cameron, J. N. and Mecklenburg, T. A. (1973). Aerial gas exchange in the coconut crab, Birgus latro with some notes on Gecaroidea lalandii. Respir. Physiol. 19: 245-261. Reese, E. S. and Kinsie, R. A. (1968). The larval development of the coconut crab or robber crab Birgus latro (L.) in the laboratory (Anomura, Paguridae). Crustaceana Suppl 2, 117-114. Reyne, A. (1939). On the food habits of the coconut crab (Birgus latro L.), with notes on its distribution. Arch. Neerland. De Zool. 3: 283-320. Rock, J. F. (1916). Palmyra Island with a description of its flora. College of Hawaii, bull. No. 4. Semper, C. (1878). Ueber die Lunge von Birgus latro. Z. Wiss. Zool. 30: 282-287. Schiller, C., Fielder, D. R., Brown, I. W. and Obed, A. (1991). Reproduction, early life history and recruitment. In: Brown, I. W. and Fielder, D. R. (eds), the Coconut Crab: aspects of Birgus latro Biology and ecology in Vanuatu. ACIAR Monograph 8, 13-35. Smatresk, N. J. and Cameron, J. N. (1981). Post-exercise acid-base balance and ventilatory control in Birgus latro, the coconut crab. J. Exp. Zool. 218: 75-82. Storch, V. and Welsch, U. (1984). Electron microscopic observations on the lungs of the coconut crab, Birgus latro (L.) (Crustacea, Decapoda). Zool. Anz. Jena 212: 73-84. Thacker, R.W. (1996). Food choices of land hermit crabs (Coenobita compressus H. Milne Edwards) depend on past experience. Journal of Experimental Marine Biology and Ecology 199: 179-191. Thacker, R.W. (1998). Avoidance of recently eaten foods by land hermit crabs, Coenobita compressus. Animal Behaviour 55: 485-496. Toole, G. & Toole, S. (1995). Understanding Biology for Advanced Level. Third Edition. Stanley Thornes. Cheltenham. Waldbauer, G. P. and Friedman, S. (1991). Insects and dietary self-selection. Ann. Rev. Ent. 36, 43-63. Wood, C. M., and Randall, D. J. (1981). Hemolymph gas transport, acid-base regulation, and anaerobic metabolism during exercise in the land crab (Cardisoma carnifex). J. Exp. Zool. 218: 23-25. WEBSITES (Website 1). At the Dawn of Time. American Indonesian Chamber Of Commerce. http://www.aiccusa.org/education/getstarted.htm. (Accessed 16/02/04) (Website 2). Indonesias Human Rights Violations In Iran Jaya / West Papua. (1998). http://www.house.gov/faleomavaega/St-papua.htm. (Accessed 16/02/04) (Website 3). Species Information: Birgus latro. International Union for Conservation and Natural Resources (IUCN) Red List of Threatened Species. http://www.redlist.org/search/details.php?species=2811 (Accessed 16/02/04) (Website 4). 1994 Categories & Criteria (version 2.3). International Union for Conservation and Natural Resources (IUCN) Red List of Threatened Species. http://www.redlist.org/info/categories_criteria1994.html#categories (Accessed 16/02/04) (Website 5). Species: Yellow crazy ant. Earth Crash Earth Spirit. http://eces.org/gallery/000439.php. (Accessed 21/03/04) (Website 6). WWF Pacific. Pacific Ecoregions In Action. Fiji Islands Marine Ecoregion. The Crossroad of Biodiversity. http://www.wwfpacific.org.fj/pacific_ecoregions_fime_index.htm. (Accessed 21/03/04) (Website 7). Riedmiller, S. (2001). Environment and development in coastal regions and in small islands. Tourists to the rescue on Chumbe Island. (2001). http://www.unesco.org/csi/wise/wise8e.htm. (Accessed 21/03/04) (Website 8). UNEP World Conservation Monitoring Centre. (1995). Protected Areas Programme. World Heritage Sites. http://www.unep-wcmc.org/sites/wh/aldabra.html. (Accessed 21/03/04) (Website 9). Singapore Zoological Gardens Docents. (2000). Coral Reef Creatures. Robber Crab Or Coconut Crab (Birgus latro). http://www.szgdocent.org/ff/f-robcrb.htm. (Accessed 21/03/04)

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(Website 10). Dodman, N. The Pet Place. Hoarding Behaviour in Cats. http://aolsvc.petplace.aol.com/articles/artShow.asp?artID=3467. (Accessed 21/03/04) (Website 11). Gray, M. T. (1998). Colorado's Wildlife Company Spring. (1998). Rodents Gallery. http://wildlife.state.co.us/Colo_Wild_Co/spr1998/gallery.htm. (Accessed 21/03/04) (Website 12). Miami University. (1999). Barracuda Attack. http://jrscience.wcp.muohio.edu/FieldCourses/MarineEcologyArticles/BarracudaAttack.html. (Accessed 21/03/04) (Website 13). Operation Wallacea. What Is Operation Wallacea? http://www.opwall.com/2004%20What%20is%20Operation%20Wallacea.htm. (Accessed 21/03/04) (Website 14). Gerlach, R. (2002). Reintroduction Of Robber Crabs To Cousine Island, Seychelles. Newsletter No. 2 Of The IUCN/SSC Southern African Invertebrates Specialist Group (SAISG). (2002). Reintroduction Of Robber Crabs To Cousine Island, Seychelles. http://www.nfi.org.za/inverts/Southern%20African%20Invertebrates%20Newsletter.htm. (Accessed 21/03/04) (Website 15). Operation Wallacea. http://www.opwall.com. (Accessed 21/03/04) (Website 16). Livestock & Environment Toolbox Home. Ranching or Grassland farming in Tree Crop Plantation in Sub-humid and Humid Zones. (http://lead.virtualcenter.org/en/dec/toolbox/Grazing/LGH3.htm). (Accessed 21/03/04) (Website 17) The University of Hawaii. Hala (Pandanus tectorius). http://www.botany.hawaii.edu/faculty/webb/BishopWeb/hala-1.htm. (Accessed 21/03/04) (Website 18) Jack Fruit Image. http://www.ocean-park.go.jp/park/new/photo/0203ns13.jpg. (Accessed 21/03/04) (Website 19) Cardisoma Image. http://www.mbi.nsysu.edu.tw/~fiddler/figure/fig_crab/carnif_1.jpg. (Accessed 21/03/04) (Website 20) Aldabra Marine Programme. Atol Image. http://www.aldabra.org/. (Accessed 21/03/04) (Website 21) J-Walk & Associates, Inc. Coconut Image. http://www.j-walk.com/digitalimages/molokai/coconut.jpg. (Accessed 21/03/04) (Website 22) Mt. San Antonio College. Coconut Image. http://elearn.mtsac.edu/mcooper/coconut.jpg. (Accessed 21/03/04) (Website 23) Emry's Reef. Flightless Rail Image. http://www.crypticcrab.com/photos/ScubaDiving/Seychelles%2099/TopSide/Land%20Trips/Rail%20Tour/Flightless% 20Rail.jpg. (Accessed 21/03/04) (Website 24) Indian Ocean Diving Acadamy. Coconut Crab Image - http://www.ioda.cx/image/036.jpg. (Accessed 21/03/04) (Website 25) Coconut Crab Image. http://home.doramail.com/arrowhead/crab5.jpg. (Accessed 21/03/04) (Website 26) Coconut Crab Image. http://home.doramail.com/arrowhead/crab2.jpg. (Accessed 21/03/04) (Website 27) Coconut Crab Image. http://home.doramail.com/arrowhead/crab2.jpg. (Accessed 21/03/04) (Website 28) PennState University. Eberly College of Science. Haemocyanin Image. http://research.chem.psu.edu/koerner/denovo2.jpg. (Accessed 21/03/04) (Website 29) The Exploring Travelogue. Exploring the Cook Islands on a motor (and foot). Cook Islands Image. http://www.exploring.nu/Raratonga_Cook_Islands.JPG. (Accessed 21/03/04) (Website 30) Cousine Island Homepage. Cousine Island Image. http://africanadrenalin.co.za/cousine/. (Accessed 21/03/04) (Website 31) Chumbe Island Coral Park Ltd. Chumbe Island Image. http://www.chumbeisland.com/Photo%20Gallery/gallarryframe.htm. (Accessed 21/03/04) (Website 32) The University of Portsmouth. Hoga Island Image. http://www.envf.port.ac.uk/geo/courses/indonesia/Hoga.JPG. (Accessed 21/03/04) (Website 33) CZAPLON ornithological club. Blue Tit Image. http://www.czaplon.most.org.pl/galeria/sniegon/parcae.jpg. (Accessed 21/03/04) (Website 34) Ken Bondy Website. Barracuda Image. http://www.kenbondy.com/images/Caribbean/Barracuda.jpg. (Accessed 21/03/04)

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(Website 35) GROUND WATER SCIENCE. Iron Leaching Image. http://www.groundwatersystems.com/ironleaching.jpg. (Accessed 21/03/04) (Website 36) Lethal Yellowing in Belize. Coconut Lethal Disease Image. http://www.ambergriscaye.com/cocopalms/ly.html. (Accessed 21/03/04)