Mangrove horseshoe crab

Carcinoscorpius rotundicauda
Conservation status
Data Deficient (IUCN 2.3) [1]
Scientific classification
Kingdom:	Animalia
Phylum:	Arthropoda
Class:	Merostomata
Order:	Xiphosura
Family:	Limulidae
Genus:	Carcinoscorpius Pocock, 1902
Species:	C. rotundicauda
Binomial name
Carcinoscorpius rotundicauda (Latreille, 1802)
Synonyms
Limulus rotundicauda Latreille, 1802

The mangrove horseshoe crab, Carcinoscorpius rotundicauda, is a marine chelicerate arthropod. Despite its name, it is more closely related to spiders and scorpions (all are in the subphylum Chelicerata) than to crabs. It is the only species in the genus Carcinoscorpius. The mangrove horseshoe crab occurs only in Asia and can be found in India, Indonesia, Malaysia, the Philippines, Singapore, Thailand and Hong Kong. There are four extant species of horseshoe crab. The biology, ecology and breeding patterns of C. rotundicauda and the two other Asian horseshoe crab species, Tachypleus gigas and Tachypleus tridentatus, have not been as well documented as those of the North American species Limulus polyphemus. All four extant species of horseshoe crabs are anatomically very similar.

Evolutionary history

Horseshoe crabs are commonly known by biologists around the world as a living fossil because they have remained practically unchanged in terms of shape and size for millions of years. Although their physiology has changed over the years, their typical three piece exoskeleton, consisting of a prosoma, opisthosoma, and telson, has remained since the mid-Paleozoic era.^[2] Fossils of horseshoe crabs that have been dated to over 400 million years ago look almost identical to those species that are still alive today.^[3] The long existence of this body plan suggests its success. It is estimated that the American species of horseshoe crab diverged from the three Indo-Pacific species approximately 135 million years ago.^[2]

Anatomy

The basic body plan of a horseshoe crab consists of three parts: the prosoma, the opisthosoma and the telson. The prosoma is the dome-shaped part at the front. The marginally-spined middle is called the opisthosoma, and the rear extension that looks like a spike is the telson, which is commonly described as the 'tail'.^[4] Contrary to popular belief, the telson is used to turn itself right side up when overturned.^[3] The mangrove horseshoe crab grows up to 40 centimeters in length (including the tail), and its whole body is protected by a hard, dark brown carapace.[4, 5] Uncommon for chelicerates, horseshoe crabs have two compound eyes. The main function of these compound eyes is to find a mate. In addition, they have two median eyes, two rudimentary lateral eyes, and an endoparietal eye on their carapace and two ventral eyes located on the underside by the mouth. Scientists believe the two ventral eyes aid in the orientation of the horseshoe crab when swimming.^[5] Each individual has six pairs of appendages. The first pair, the chelicerae, is used to place food in its mouth. The next pair of legs are the pedipalps, which are used for walking. The rest of the legs are used for locomotion and are known as pusher legs. Located behind their legs are book gills. These gills are used for propulsion to swim and to exchange respiratory gases.^[5]

Distribution and habitat

This species occurs only in Asia around the Indo-West Pacific region where the climate is tropical or subtropical.^[6] These horseshoe crabs can be found to exist throughout the Southeast Asia region in shallow waters with soft, sandy bottoms or extensive mud flats. The mangrove horseshoe crab is benthopelagic, spending most of its life close to or at the bottom of a body of their brackish, swampy water habitat, such as mangroves.^[6] This is the habitat for which it gains its common name: mangrove horseshoe crab. Scientists have studied the distribution of mangrove horseshoe crabs in Hong Kong specifically. The researchers noted their abundance on the beaches of Hong Kong before the sharp decline in population over the past ten years. In the study, they found an uneven distribution of the horseshoe crabs throughout Hong Kong, with a greater abundance found in the western waters. They predict this unevenness is due to the estuarine hydrography in the western waters, influenced by the Pearl River. They also found that T. tridentatus can coexist in the same habitat as the mangrove horseshoe crab.^[7]

Diet

Mangrove horseshoe crabs are selective benthic feeders, feeding mainly on insect larvae, small fish, oligochaetes, small crabs and thin-shelled bivalves.^[8] Lacking jaws, it grinds up the food with bristles on its legs and places it in its mouth using its chelicerae. The ingested food then enters the cuticle-lined oesophagus and then the proventriculus. The proventriculus is made up of a 'crop' and a 'gizzard'. The crop can expand to fit the ingested food, while the gizzard grinds the food into a 'pulp'.^[5] Studies have found that mangrove horseshoe crabs have a strong preference for insect larvae over the other organisms on which it also feeds.^[8]

Mating biology

In the spring, horseshoe crabs migrate from the deeper water to the shallow, muddy areas. Nesting usually follows the cycle of the high tide of the full and new moons. During the mating period, the males will follow and cling to the backs of their potential mates using modified prosomal appendages for long periods of time before the egg-laying has occurred. Horseshoe crab species with low spawning densities and 1:1 sex ratios, such as the mangrove horseshoe crab, are found to be monogamous. In addition, the female does not choose her mate. Males find their female mates with the use of visual and chemoreceptive signals.^[9] Once a mate is found, the female digs a hole and lays the eggs while the male externally fertilizes them. Once the eggs are laid, the male and female head back to the ocean, and the eggs develop on their own.^[9] Their eggs are large, and after a couple weeks, the eggs hatch into miniature versions of the adults.

Mangrove horseshoe crabs in Singapore breed from August to April. Juveniles grow about 33% bigger each time they moult, and it takes the juveniles about five moults to grow from 2 centimetres (0.79 in) to adult size.^[10]

Human impact

Thousands of the horseshoe crabs are killed by local fishermen.^[11] In some areas, the crab is cooked, and the unlaid eggs are eaten. However, some reports show that the crabs are toxic.^[3] In addition, horseshoe crabs are prized for their blue blood, as it is widely used in biomedical sciences for the development of drugs for diseases like mental exhaustion and gastroenteritis. The blood contains a chemical called Limulus amoebocyte lysate (LAL) that can be used to detect pathogens and their endotoxins.^[11]

References

1. ^ World Conservation Monitoring Centre (1996). "Carcinoscorpius rotundicauda". IUCN Red List of Threatened Species. Version 2.3. International Union for Conservation of Nature. http://www.iucnredlist.org/apps/redlist/details/3856. Retrieved April 18, 2010. 
2. ^ ^{a b} John T. Tanacredi, Mark L. Botton & David Smith (2009). Biology and Conservation of Horseshoe Crabs. Dordrecht: Springer. 
3. ^ ^{a b c} Kelvin K. P. Lim, Dennis H. Murphy, T. Morgany, N. Sivasothi, Peter K. L. Ng, B. C. Soong, Hugh T. W. Tan, K. S. Tan & T. K. Tan (2001). "Mangrove horseshoe crab, Carcinoscorpius rotundicauda, family Limulidae". In Peter K. L. Ng & N. Sivasothi. A Guide to Mangroves of Singapore 1. Singapore Science Centre. http://mangrove.nus.edu.sg/guidebooks/text/2076.htm. 
4. ^ Helen M. C. Chiu & Brian Morton (2003). "The morphological differentiation of two horseshoe crab species, Tachypleus tridentatus and Carcinoscorpius rotundicauda (Xiphosura), in Hong Kong with a regional Asian comparison". Journal of Natural History 37 (19): 2369–2382. doi:10.1080/00222930210149753. 
5. ^ ^{a b c} Ecological Research & Development Group (April 23, 2011). "Natural history: the amazing horseshoe". The Horseshoe Crab: Natural History, Anatomy, Conservation and Current Research. http://www.horseshoecrab.org/nh/species.html. Retrieved June 1, 2011. 
6. ^ ^{a b} "Carcinoscorpius rotundicauda, mangrove horseshoe crab". SeaLifeBase. 2010. http://www.sealifebase.org/Summary/SpeciesSummary.php?ID=25. Retrieved April 23, 2011. 
7. ^ Helen M. C. Chiu & Brian Morton (1999). "The distribution of horseshoe crabs (Tachypleus tridentatus and Carcinoscorpius rotundicauda) in Hong Kong". In Brian Morton. Asian Marine Biology. 16. Hong Kong University Press. pp. 185–196. ISBN 9789622095205. http://books.google.co.uk/books?id=GVSREB4L5kIC&pg=PA185. 
8. ^ ^{a b} H. Zhou & Brian Morton (2004). "The diets of juvenile horseshoe crabs, Tachypleus tridentatus and Carcinoscorpius rotundicauda (Xiphosura), from nursery beaches proposed for conservation in Hong Kong". Journal of Natural History 38 (15): 1915–1925. doi:10.1080/0022293031000155377. 
9. ^ ^{a b} Jennifer H. Mattei, Mark A. Beekey, Adam Rudman & Alyssa Woronik (2010). "Reproductive behavior in horseshoe crabs: does density matter?" (PDF). Current Zoology 56 (5): 634–642. http://www.sacredheart.edu/download/3939_current_zoology_final.pdf. 
10. ^ Lesley Cartwright-Taylor, Julian Lee & Chia Chi Hsu (2009). "Population structure and breeding pattern of the mangrove horseshoe crab Carcinoscorpius rotundicauda in Singapore". Aquatic Biology 8: 61–69. doi:10.3354/ab00206. http://www.int-res.com/abstracts/ab/v8/n1/p61-69/. 
11. ^ ^{a b} Bibhuti Pati (June 24, 2008). "Horseshoe crabs galloping towards extinction". Merinews. http://www.merinews.com/article/horseshoe-crabs-galloping-towards-extinction/136265.shtml. Retrieved June 1, 2011.