Crustacean larvae

Crustaceans may pass through a number of larval and immature stages between hatching from their eggs and reaching their adult form. Each of the stages is separated by a moult, in which the hard exoskeleton is shed to allow the animal to grow. The larvae of crustaceans often bear little resemblance to the adult, and there are still cases where it is not known what larvae will grow into what adults. This is especially true of crustaceans which live as benthic adults (on the sea bed), more-so than where the larvae are planktonic, and thereby easily caught. Crustaceans may pass through a number of larval and immature stages between hatching from their eggs and reaching their adult form. Each of the stages is separated by a moult, in which the hard exoskeleton is shed to allow the animal to grow. The larvae of crustaceans often bear little resemblance to the adult, and there are still cases where it is not known what larvae will grow into what adults. This is especially true of crustaceans which live as benthic adults (on the sea bed), more-so than where the larvae are planktonic, and thereby easily caught. Many crustacean larvae were not immediately recognised as larvae when they were discovered, and were described as new genera and species. The names of these genera have become generalised to cover specific larval stages across wide groups of crustaceans, such as zoea and nauplius. Other terms described forms which are only found in particular groups, such as the glaucothoe of hermit crabs, or the phyllosoma of slipper lobsters and spiny lobsters. At its most complete, a crustacean's life cycle begins with an egg, which is usually fertilised, but may instead be produced by parthenogenesis. This egg hatches into a pre-larva or pre-zoea. Through a series of moults, the young animal then passes through various zoea stages, followed by a megalopa or post-larva. This is followed by metamorphosis into an immature form, which broadly resembles the adult, and after further moults, the adult form is finally reached. Some crustaceans continue to moult as adults, while for others, the development of gonads signals the final moult. Any organs which are absent from the adults do not generally appear in the larvae, although there are a few exceptions, such as the vestige of the fourth pereiopod in the larvae of Lucifer, and some pleopods in certain Anomura and crabs. Also, the Sacculina and other Rhizocephala have a distinctive nauplius larva with its complex body structure, but the adult form lacks many organs due to extreme adaptation to its parasitic life style. Antonie van Leeuwenhoek was the first person to observe the difference between larval crustaceans and the adults when he watched the eggs of Cyclops hatching in 1699. Despite this, and other observations over the following decades, there was controversy among scientists about whether or not metamorphosis occurred in crustaceans, with conflicting observations presented, based on different species, some of which went through a metamorphosis, and some of which did not. This controversy persisted until the 1840s, and the first descriptions of a complete series of larval forms were not published until the 1870s (Sidney Irving Smith on the American lobster in 1873; Georg Ossian Sars on the European lobster in 1875, and Walter Faxon on the shrimp Palaemonetes vulgaris in 1879). The genus name Nauplius was published posthumously by Otto Friedrich Müller in 1785 for animals now known to be the larvae of copepods. The nauplius stage (plural: nauplii) is characterised by the use of the appendages of the head (the antennae) for swimming. The nauplius is also the stage at which a simple, unpaired eye first appears. The eye is known for that reason as the 'naupliar eye', and is often absent in later developmental stages, although it is retained into the adult form in some groups, such as the Notostraca. The genus Zoea was initially described by Louis Augustin Guillaume Bosc in 1802 for an animal now known to be the larva of a crab. The zoea stage (plural: zoeas or zoeae) is characterised by the use of the thoracic appendages for swimming and a large dorsal spine. The post-larva is characterised by the use of abdominal appendages (pleopods) for propulsion. The post-larva is usually similar to the adult form, and so many names have been erected for the stage in different groups. William Elford Leach erected the genus Megalopa in 1813 for a post-larval crab; a shrimp post-larva is called a parva; hermit crab post-larva are called glaucothoe. In the Branchiopoda, the most basal group of crustaceans, there is no metamorphosis; instead, the animal grows through a series of moults, with each moult adding various numbers of segments to the body, but without any dramatic changes in form. Every other crustacean group with free larvae shows a metamorphosis, and this difference in the larvae is thought to reflect 'a fundamental cleavage' of the crustaceans.