(1) It leaves the egg as a ciliated or non-ciliated free larva.
(2) This larva makes its way on to the gills or into the body of some Mollusc or other host, throws off its epidermis and becomes a Redia or Sporocyst.
(3) In the body cavity of the Redia or Sporocyst numerous tailed larvæ, known as Cercariæ, are developed by a process of internal gemmation.
(4) The Cercariæ pass out of the body of their parent, and out of their host, and become for a short time free. They then pass into a second, usually invertebrate host, and encyst.
(5) If their second host is swallowed by the vertebrate host of the adult of the species, the encysted forms become free, and attain to sexual maturity.
The majority of these stages are simply parts of a complicated metamorphosis, but in the coexistence of larval budding (giving rise to Cercariæ or fresh Rediæ) with true sexual reproduction there is in addition a true alternation of generations.
Polystomeæ. The ova of the Polystomeæ are usually large and not very numerous, and they are in most cases provided with some process for attachment. Some species of Polystomeæ, e.g. Gyrodactylus, are however viviparous. The young leave the egg in a nearly perfect state, and at the utmost undergo a slight metamorphosis and no alternations of generations. Some however (Polystomum, Diplozoon) are provided with temporary cilia, but the number investigated is too small to determine whether ciliation is the rule or the exception. The ciliated larvæ have a short free existence. The cilia are developed on special cells which may be arranged in transverse bands in the same way as in the larvæ of many Chætopods, but are not, in the larvæ at present known, distributed uniformly. When the free larvæ become parasitic the cells with cilia shrink up.
In Polystomum integerrimum, which lives in the urinary bladder of Rana temporaria, the eggs when laid in the spring pass out into the water. The segmentation is complete, and the embryo when hatched is provided with most of the adult organs, but presents certain striking larval characters. It has five rings of ciliated cells. Three of these are placed anteriorly, and are especially developed on the ventral surface, the posterior one being incomplete dorsally; two are placed posteriorly, and are especially developed on the dorsal surface. Anteriorly there is a tuft of cilia.
The larva itself resembles somewhat an adult Gyrodactylus, and is provided (1) with a large posterior disc armed with hooks, and (2) with two pairs of eyes which persist in the adult state. After a certain period of free existence the larva attaches itself to the gills of a tadpole. The rings of ciliated cells shrink up, and some of the six pairs of suckers found in the adult commence to be formed on the posterior disc. When the bladder of the tadpole is developed, the young Polystomum passes down the alimentary tract to the cloaca, and thence to the urinary bladder, where it slowly attains to sexual maturity. When the larva becomes attached to the gills of a very young tadpole, its development is somewhat more rapid in consequence of better nutrition from the more delicate gills. It then reaches its full development in the gill cavity, and, though smaller and provided with differently organised generative organs to the normal form, produces generative products and dies without being transported to the bladder (vide Zeller, Nos. [216] and [217]).
The ova of Diplozoon, a form parasitic on the gills of freshwater fish (Phoxinus, etc.), are provided with a long spiral filament (Zeller, No. [215]). The embryo has five ciliated areas, four lateral and one posterior. The young form is known as Diporpa. Sexual maturity is not attained till two individuals unite permanently together. They unite by the ventral sucker of each of them becoming attached to the dorsal papilla of the other. Subsequently these parts coalesce, and the ventral suckers disappear in the process. Gyrodactylus, parasitic, like Diplozoon, on the gills of freshwater fishes (Gasterosteus, etc.), is remarkable for its mode of reproduction. It is viviparous, producing a single young one at a time, and, what is still more remarkable, the young while still within its parent produces a young one, and this again a young one, so that three generations may be present within the parent. It seems probable that the second and third generations are produced asexually, the generative organs not being developed; while the young Gyrodactylus of the first generation springs from a fertilized ovum (Wagener, No. [214]).