Chapter 14. Terrestrial environments.

Among crustaceans, only Amphipoda, Isopoda, and Decapoda have invaded truly terrestrial environments, but only two groups show full adaptations to live on land: the family Talitridae among the Amphipoda and the suborder Oniscidea among the Isopoda. The Talitridae occur primarily in forest leaf litter, but a number of other habitats, including caves, are recorded. Talitrids are important ecological contributors to the litter fauna, often occurring in high densities. Their adaptations to a terrestrial way of life include the retention of the mitten-shaped second gnathopods, a neotenic condition among males; the first article of antenna 2 greatly enlarged and fixed to the side of the head; and enlarged gills and pleopods often reduced, sometimes to vestigial stumps. Talitrids have a skewed world distribution being at their most diverse in New Zealand, Tasmania, and Japan/ Taiwan. They occur in the Caribbean and Central America but are absent from South and North America except as introduced taxa. Their distribution is largely a result of tectonic activity during the past 150 million years and of extinctions during the Tertiary due to increasing aridity of the climate. The Oniscidea (terrestrial isopods) are the only crustaceans that have managed to adapt to almost all habitat types on land and have become the most species-rich suborder of Isopoda. Although monophyly of the Oniscidea is generally accepted, current taxonomy, based almost entirely on morphological characters, needs extensive revision. Terrestrial isopods present a number of unique adaptations to life on land, some of which result from what can be considered as pre-adaptations of ancestral marine isopods, such as egg development in a marsupium, being dorso-ventrally oblate and having a pleopodal respiration. Other crucial adaptations of Oniscidea include the waterconducting system, the structure of their cuticle, and the "covered" type of pleopodal lungs, all of which are responses to the acute problem of desiccation. They are also among the most speciose taxa in caves, some species have even returned to an aquatic life, and a few species have evolved social behavior. Oniscidea are increasingly being used in biogeographical, phylogeographical, ecological, and evolutionary research and can become model organisms for a broad range of biological studies.