Evolution and Extinction of Partulidae, Endemic Pacific Island Land Snails

Robert H. Cowie

Abstract

The broad outline of the systematics of the endemic Pacific island land snail family Partulidae has been understood for some time. The family is divided into three genera: Eua has four species, confined to Tonga and Samoa; Samoana has about 23 species, widely but sporadically distributed in Polynesia and the Mariana Islands; Partula has about 100 species, distributed from Belau to the Society Islands. This review integrates this systematic and biogeographic knowledge with work on ecology, population genetics and speciation that has concentrated especially on the recently speciated Partula spp. of Moorea in the Society Islands. Explanations of Moorean diversity (much of which seems unrelated to ecological factors) based on parapatric speciation and the evolution of morph ratio clines in the absence of isolation have predominated, although without incontrovertible support. Unitary explanations are probably not appropriate. Rather little is known of the basic biology of partulids. They are generally arboreal; feed on a wide range of partially decayed and living plant material; and are relatively long-lived, slow reproducing, ovoviviparous, cross- or self-fertilizing hermaphrodites. The phylogenetic and geographical origins of the three genera are unknown. Partula may have evolved, somewhere unknown, from Samoana, which evolved from Eua in the Tonga-Samoa region, this being the region of origin of Eua; but the opposite sequence has also been postulated. The question is unresolved. Origins of the Moorean species are better understood as their inter-relationships are relatively clear. Rather few dispersal events probably took place and the Society Island fauna as a whole may be derived from but two colonization events - first by a Samoana sp. and later by a Partula sp., both of which then speciated in situ - with a few intra-archipelago colonization events taking place subsequently, predominantly in a southwesterly direction from the older to the younger islands. Many of the questions posed by the group may never be answered. Some species, notably those of Moorea, are already extinct in the wild; others are severely threatened. Artificial introductions of both plants and animals, combined with urban and agricultural development, have had significant impacts, but ill-conceived biological control programmes, targeted at the Giant African Snail, Achatina fulica, constitute currently the most serious threat. However, significant areas are still open to research. Analysis of DNA variation, combined with modern ideas of Pacific biogeography, should allow the whole range from the broad origins of the fauna to the detailed evolution within groups of species to be addressed. Some species may yet be relatively secure in the wild and allow field studies, but extraction of DNA from museum specimens provides an exciting opportunity to continue unravelling the evolutionary history of these endangered snails and to contribute further to our understanding of evolutionary processes and the biogeography of the Pacific.