Colonization and speciation of cave animals in the Philippines

Island-like situation of caves resulted to species isolation while organism's phenotypic plasticity allows the animal to cope with the cave's environment. These conditions eventually lead to organism's speciation through genetic differentiation. Combined morphological and molecular analyses provided insights on the speciation events and colonization of the subterranean ecosystem. Morphological analysis of hypogean species, known as troglobite, and its epigean congeners showed the interesting differences in their characters. Troglobite exhibited cave adaptations such as degenerated eyesight, enlargement or elongation of ambulatory organs, loss of pigmentation and development of other useful organs that favors their survival in the dark cave environment. Molecular clock estimation based on the substitution rate of 0.88% per million years established for 16S rRNA for the grapsid crab genus Sesarma suggested that the troglobitic Sundathelphusa species colonized the cave habitat in Samar Island in the late Miocene epoch and started to diverge from its epigean ancestor ca. 5.92 mya. Interestingly, the five species of the genus Sundathelphusa from Bohol Island comprising of both hypogean and epigean species (S. cavernicola, S. sottoae, S. vediniki, S. urichi and S. boex) occupy a single clade with divergence time from its sister clade ca. 2.58 mya. This phenomenon suggests two possible interpretations of the existence of Bohol species: (1) they belong to a single species with regular genetic flow from their surface relative and that their character differences can be best interpreted as ecophenotypic, or, (2) the speciation event was very rapid and quite recent. Mitochondrial DNA sequences of 430 base pairs of the large subunit rRNA (16S rRNA) revealed the phylogenetic relationships of the genus Sundathelphusa suggesting a multiple colonizations of caves. The speciation events coincided with the timing of the eustatic sea level fluctuation and geologic changes in the Philippine archipelago. This eustatic event and subsequent geologic changes must have influenced the epigean organisms to migrate and invade the subterranean ecosystem. The bizarre feature of hypogean fauna is the result of many years of evolution due to long confinement in the underground ecosystem.