Three new extinct species from the endemic Philippine cloud rat radiation (Rodentia, Muridae, Phloeomyini)

The 18 extant members of the Tribe Phloeomyini, the “cloud rats,” constitute an endemic Philippine radiation of arboreal herbivores that range in size from ca. 18 g to 2.7 kg, most occurring in cloud forest above 1,200 m elevation. Although calibrated phylogenies indicate that the Phloeomyini is estimated to have begun diversifying within the Philippines by ca. 10–11 million years ago, no extinct fossil species have been described, severely limiting our understanding of this distinctive radiation. Our studies of fossil and subfossil small mammal assemblages from the lowland Callao Caves complex in NE Luzon, Philippines, have produced specimens of Phloeomyini that date from ca. 67,000 BP to the Late Holocene (ca. 4,000 to 2,000 BP). We identify three extinct species that we name as new members assigned to the genera Batomys, Carpomys, and Crateromys, distinguished from congeners by body size, distinctive dental and other morphological features, and occupancy of a habitat (lowland forest over limestone) that differs from the high-elevation mossy forest over volcanic soils occupied by their congeners. Batomys cagayanensis n. sp. is known only from two specimens from ca. 67,000 BP; Carpomys dakal n. sp. and Crateromys ballik n. sp. were present from ca. 67,000 BP to the Late Holocene. These add to the species richness and morphological diversity of this endemic Philippine radiation of large folivores, and show specifically that the lowland fauna of small mammals on Luzon was more diverse in the recent past than it is currently, and that Luzon recently supported five species of giant rodents (ca. 1 kg or more). All three occurred contemporaneously with Homo luzonensis, and two, the new Carpomys and Crateromys, persisted until the Late Holocene when multiple exotic mammal species, both domestic and invasive, were introduced to Luzon, and new cultural practices (such as making pottery) became evident, suggesting that modern humans played a role in their extinction.