starting at the rear of the upper eyelid. Skin very porous, slightly rough on the back and the head, stomach side smooth; lower back of the legs very rough. Back a dark red-brown, rest of the body black." This species was later synonymised with M. betsileo. Description Species of this genus are small, varying in length between . Most Mantella species are sexually dimorphic in size, with females being larger than males. Mantella vary in shape from streamlined to plump/rounded bodies, with skin that is either smooth or granular. They have small, angular heads, with large eyes that are either entirely dark or have lighter coloration around the edge of the iris. Mantella have a very distinct tympanum. The tips or discs of the fingers are slightly enlarged, though those of the climbing mantella (Mantella laevigata) are distinctly larger than in other members of the genus. They have four fingers on each forelimb and five on each hindlimb; some species have webbed digits, while others do not. The tibiotarsal articulation is roughly between the shoulder and the nostrils. Many species of Mantella are similar to the neotropical family Dendrobatidae in their use of aposematism (from Greek ἀπό apo away, σῆμα sema sign), a defense mechanism that uses dramatic coloration to deter potential predators. Coloration and markings vary between species, with combinations of green, red, orange, yellow, blue, brown, white and black. These colorations are often evidence that the specimen produces toxic, pharmacologically active alkaloid secretions. There are significant similarities between a few species of Mantella and Dendrobatidae, notably the golden mantella (Mantella aurantiaca) and the golden poison frog (Phyllobates terribilis). Cowan's mantella (Mantella cowanii) and certain variations of the Harlequin poison frog (Oophaga histrionica) are also very similar in coloration. Most members of the genus also exhibit aposematism on the ventral region, excluding the golden mantella and black-eared mantella (Mantella milotympanum). The venter is normally uniform black, dark grey, or brown and are often marked with blueish or white spots, flecks, or blotches. There are similar blueish to white markings in the form of either spots or a continuous horseshoe-shaped marking on the vocal sac. These characteristics can be used to distinguish between species, such as Baron's mantella (Mantella baroni) and the Malagasy mantella (Mantella madagascariensis), when locality data is unavailable. Mantella show a variety in alkaloid profiles between individual frogs of the Ranomafana region. These same alkaloids have been