A NEW OVOVIVIPAROUS SPECIES OF TECTARIUS (GASTROPODA: LITTORINIDAE) FROM NIUE, SOUTH PACIFIC, WITH A MOLECULAR PHYLOGENY OF THE GENUS

A new ovoviviparous littorinid gastropod, Tectarius (Echininiopsis)niuensis, from Niue, west of the Cook Islands, is described.This is distinguished from the only other ovoviviparous memberof the genus, T. (E.) viviparus (Rosewater, 1982) from the MarianaIslands, here redescribed in detail. The new species is remarkabletor its high-level habitat in the littoral fringe on wave-exposedkarstic limestone cliffs, for its variation in shell shape accordingto tidal level, shell colour polymorphism, calcified operculumand penis with a single mamilliform penial gland. In a cladisticanalysis of morphological characters, including single representativesof each of the three other subgenera of Tectarius (Tectarius,Echininus, Tectininus), these two ovoviviparous species appearas sister-taxa. This is confirmed by a molecular phylogeneticanalysis of the same species, based on the sequence of a portionof the 16S ribosomal RNA mitochondrial gene. Neither analysisunequivocally confirms the monophyly of Tectarius. The divergenceof DNA sequences within Tectarius suggests that the genus arosein the Upper Cretaceous, much earlier than the oldest (UpperEocene) fossils. Only 4 of the 175 species of Littorinidae areknown to be ovoviviparous (with brooding through metamorphosis)and the possible adaptive significance of this type of developmentis discussed. Hitherto, its rarity had been explained by earlyextinction of poorly-dispersed brooding taxa. However, ovoviviparitymay have persisted in Echininiopsis for at least 35 millionyears, and has not precluded colonization of islands 6300kmapart. (Received 17 September 1996; accepted 15 November 1996)