The dispersal of Halimeda in northern hemisphere mid-latitudes: Palaeobiogeographical insights

The bryopsidalean alga Halimeda gained an important role as carbonate producer in Cenozoic tropical coral reefs and became a significant constituent of the modern Mediterranean seaweed flora. There are, however, open questions at which time the thermophile alga appeared in the cooler Mediterranean Sea and why it is not detected in coral reefs of the modern Persian Gulf. To unravel the biogeography and ecology of Halimeda at its northern margin of distribution, we use fossil Halimeda records of the Central Paratethys/Medditerranean for comparison of the geological, (palaeo)ecological and evolutionary dispersal constraints of the alga in the Miocene and Holocene Persian Gulf. The revealed spatial and temporal distribution patterns of Halimeda in the regions of the Mediterranean and Arabian seas identify water temperature as the major ecological constraint and the extreme Plio-Pleistocene climate changes as the motor for the dispersal and evolution of Halimeda in higher latitudes. Generally, the distribution of tropical species in higher latitudes was related to warm climate intervals during the Neogene. Accordingly, the available (palaeo)biogeographic data implies that the warm-adapted ancestors of the present-day Mediterranean H. tuna population possibly entered the Mediterranean Sea during the mid-Pliocene global warmth and became isolated during subsequent cooling. It also implies that the warm Persian Gulf water is probably unsuitable for the cool-adapted H. discoidea population in the Gulf of Oman and that its tropical ancestors could have reached the Gulf of Oman only during a Pleistocene glacial phase when monsoon-induced upwelling of cold water in the Arabian Sea was reduced and the Persian Gulf fell dry. This example demonstrates the limitation of the actualistic palaeontological approach when using biota at the edges of their distribution range as palaeoclimate proxy.