Palaeobiogeographic implications of Middle Permian brachiopods from Johore (Peninsular Malaysia)

A new Middle Permian locality in northern Johore, Peninsular Malaysia, yields a small-sized, but compositionally unique, brachiopod fauna consisting of eight species: 'Pseudoleptodus' sp., 'Caricula' cf. 'salebrosa' Grant, 'Neochonetes' ('Nongtaia') aff. 'arabicus' (Hudson & Sudbury), 'Karavankina' sp., 'Transennatia' cf. 'insculpta' (Grant), 'Hustedia' sp., 'Orthothetina' sp., and martiniid indet. The first four genera are new records for Malaysia; in particular, the rare taxa 'Pseudoleptodus' and 'Caricula' characterize the fauna. The brachiopods occur together with the ammonoid 'Agathiceras' sp., the nautiloid 'Foordiceras'? sp., bivalves, and crinoid stems. The locality belongs to the East Malaya terrane of the Cathaysian biotic region, but some affinities to species of the Sibumasu province are recognized. The Malaysian forms of 'Pseudoleptodus', 'Caricula' and 'Transennatia' are similar to those of the Ratburi Limestone (southern Thailand). A Roadian–early Wordian age is interpreted for the Johore fauna. The similarity of brachiopods reported here with those from the Ratburi Limestone suggests that there was species interchange or one-way migration between shallow waters of East Malaya and Sibumasu across the main Palaeo-Tethys. The Tethyan seaway between the two terranes must have been narrower than previously interpreted by some authors to allow such faunal traffic during the Roadian–Wordian time period.