Synapsids are unique in having developed multirooted teeth and complex occlusions. These innovations evolved in at least two lineages of mammaliamorphs (Tritylodontidae and Mammaliaformes). Triassic fossils demonstrate that close to the origins of mammals, mammaliaform precursors were "experimenting" with tooth structure and function, resulting in novel patterns of occlusion. One of the most surprising examples of such adaptations is present in the haramiyidan Glade, which differed from contemporary mammaliaforms in having two rows of cusps on molariform crowns adapted to omnivorous/herbivorous feeding. However, the origin of the multicusped tooth pattern present in haramiyidans has remained enigmatic. Here we describe the earliest-known mandibular fossil of a mammaliaform with double molariform roots and a crown with two rows of cusps from the Late Triassic of Greenland. The crown morphology is intermediate between that of morganucodontans and haramiyidans and suggests the derivation of the multicusped molariforms of haramiyidans from the triconodont molar pattern seen in morganucodontids. Although it is remarkably well documented in the fossil record, the significance of tooth root division in mammaliaforms remains enigmatic. The results of our biomechanical analyses (finite element analysis [FEA]) indicate that teeth with two roots can better withstand stronger mechanical stresses like those resulting from tooth occlusion, than teeth with a single root.

The end-Permian mass extinction constituted the most devastating biotic crisis of the Phanerozoic. Its aftermath was characterized by harsh marine conditions incorporating volcanically induced oceanic warming, widespread anoxia and acidification. Bio-productivity accordingly experienced marked fluctuations. In particular, low palaeolatitude hard substrate communities from shallow seas fringing Western Pangaea and the Tethyan Realm were extremely impoverished, being dominated by monogeneric colonies of filter-feeding microconchid tubeworms. Here we present the first equivalent field data for Boreal hard substrate assemblages from the earliest Triassic (Induan) of East Greenland. This region bordered a discrete bio-realm situated at mid-high palaeolatitude (> 30 degrees N). Nevertheless, hard substrate biotas were compositionally identical to those from elsewhere, with microconchids encrusting Claraia bivalves and algal buildups on the sea floor. Biostratigraphical correlation further shows that Boreal microconchids underwent progressive tube modification and unique taxic diversification concordant with changing habitats over time. We interpret this as a post-extinction recovery and adaptive radiation sequence that mirrored coeval subequatorial faunas, and thus confirms hard substrate ecosystem depletion as a hallmark of the earliest Triassic interval globally.

Marine benthic ecosystems collapsed during the catastrophic end-Permian mass extinction, and subsequently endured a protracted phase of biotic recovery under harsh environmental conditions. In particular, metazoan reef communities almost totally disappeared and were replaced by microbe-dominated mounds during the latest Permian-earliest Triassic. Here we report the stratigraphically oldest exclusively metazoan bioconstructions from earliest Triassic (mid-Induan) strata in East Greenland - these formed within the first ca 300 ka after the Permian-Triassic boundary. Unlike the multitaxic sponge-microbe and bivalve-based buildups recorded from the Early Triassic peri-paleoequatorial Panthalassan and Tethyan margins, the East Greenland bioaccumulations developed within a restricted Boreal mid-paleolatitude seaway, and comprised a monospecific primary framework of microconchid 'lophophorate tubes with shell fragments and phosphatic debris cemented by biogenic calcite. Prostrate growth of the microconchids likely facilitated their accretion into successive sheet-like biostromes and small bioherms. These are associated with a regional paleoenvironmental shift towards well-oxygenated bottom waters, and locally punctuated sedimentation that created a favorable habitat. Although microconchids were both abundant and geographically widespread throughout the earliest Triassic, such buildups formed solely by these metazoans have not been reported from that time frame outside the Boreal Realm. These apparently flourished in the absence of more stable complex communities, and suggest that a locally variable, rather than ubiquitously sequential revival of metazoan bioconstruction activity took place in the immediate aftermath of the end-Permian extinction. However, these may also suggest that ecological recovery of benthic marine ecosystems following the end-Permian mass extinction might have started earlier in higher paleolatitudes.