Adaptive strategies and environmental significance of lingulid brachiopods across the late Permian extinction

Climate warming during the late Permian is associated with the most severe mass extinction event of the Phanerozoic, and the expansion of hypoxic and anoxic conditions in shallow shelf settings. It has been hypothesised that wave aeration provided a ‘habitable zone’ in the shallowest environments that allowed the survival and rapid recovery of benthic invertebrates during the Early Triassic. We test this hypothesis by studying the rock and fossil records of the Aggtelek Karst, Hungary. Nearshore settings recorded in the Bódvaszilas Sandstone Formation and units A and D of the Szin Marl Formation are characterised by taxonomically homogenous fossil assemblages of low diversity and low evenness. Ecological and taxonomic recovery in this environmental setting was hampered by persistent environmental stress. This stress is attributed to increased runoff related to climate warming during the Early Triassic that resulted in large salinity fluctuations, increased sedimentation rates and eutrophication that led to seasonal hypoxia and an environment only favourable for opportunistic taxa. In contrast, shoal and mid-ramp settings further offshore are characterised by high diversity faunas with a greater functional complexity. Prior to the late Spathian Tirolites carniolicus Zone, the shelly fossils and trace fossils are limited to settings aerated by wave activity, which supports the habitable zone hypothesis. In the Tirolites carniolicus Zone, however, the oxygen minimum zone retreats offshore and the habitable deeper shelf settings are rapidly colonised by shallow water taxa, evidenced by the highest levels of diversity and bioturbation recorded in the study. Locally, full recovery of marine ecosystems is not recorded until the Illyrian, with the establishment of a sponge reef complex.

A reduction in body size (Lilliput effect) has been repeatedly proposed for many marine organisms in the aftermath of the Permian–Triassic (PT) mass extinction. Specifically-reduced maximum sizes of benthic marine invertebrates have been proposed for the entire Early Triassic. This concept was originally based on observations on Early Triassic gastropods from the western USA basin and the Dolomites (N Italy) and it stimulated subsequent studies on other taxonomic groups. However, only a few studies have tested the validity of the Lilliput effect in gastropods to determine whether the paucity of large-sized gastropods is a genuine signal or the result of a poor fossil record and insufficient sampling. In combination with a review of the literature, we document numerous new, abundant, large-sized gastropods from the Griesbachian outcrops of Greenland and from the Smithian–early Spathian interval in the southwestern USA. We show that large-sized (“Gulliver”) gastropods (i) were present soon after the PT mass extinction, (ii) occurred in various basins, sedimentary facies and environmental contexts (from shallow to deeper settings), and (iii) belong to diverse higher-rank taxa. Focusing on the western USA basin, we investigate areas from which microgastropod shell-beds were previously presented as being typical. However, we show that Gulliver gastropods do occur in the very same areas. Insufficient sampling effort is probably the main reason for the rarity of reports of large Early Triassic gastropods, which is supported by preliminary rarefaction-based simulations. Finally, it appears that the recently documented middle to late Smithian climate shifts and the severe end-Smithian extinction of nekto-pelagic faunas did not reduce maximum shell sizes of gastropods.

A new terrestrial–marine assemblage from the lower beds of a thin outcrop section of the Kockatea Shale in the northern Perth Basin, Western Australia, contains a range of fossil groups, most of which are rare or poorly known from the Lower Triassic of the region. To date, the collection includes spinose acritarchs, organic-cemented agglutinated foraminifera, lingulids, minute bivalves and gastropods, ammonoids, spinicaudatans, insects, austriocaridid crustaceans, actinopterygians, a temnospondyl-like mandible, plant remains, and spores and pollen. Of these groups, the insects, crustaceans and macroplant remains are recorded for the first time from this unit.

Palynomorphs permit correlation to nearby sections where conodonts indicate an early Olenekian (Smithian) age. The locality likely represents the margin of an Early Triassic shallow interior sea with variable estuarine-like water conditions, at the southwestern end of an elongate embayment within the East Gondwana interior rift–sag system preserved along the Western Australian margin. Monospecific spinose acritarch assemblages intertwined with amorphous organic matter may represent phytoplankton blooms that accumulated as mats, and suggest potentially eutrophic surface waters. The assemblage represents a mixure of marine and terrestrial taxa, suggesting variations in water conditions or that fresh/brackish-water and terrestrial organisms were transported from adjacent biotopes. Some of the lower dark shaly beds are dominated by spinicaudatans, likely indicating periods when the depositional water body was ephemeral, isolated, or subjected to other difficult environmental conditions.

The biota of the Kockatea Shale is insufficiently known to estimate biotic diversity and relationships of individual taxa to their Permian progenitors and Triassic successors, but provides a glimpse into a coastal-zone from the interior of eastern Gondwana. Specialist collecting is needed to clarify the taxonomy of many groups, and comparisons to other Lower Triassic sites are required to provide insights into the pattern of biotic decline and recovery at the end-Permian crisis.