Clustered, Stacked and Imbricated Large Coastal Rock Clasts on Ludao Island, Southeast Taiwan, and Their Application to Palaeotyphoon Intensity Assessment

This work investigated the characteristics of a boulder field on the exposed south east coast of Ludao Island (Green Island) in southern Taiwan. Although the region regularly experiences seasonal Pacific typhoons, fieldwork on Ludao was prompted following the double-strike of Typhoon Tembin in August 2012, which followed an unusual looping track and was one of the strongest storms to affect the island in recent decades. In Wen Cuen Bay, large limestone and volcanic clasts (103–105 kg) occur both as isolated individuals and also grouped into distinct clusters across the gently-sloping emerged reef platform of Holocene age. Some individuals reach megaclast proportions. Observations revealed limited evidence for the production of new coastal boulders by Typhoon Tembin. However, clustering, stacking and notable imbrication of old large clasts provide evidence for multiple high-energy palaeoevents. Stacking and imbrication are significant depositional features, implying that (partial) lifting by wave transport was responsible. Boulders deposited by Typhoon Tembin suggest that storm produced minimum flow velocities of 3.2–5.1 m/s. This range of minimum flow velocity (MFV) values is lower than the 4.3–13.8 m/s range inferred from the pre-Tembin boulders, which indicates that older storm washovers must have been stronger, judging from their ability to stack and imbricate large clasts. One explanation for high upper values of palaeoevent MFVs is that localized funnelling of water flow through narrow relict channels (inherited spur-and-groove morphology, oriented perpendicular to the modern reef edge) concentrates onshore flow energy into powerful confined jets. Support for this hypothesis is the positioning and train-of-direction of the main imbricated boulder cluster at the landward head of one such feature. Geomorphic controls amplifying wave-driven flow velocities across the emerged Holocene reef mean that a palaeotyphoon origin is sufficient for explaining large clast stacking and imbrication, without the need to invoke a tsunami hypothesis.

Fossil dermal denticles reveal the preexploitation baseline of a Caribbean coral reef shark community

Preexploitation shark baselines and the history of human impact on coral reef–associated shark communities in the Caribbean are tpoorly understood. We recovered shark dermal denticles from mid-Holocene (∼7 ky ago) and modern reef sediments in Bocas del Toro, Caribbean Panama, to reconstruct an empirical shark baseline before major human impact and to quantify how much the modern shark community in the region had shifted from this historical reference point. We found that denticle accumulation rates, a proxy for shark abundance, declined by 71% since the mid-Holocene. All denticle morphotypes, which reflect shark community composition, experienced significant losses, but those morphotypes found on fast-swimming, pelagic sharks (e.g., families Carcharhinidae and Sphyrnidae) declined the most. An analysis of historical records suggested that the steepest decline in shark abundance occurred in the late 20th century, coinciding with the advent of a targeted shark fishery in Panama. Although the disproportionate loss of denticles characterizing pelagic sharks was consistent with overfishing, the large reduction in denticles characterizing demersal species with low commercial value (i.e., the nurse shark Ginglymostoma cirratum) indicated that other stressors could have exacerbated these declines. We demonstrate that the denticle record can reveal changes in shark communities over long ecological timescales, helping to contextualize contemporary abundances and inform shark management and ecology.

Nanocrystals as phenotypic expression of genotypes—An example in coralline red algae

Coralline red algae (CRA) are important ecosystem engineers in the world’s oceans. They play key roles as primary food source and carbonate producers in marine habitats. CRA are also vital for modern reef systems where they act as substrate for coral growth and stabilizers of reef frameworks. However, morphotaxonomic identification of these important marine organisms is hampered by the fact that morphological concepts used for their classification do not correspond to molecular data. We present the first analysis of nanoscale features in calcified cell walls of CRA in a globally distributed sample set. We use new morphological traits based on these cell wall ultrastructures to construct an independent morphological phyletic tree that shows a promising congruency with existing CRA molecular phylogenies. Our results highlight cellular ultrastructures as a tool to define the phenotypic expression of genotypic information showing their potential to unify morphology with molecular phylogeny.

Prey fractionation in the Archaeocyatha and its implication for the ecology of the first animal reef systems

Archaeocyaths are the most abundant sponges from the Cambrian period, having formed the first animal reef communities more than 500 million years ago. The Archaeocyatha are index fossils for correlating rocks of similar ages globally because of their abundance, extensive geographic distribution, detailed anatomy, and well-established taxonomy. Their ecological significance remains incompletely explored, yet they are known to have strongly competitively interacted, unlike modern sponges. This study examines the feeding ecology of the fossil remains of Siberian archaeocyath assemblages. As suspension feeders, archaeocyaths filtered plankton from the water column through pores in their outer wall. Here we outline a new method to estimate the limit on the upper size of plankton that could be consumed by an archaeocyath during life. The archaeocyaths examined were predominantly feeding on nanoplankton and microplankton such as phytoplankton and protozooplankton. Size-frequency distributions of pore sizes from six different Siberian archaeocyath assemblages, ranging from Tommotian to Botoman in age, reveal significantly different upper limits to the prey consumed at each locality. Some of the assemblages contain specimens that could have fed on larger organisms extending into the mesoplankton, including micro-invertebrates as a possible food resource. These results show that during the establishment of the first animal reef systems, prey partitioning was established as a way of reducing competition. This method has applicability for understanding the construction and the functioning of the first reef systems, as well as helping to understand modern reef systems and their development though time and space.

Freshwater input, upwelling, and the evolution of Caribbean coastal ecosystems during formation of the Isthmus of Panama

Caribbean biota underwent major ecological and evolutionary transformation in the Pliocene–Pleistocene, but a lack of detailed paleoenvironmental reconstruction prevents thorough resolution of cause and effect. We quantify levels of upwelling and freshwater input into Caribbean coastal shelf ecosystems over the last ∼6 m.y. with >3300 stable isotope measurements from 74 fossil serially sampled gastropods by normalizing δ18O values to open-ocean δ18O from planktonic foraminifera. We find that the influence of Pacific-like upwelling in the southwestern Caribbean was low after 4.25 Ma but coastal ecosystems were heavily influenced by seasonal freshening until ca. 2.5 Ma, after which time low-freshwater conditions were established. The origination of modern oligotrophic coastal conditions was therefore a result of oceanographic change causing declining upwelling, and declining nutrients from terrestrial sources. We speculate that a southward shift of the Intertropical Convergence Zone, associated with Northern Hemisphere glaciation, reduced rainfall and terrestrial nutrient input and contributed to biotic turnover in the southwestern Caribbean, including the proliferation of modern reef communities.

Photosymbiosis and the expansion of shallow-water corals

Roughly 240 million years ago (Ma), scleractinian corals rapidly expanded and diversified across shallow marine environments. The main driver behind this evolution is uncertain, but the ecological success of modern reef-building corals is attributed to their nutritional symbiosis with photosynthesizing dinoflagellate algae. We show that a suite of exceptionally preserved Late Triassic (ca. 212 Ma) coral skeletons from Antalya (Turkey) have microstructures, carbonate13C/12C and18O/16O, and intracrystalline skeletal organic matter15N/14N all indicating symbiosis. This includes species with growth forms conventionally considered asymbiotic. The nitrogen isotopes further suggest that their Tethys Sea habitat was a nutrient-poor, low-productivity marine environment in which photosymbiosis would be highly advantageous. Thus, coral-dinoflagellate symbiosis was likely a key driver in the evolution and expansion of shallow-water scleractinians.

Two waves of colonization straddling the K–Pg boundary formed the modern reef fish fauna

Living reef fishes are one of the most diverse vertebrate assemblages on Earth. Despite its prominence and ecological importance, the origins and assembly of the reef fish fauna is poorly described. A patchy fossil record suggests that the major colonization of reef habitats must have occurred in the Late Cretaceous and early Palaeogene, with the earliest known modern fossil coral reef fish assemblage dated to 50 Ma. Using a phylogenetic approach, we analysed the early evolutionary dynamics of modern reef fishes. We find that reef lineages successively colonized reef habitats throughout the Late Cretaceous and early Palaeogene. Two waves of invasion were accompanied by increasing morphological convergence: one in the Late Cretaceous from 90 to 72 Ma and the other immediately following the end-Cretaceous mass extinction. The surge in reef invasions after the Cretaceous–Palaeogene boundary continued for 10 Myr, after which the pace of transitions to reef habitats slowed. Combined, these patterns match a classic niche-filling scenario: early transitions to reefs were made rapidly by morphologically distinct lineages and were followed by a decrease in the rate of invasions and eventual saturation of morphospace. Major alterations in reef composition, distribution and abundance, along with shifts in climate and oceanic currents, occurred during the Late Cretaceous and early Palaeogene interval. A causal mechanism between these changes and concurrent episodes of reef invasion remains obscure, but what is clear is that the broad framework of the modern reef fish fauna was in place within 10 Myr of the end-Cretaceous extinction.