Philippa Kaur

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Ecology of Yellowfoot limpetIn the Tropical Pacific, sympatric limpets (Cellana melanostoma, Cellana exarata, Cellana sandwicensis, Cellana talcosa) inhabit the Hawaiian rocky intertidal ecosystem, where they graze on crustose coralline algae (CCA) and epibenthic microorganisms. Distribution ranges from the splash zone (upper-intertidal) to subtidal zone, and across the entire Hawaiian Archipelago26. They are dispersed across the majority of seamounts, atolls, and islands, however, not all species are present in every rocky intertidal locality, which reflects species-specific micro-habitat preferences.The reproduction cycles for each species appears to vary in time and space, and on-going long-term monitoring efforts are in progress to define this critical life-history trait. Previous studies on the yellowfoot limpet C. sandwicensis, reveal that reproduction is highly synchronized from December to March27,29. Gametogenesis also occurs from June to August, however, the level synchronicity and intensity of this second spawn period are inconsistent.These limpets are gonochoristic and considered to be sequential hermaphrodites44. The sex ratio is near 1:1(M:F) during spawning season, however, we have directly observed populations to maintain disproportionate sex ratios.Development of this broad-cast spawning limpet has been described from egg to post-larvae, where settlement occurs in less than 4 days post-fertilization29. This short larval duration ensures recruitment to the same localized intertidal environment, and reduces likelihood of hybridization between sympatric species with similar life-histories26.For wild limpets, growth rates shift through ontogeny—average monthly growth decreasing from 4–5 mm shell length (SL) as juveniles to 2–3 mm SL as adults27. Limpets also exhibit seasonal growth patterns—influenced by temperature and feeding28,37. Currently, growth rates of large individuals ( >50 mm SL) and species longevity are absent in the literature.Regional climate and coastal oceanographyKa’alawai is located on the south-facing shoreline of Oahu Island, Hawai’i (21°15’20.7“N 157°47’30.8“W). This area, defined as a rocky intertidal zone, is primarily comprised of basalt outcrops, boulders and benches, and supports a diverse community of epibenthic flora and fauna. The area is relatively easy to access by foot, and has been continuously exposed to various anthropogenic factors, which includes development, urban run-off, and subsistence fishing.The microclimate of the region is characterized by mild, wet winters (January to March) and dry, hot summers (July to September). The mean daily atmospheric temperature range and mean daily sea-surface temperature range are 18.44–31.38 °C and 22.67–30.18 °C, respectively. The annual precipitation is low relative to windward sides of the island, with maximum rainfall of 6.35 cm (data sources: US climate station USC00519397: Waikiki 717.2; PacIOOS Nearshore Sensor 04 (NS04): Waikiki Aquarium). Although freshwater input from precipitation along this coastline is considered to be marginal, the mixing of submarine groundwater discharge generates a unique geochemical profile for surface seawater at Ka’alawai. In particular, the mean surface salinity for this study site has been reported to be 25.4 ‰, which reflects this highly localized land-sea interaction45.The coastal oceanography of this region is predominantly influenced by wave, wind, and tidal forces. The south-shore region experiences a mixed tidal cycle—having both diurnal and semi-diurnal sinusoidal constituents per lunar day—with a tidal range of 58 cm and 91 cm during neap tide and spring tide, respectively; The trade winds from north-easterly direction (between 22.5°–67.5°) account for ~63% of the year with mean annual intensity around 5 m/s;46 and South swells with wave amplitudes of ~3 m are generated by storms in the Tasmanian Sea during Northern Hemisphere Summer47,48.Modern and historical specimensOn June 28th of 2018, live Yellowfoot limpet (Cellana sandwicensis) specimens CW1 and CW2 were collected from the rocky intertidal zone at Ka’alawai, Oahu, Hawai’i (Fig. 7). The animals were immediately sacrificed/dissected using scalpel blade, and measured for shell dimensions using a caliper. Limpets were weighed to determine gonadosomatic index, and gonads were preserved for histological examination. Shells were rinsed in an ultrasonic bath and air-dried.Fig. 7: Study site map.Hawaiian limpet specimens (Cellana sandwicensis) were collected along the rocky intertidal shoreline of Ka’alawai (Oahu, Hawaii). Instrumental sea-surface temperatures were measured in-situ by PacIOOS Nearshore Sensor 04 (NS04) at the Waikiki Aquarium.Full size imageA historical specimen BPBM (identification number 250851-200492) was loaned from the Bernice Pauahi Bishop Museum Malacology Department Collection. This specimen’s geographical and ecological origin is unknown, but was identified as C. sandwicensis by its characteristic shell morphology49. This specimen was selected for its large size to estimate life-expectancy of this limpet species, as well as to evaluate this method for paleoclimatology studies.Permission was not required to obtain specimens used in this study, and limpets were collected at a size exceeds the legal minimum shell length of 31.8 mm (Hawaii State Law is enforced by Department of Land and Natural Resources). Ethical approval was not required to conduct analysis.Characterization of shell microstructureShell microstructure was identified before isotopic analysis could be attempted. Each shell was cross-sectioned from anterior to posterior direction using a low speed saw (Isomet 1000, Buehler) equipped with a 0.5 mm diamond coated blade. Parallel cuts were made at the apex or maximal growth-axis to obtain two replicate 1.3 mm thick-sections per specimen. The first replicate thick-sections, prepared for micro-sampling, were further cut into More

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