Tooth Mg/Ca ratios and Aristotle's lantern morphometrics reflect trophic types in echinoids.

Several studies have inferred the ecological significance regarding the morphometrics of Aristotle's lantern and the mechanical properties of magnesium in echinoid teeth. This study attempts to combine these aspects, connecting them to the trophic habits of three native and an invasive echinoid in the Eastern Mediterranean Sea. Spatiotemporal data from the central and southern Aegean Sea were obtained, regarding the relative size of lanterns and demi-pyramids of Arbacia lixula, Paracentrotus lividus, Sphaerechinus granularis, and the invasive echinoid Diadema setosum and the Mg/Ca ratios of four zones on the tooth cross-section. Since environmental factors affect the examined factors, data for temperature, salinity, and concentration of chlorophyll-a were included in a principal component analysis. A. lixula and P. lividus presented intraspecific differences in the relative size of the lantern and demi-pyramid, while S. granularis and D. setosum exhibited variation in the elongation index. Differences in the Mg/Ca ratios were observed for all species although in different zones. Temperature appears to be related to all Mg/Ca zones except for the stone part, while the elongation index showed an inverse trend to all other morphometric parameters. The results of the PCA for the four species on the spatiotemporal level exhibited a distinction of individuals with season but not species, except for A. lixula, an omnivore with a carnivorous tendency, which was clearly separated from the herbivorous species. Using hierarchical clustering on the principal components it was evident that the three native species occupy different clusters, but when D. setosum was added, it shared the same cluster with S. granularis. This might infer similar feeding preferences, specifically for coralline algae, which might lead to a swift in the ecological equilibrium in regions, where D. setosum is found, either by affecting habitat type, or by restricting the distribution of S. granularis as was previously observed with Diadema africanum.

Abundance and population characteristics of the invasive sea urchin Diadema setosum (Leske, 1778) in the south Aegean Sea (eastern Mediterranean).

BACKGROUND: The Indo-Pacific sea urchin Diadema setosum has invaded the Mediterranean Sea and has spread along many locations in the southeastern part of the basin, where established populations exist on the shallow subtidal rocky shore. Diadema setosum is a ubiquitous species, of particular ecological importance due to the high levels of grazing pressure it imposes on benthic communities. Its biology, however, is not adequately studied, especially along its introduced range of distribution. The present study examines the population status of D. setosum outside its native range, in the Dodecanese island complex, south Aegean Sea. Thirty-four stations located across 16 islands were surveyed by scientific SCUBA-diving (up to a depth of 10 m) in December 2019 and June-July 2020. Samplings included: (i) visual census along transects to estimate relative abundance and population density, and (ii) random collection of specimens from densely populated stations to assess biometry and reproductive condition (histological examination of gonads) of D. setosum. RESULTS: Diadema setosum was found in 21 out of the 34 surveyed stations. The species had sparse populations of well-hidden individuals in rocky crevices, but with dense localized patches in Agathonisi, Leros, Kalymnos, Pserimos, Symi, Alimia and Chalki islands. In those seven islands, mean population density was 2.5 ± 1.48 individuals m-2. Diadema setosum had denser populations in shallower depths but larger dimensions in deeper; these results suggest segregated density and size patterns along a depth gradient. The size structure, according to the size frequency distribution of the test diameter, was unimodal with a fitted mode at 4.0-4.5 and 6.5-7.0 cm in shallow and deep populations, respectively. The examined morphometric relationships followed negative allometry, as previously suggested for the species within its native range of distribution, and test diameter appeared to be a good predictor of biomass. Diadema setosum specimens had immature gonads in winter and mature in summer, suggesting a synchronous reproductive pattern. These results conform to previous data from temperate populations of the species. CONCLUSIONS: Differences in local environmental conditions, e.g. hydrodynamics and habitat type, together with biotic interactions, e.g. recruitment and competition, probably shape D. setosum population in the south Aegean distributional range. The establishment of D. setosum has severe implications on benthic communities and local sea urchin populations demanding management measures to prevent the forecasted further expansion of this invasive species.

Mechanical defensive adaptations of three Mediterranean sea urchin species.

In the Mediterranean, Paracentrotus lividus and Sphaerechinus granularis are important drivers of benthic ecosystems, often coexisting in sublittoral communities. However, the introduction of the invasive diadematoid Diadema setosum, which utilizes venomous spines, may affect these communities. To describe the mechanical properties of the test and spines of these three species, specimens were collected in winter of 2019 from the sublittoral zone of the Dodecanese island complex, southeastern Aegean Sea. This region serves as a gateway for invasive species to the Mediterranean Sea. Crushing test was conducted on live individuals, while 3-point bending test was used to estimate spine stiffness. Porosity and mineralogy of the test and spine, thickness of the test, and breaking length of the spine were measured and compared, while the microstructural architecture was also determined. The test of S. granularis was the most robust (194.35 ± 59.59 N), while the spines of D. setosum (4.76 ± 2.13 GPa) exhibited highest flexibility. Increased porosity and thickness of the test were related to increased robustness, whereas increased flexibility of the spine was attributed to high porosity, indicating that porosity in the skeleton plays a key role in preventing fracture. The spines of S. granularis exhibited highest length after fracture % (71.54 ± 5.5%). D. setosum exhibited higher values of Mg concentration in the test (10%) compared with the spines (4%). For the first time, the mineralogy of an invasive species is compared with its native counterpart, while a comparison of the mechanical properties of different species of the same ecosystem also takes place. This study highlights different ways, in which sea urchins utilize their skeleton and showcases the ecological significance of these adaptations, one of which is the different ways of utilization of the skeleton for defensive purposes, while the other is the ability of D. setosum to decrease the Mg % of its skeleton degrading its mechanical properties, without compromising its defense, by depending on venomous bearing spines. This enables this species to occupy not only tropical habitats, where it is indigenous, but also temperate like the eastern Mediterranean, which it has recently invaded.

Chemical Composition and Microstructural Morphology of Spines and Tests of Three Common Sea Urchins Species of the Sublittoral Zone of the Mediterranean Sea.

In the Mediterranean Sea, the species Arbacia lixula, Paracentrotus lividus and Sphaerechinus granularis often coexist, occupying different subareas of the same habitat. The mechanical and chemical properties of their calcitic skeletons are affected both by their microstructural morphology and chemical composition. The present study describes the main morphologic features and the possible temporal differences in elemental composition of the test and spines of the three species, while also determining the molar ratio of each element of their crystalline phase. Scanning electron microscopy showed major differences in the ultrastructure of the spines, while minor differences in the test were also noticed. More specifically, the spines of all three sea urchins possess wedges, however A. lixula exhibits bridges connecting each wedge, while barbs are observed in the wedges of S. granularis. The spines of P. lividus are devoid of both microstructures. Secondary tubercles are absent in the test of A. lixula, while the tests and spines of all three species are characterized by different superficial stereom. Energy dispersive x-ray spectroscopy detected that Ca, Mg, S, Na and Cl were present in all specimen. Mg and Mg/Ca showed significant differences between species both in test and spines with S. granularis having the highest concentration. The spines of P. lividus exhibited lowest values between all species. Differences between spines and test were observed in all elements for P. lividus except S. A. lixula exhibited different concentrations between test and spines for Ca, Mg and Mg/Ca, whereas S. granularis for Mg, Cl and Mg/Ca. Finally, temporal differences for Ca were observed in the test of P. lividus and the spines of S. granularis, for Mg in test of S. granularis, for S in the spines of A. lixula and the test and spine of S. granularis, for Na in the test of P. lividus and A. lixula and for Cl and Mg/Ca in the test P. lividus. Powder X-ray diffractometry determined that, out of all three species, the spines of P. lividus contained the least Mg, while the test of the same species exhibited higher Mg concentration compared to A. lixula and S. granularis. The current study, although not labeling the specimens attempts to estimate potential time-related elemental differences among other results. These may occur due to changes in abiotic factors, probably water temperature, salinity and/or pH. Divergence in food preference and food availability may also play a key role in possible temporal differences the skeletons of these species.