Importance, biodiversity and dynamics of coccolithophorids in the coastal area of the north-east Adriatic sea

Dijatomeje, dinoflagelati i kokolitoforidi su najvažnije fitoplanktonske skupine u obalnim morima, od kojih je posljednja najslabije istražena u Jadranu. Iako je za sjeverni Jadran u literaturi zabilježeno 86 vrsta kokolitoforida, redovito ih se zamjećuje tek 16. Sukladno novim spoznajama o složenosti njihovog životnog ciklusa, potrebita je revizija njihove bioraznolikosti te ekološkog značaja u Jadranu. Uzorci za analizu fitoplanktonske zajednice i popratnih fizičko-kemijskih čimbenika sakupljani su mjesečno tijekom 2008. i 2009. godine duž zapadne obale Istarskog poluotoka. Za dodatne analize fotosintetskih pigmenata i kokolitoforida uzorkovano je svakih deset dana. Tijekom studija zabilježeno je ukupno 52 taksona kokolitoforida, a dominirao je rod Syracosphaera, sa zastupljenih 13 vrsta. Kokolitoforidi heterofaze (HET) i holofaze (HOL) bili su prisutni tijekom cijelog istraživanog razdoblja, a sezonalnost HET/HOL faze primijećena je za Coronosphaera mediterranea (jesen)/ HOL Calyptrolithina wettsteini (proljeće) i Syracosphaera pulchra (ljeto)/ HOL Calyptrosphaera oblonga (proljeće). Analiza pigmenata ukazuje na dominaciju 19'-heksanoilofukoksantina, pigmenta svojstvenog za haptofite. Visoke koncentracije tog pigmenta zabilježene su tijekom cijele godine s najvišim vrijednostima u kasno proljeće. Razdoblje proljeća (ožujak - lipanj) obilježila je visoka raznolikost vrsta i veći doprinos kokolitoforida ukupnoj fitoplanktonskoj zajednici (> 60%). Brojnost kokolitoforida Rhabdosphaera clavigera i Syracosphaera pulchra, značajno je povezana s visokim temperaturama i niskim koncentracijama nitrata, što je uobičajeno za ljetne mjesece. Nasuprot, brojnost vrsta Calciosolenia murrayi i C. brasiliensis značajno je bila povezana s povišenim koncentracijama hranjivih soli, te je njihov razvoj bio karakterističan za jesensku fitoplanktonsku zajednicu. Navedene vrste su se pokazale uspješnim kompetitorima s dijatomejama koje su inače dominirale fitoplanktonskom zajednicom. Najbrojnija je bila vrsta Emiliania huxleyi (106 stanica L-1 ), tijekom monospecifičnog cvata (92% od ukupnog fitoplanktona) u siječnju. E. huxleyi je najčešći kokolitoforid, povezan s niskim temperaturama i visokim koncentracijama hranjivih soli, prisutan u velikom broju i zimi i ljeti.Diatoms, dinoflagellates and coccolithophorids are the most important groups in the marine phytoplankton community. Studies of coccolithophorid taxonomy and ecology in the Adriatic are scarce, unlike those on diatoms and dinoflagellates. Although 86 species of coccolithophorids were reported for the northern Adriatic, only 11 species are regularly recorded. A revision of coccolithophorid biodiversity and their ecological importance in the Adriatic was necessary due to new insights on the life cycles of species. This study was conducted during one year (2008 2009) and samples were collected every ten days in the coastal area in front of Rovinj. The research was completed using light and electron microscopic examinations together with pigment analysis by high-performance liquid chromatography. A total of 52 coccolithophore morphospecies were recorded during the study. Dominating genus was Syracosphaera, represented by 13 species. Coccolithophorids of both diploid (HET) and haploid (HOL) phases were present throughout the investigated period. Seasonality in HET/HOL phases was observed for Syracosphaera pulchra (summer)/ HOL Calyptrosphaera oblonga (spring) and Coronosphaera mediterranea (autum)/ HOL Calyptrolithina wettsteini (spring). Pigment analysis pointed to a clear domination of 19-hexanoyloxyfucoxanthin, the pigment characteristic for haptophytes. High concentrations of 19-hexanoyloxyfucoxanthin were recorded throughout the year with a distinct peak in the spring. The spring period (March - June) was characterised by a high species diversity and higher contribution of coccolithophorids to the total phytoplankton assemblage (>60%). The typical summer coccolithophores, Rhabdosphaera clavigera and Syracosphaera pulchra, were related to high temperatures and low nitrate concentrations. Coccolithophorids Calciosolenia murrayi and C. brasiliensis were related to elevated nutrient concentrations, and were characteristic for the autumn assemblage. They were able to find favourable living conditions despite the abundant occurrence of diatoms and low light transmission values. Highest abundances of the species Emiliania huxleyi (106 cells L-1 ) were recorded in January during a monospecific bloom (92% of whole phytoplankton community). E. huxleyi was the most frequently recognised species and was related to low temperature and nutrient rich waters, consequently present in high numbers both during winter and summer

Osmotrophy of dissolved organic carbon by coccolithophores in darkness

• The evolutionary and ecological story of coccolithophores poses questions about their heterotrophy, surviving darkness after the end-Cretaceous asteroid impact as well as survival in the deep ocean twilight zone. Uptake of dissolved organic carbon might be an alternative nutritional strategy for supply of energy and carbon molecules. • Using long-term batch culture experiments, we examined coccolithophore growth and maintenance on organic compounds in darkness. Radiolabeled experiments were performed to study the uptake kinetics. Pulse-chase experiments were used to examine the uptake into unassimilated, exchangeable pools versus assimilated, non-exchangeable pools. • We found that coccolithophores were able to survive and maintain their metabolism for up to 30 days in darkness, accomplishing about one cell division. The concentration dependence for uptake was similar to the concentration dependence for growth in Cruciplacolithus neochelis, suggesting that it was taking up carbon compounds and immediately incorporating them into biomass. We recorded net incorporation of radioactivity into the particulate inorganic fraction. • We conclude that osmotrophy provides nutritional flexibility and supports long-term survival in light levels well below threshold for photosynthesis. The incorporation of dissolved organic matter into particulate inorganic carbon, raises fundamental questions about the role of the alkalinity pump and the alkalinity balance in the sea