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Review

Back to Basics: Revision of Coccolithophore Species List in the Adriatic Sea

1
Division for Marine and Environmental Research, Ruđer Bošković Institute, 10000 Zagreb, Croatia
2
Laboratory of Plankton and Shellfish Toxicity, Institute of Oceanography and Fisheries, 21000 Split, Croatia
*
Author to whom correspondence should be addressed.
Water 2023, 15(3), 603; https://doi.org/10.3390/w15030603
Submission received: 30 December 2022 / Revised: 26 January 2023 / Accepted: 29 January 2023 / Published: 3 February 2023
(This article belongs to the Special Issue Marine Phytoplankton Diversity)

Abstract

:
Coccolithophores are unicellular marine algae belonging to the haptophyte group, characterized by the production of intricate calcium carbonate plates that adorn their cells and exhibit species-specific morphology. The Adriatic Sea has historically been the type locality of numerous phytoplankton species, including coccolithophores. However, since the initial description, many species have not been recorded during the frequent phytoplankton surveys in the Adriatic Sea. This was mainly because these surveys did not use electron microscopy, which is necessary for accurate species identification. In this study, we re-evaluate the coccolithophore species lists using historical records and compare them with recent surveys in the coastal and open waters of the Adriatic Sea. In light of changes in nomenclature resulting from clarification of the species’ life cycles, we update the taxonomic list of coccolithophore species occurring in the Adriatic.

1. Introduction

Coccolithophores are single-celled algae characterized by their ability to calcify. They are part of the marine phytoplankton and are among the most important primary producers in the sea. Because of their ability to both calcify and photosynthesize, they fundamentally affect the chemistry of seawater and are one of the most important drivers of the oceanic carbon cycle and thus the Earth’s climate [1]. Coccolithophores occur in both open and coastal marine waters and sometimes form large blooms [2] which are detectable in ocean colour satellite imagery [3].
The characteristic feature of coccolithophores is their intricately shaped calcite plates, coccoliths, which adorn their cells in at least one phase of their life cycle [4]. Coccolithophores have a complex haplo-diplontic life cycle in which both haploid (with a single set of chromosomes) and diploid (with two copies of each chromosome) phases can reproduce vegetatively. Coccolith morphology differs between these two phases. In the diploid phase, coccolithophores produce heterococcoliths, plates consisting of a series of intricately shaped calcite crystals that deviate from the typical rhombohedral geometry [5]. In their haploid phase, their cells are covered with holococcoliths, nannoliths, or they do not calcify at all and have only organic scales covering their cells. Holococcoliths consist of single rhombohedral crystals, while nannoliths are calcareous structures that do not have the characteristic features of either heterococcoliths or holococcoliths.
The taxonomy of coccolithophores is based primarily on the characteristic, species-specific morphology of the coccoliths that cover their cells, the coccospheres [6]. Coccospheres bearing different types of coccoliths have been considered to belong to separate taxa because of their morphological differences, so coccolithophores of the same taxa but at different stages of their life cycle have been incorrectly classified as different species. Although the dimorphic life cycle of coccolithophores was first described by Parke and Adams [7], life cycle associations have only been clarified for ~60 coccolithophore species, representing ~20% of the described diversity, with examples distributed throughout the diversity of coccolithophores [8]. The main tool for clarifying life cycle associations is the detection of “combination cells” (coccospheres, bearing both hetero- and holoccoliths, first defined in Thomsen et al. [9]) in natural samples using scanning electron microscopy (SEM) [8].
The currently known diversity of coccolithophores includes ~280 morphospecies [8], including ~60 holococcolithophores, which should probably be excluded as they likely represent the life cycle stage of heterococcolithophores [10]. Coccolithophores are most diverse in subtropical and tropical waters [11]. In northwestern Mediterranean waters, Cros and Fortuño [12] reported 168 coccolithophores (including various taxonomic and morphotypic units). In the last comprehensive review of phytoplankton species in the Adriatic Sea, Viličić et al. [13] list 95 coccolithophore species in their Table 1. However, the authors note that the species diversity of coccolithophores throughout the Mediterranean is not comparable due to differences in identification methods, suggesting that the use of SEM for coccolithophores was not widespread at the time.
In this study, we aimed to investigate and review the taxonomic work on coccolithophores in the Adriatic Sea from the early 1900s to the early 2000s. We intended to update the taxonomy of these old works with new knowledge from the biology of coccolithophores. We also aimed to provide a comprehensive picture of the current diversity of coccolithophores in the Adriatic Sea by summarising published taxonomic surveys using SEM as a gold standard for coccolithophore identification. We anticipate that our efforts to update the checklist of coccolithophore species in the Adriatic Sea will be considered an important source of information for both research and policy.

2. Materials and Methods

In our analysis of historical and modern records of diversity in the Adriatic, we followed the recommendations of the taxonomy of the Nannotax3 website [14]. This online taxonomy project was originally based on the creation of online versions of the well-established syntheses of Bown [15] and Young et al. [6] and was later updated to include data from later publications. It is a useful basis for analysis, both because it is the most up-to-date review and because an attempt has been made to make it comprehensive, including both formally described species and many informally described morphotypes that are very likely to be true species.
For the analysis of the historical record, we used the works of Steuer [16], Brunnthaler [17], Schiller [18,19,20], Kamptner [21,22], Revelante [23], and Viličić et al. [13].
In 1903, Steuer [16] conducted studies in the Gulf of Trieste, while Brunnthaler [17] studied coastal waters near Rovinj, Croatia in 1910. Schiller’s studies [18,19,20] were based on material obtained during the seasonal voyages of the research steamer “Najade”, which investigated transects in the Adriatic Sea: Venice to Rovinj, Ancona to Dugi Otok Island, Bari to Dubrovnik, Otranto to Cap Linquetta in February, May, August, and November 1912. The investigation of coccolithophores by Kamptner [21,22] was mainly related to the coastal waters of the town of Rovinj and various other places on the southwestern coast of Istria, Croatia. The investigations of the coastal waters of the town of Rovinj were carried out quantitatively for a whole year, from the beginning of May 1935 to the end of April 1936. Samples of 2 L were completely sedimented and examined by light microscopy (LM) under high magnifications (1750×, 3750×, and 6000×).
Recent studies on coccolithophores in the northern Adriatic included Cerino et al. [24], Godrijan et al. [25], and Neri et al. [26]. Cerino et al. [24] analyzed data from monthly sampling between May 2011 and February 2013 at the C1-LTER coastal station in the Gulf of Trieste. Coccolithophore composition was determined using a polarized light microscope (PLM) and an SEM. Godrijan et al. [25] analyzed the coccolithophore community on the eastern Adriatic coast from September 2008 to December 2009, sampling fortnightly at 5 m depth. Morphospecies composition was determined using an SEM. Neri et al. [26] presented the entire phytoplankton community in the surface layer using LM (Utermöhl method) for the 1988–2018 data series from offshore station SG05, located 15 nm off the Italian coast, with variable temporal frequency (monthly to quarterly).
For the middle Adriatic, coccolithophores were studied by Šupraha et al. [27,28,29], Skejić et al. [30,31], and Young et al. [32], and in all taxonomic identification was performed by SEM. The works of Šupraha et al. [27,28,29] deal with the coccolithophore community in the coastal region of the Krka estuary in February and July 2013, and quantitative and qualitative analyses were performed for the vertical profile of the coccolithophore community. In their 2018 work, Skejić et al. [30] focused on data collected in December 2015 and April 2016. The study area includes the Jabuka Pit depressions (depth 260 m) and an Italy–Croatia transect (maximum depth 160 m). In their 2021 study, Skejić et al. [31] investigated the sampling site in the highly stratified Krka River estuary on the eastern Adriatic coast. Samples were collected from August to October 2017. Sampling occurred every two weeks at the surface (0 m), in the halocline (mainly 1.5 m), and at 7 m depth. Finally, Young et al. [32] report a single event from a surface water sample collected at an offshore station in March 2018.
Studies on coccolithophores in the southern Adriatic were scarce and, in the last 15 years, have included only the investigation by Balestra et al. [33]. The samples for this study were collected in October 2000 in a grid along four transects in the Gulf of Manfredonia, in the western part of the southern Adriatic. The data presented in this article are from quantitative analyses performed on the vertical water profile using the PLM and SEM.
Images were processed using the Adobe Lightroom 6.1. (Adobe, Mountain View, California, United States). Some of the images in figures have been published elsewhere, e.g., in Godrijan et al. [25], Skejić et al. [30], and Young et al. [32].

3. Results and Discussion

3.1. History of Coccolithophore Research in the Adriatic Sea

In 1903, Steuer [16] conducted the first survey of coccolithophores in the Adriatic Sea, in the Gulf of Trieste and found Syracosphaera pulchra, Rhabdosphaera stylifer, and Syracosphaera robusta. In 1910, Brunnthaler [17] found a much larger number of cocccoltithophores in the coastal waters near Rovinj. Overall, the Adriatic Sea is the type locality of many phytoplankton species, including coccolithophores. New species of coccolithophores in the Adriatic Sea were described by (i) Brunnthaler [17], who listed one new species of coccolithophores, Syracosphaera lohmanni Brunnthaler 1911; (ii) Schiller, who listed 23 new species in 1913 [18] and another 23 new species in 1925 [19]; and (iii) Kamptner, who listed 4 new species in his 1927 article [22] and 16 new species in 1941 [21].
The list of morphospecies in Table 1 was published by Erwin Kamptner in 1941 [21] as an overview of his efforts in the study of coccolithophores along the coast of the Istrian Peninsula in the northern Adriatic. The list includes 45 taxa. However, due to sparse information in the descriptions of some species and limited details in some line drawings of cocospheres, some of the listed species cannot be identified to species level. Moreover, some of the species listed are duplicates resulting from taxonomic revisions [14]. We note, then, that the Kamptner species list contains 35 morphotypes of coccolithophores from the northern Adriatic, for which we trust ourselves to give a currently accepted name. Furthermore, two of the listed species later turned out not to be coccolithophores but calcified dinoflagellate cysts of the species Scrippsiella acuminata (Ehrenberg) Kretschmann, Elbrächter, Zinssmeister, S.Soehner, Kirsch, Kusber, & Gottschling 2015. Kamptner also described Acanthoica rubus, but the line drawing of the coccosphere is rather insufficient to identify the species, and subsequently this morphospecies was never recorded again. Here, we propose that Corisphaera corona shares similarities with Syracosphaera periperforata Kleijne 1991 var. periperforata HOL in the highly arched circum-flagellar coccoliths, while Pontosphaera steueri resembles Hymenomonas globosa (Magne 1954) Gayral & Fresnel 1976, due to the organisation of its coccoliths and elliptical muroliths, and finally that Thalassopapus pellucidus Kampt has similarities to Calciopappus caudatus Gaarder & Ramsfjell, 1954 in the shape of its coccosphere and its circum-flagellar coccoliths modified into elongate spines.
Table 2 consists of a list of morphospecies compiled by Noelia Revelante in 1985 [23]. This work mentions a total of 81 morphospecies that have been recorded in the northern Adriatic. However, this list was compiled from data from the literature and lists species reported for the northern Adriatic by Brunnthaler, Schiller, and Kamptner. Going back to the original descriptions, some of the species cannot be recognized to the species level due to the lack of details in species descriptions and some line drawings of the cocospheres. Additionally, some of the listed species are duplicates as a consequence of taxonomic revisions [14]. We have marked all morphospecies that are not identifiable. We therefore conclude that the Revelante species list contains 41 morphotypes of coccolithophores from the northern Adriatic, for which we can give the currently accepted names.
Finally, in our historic analysis we present the list of species compiled by Viličić et al. in 2002 [13] (Table 3). The list indicates the distribution of coccolithophore morphospecies for the eastern Adriatic, given as follows: N—Northern, M—Middle, S—Southern. A total of 95 species are listed. Although the text indicates that there are 101 species of prymnesiophytes (Prymnesiophyceae—class of haptophytes, including coccolithophores), we could not identify the 6 additional species in any of the tables in the paper. Going back to the original descriptions, again, some of the listed species cannot be determined to the specific species level due to the lack of information in species descriptions and the lack of features in some line drawings of the cocosphere. Additionally, due to taxonomic revisions, several of the listed species are duplicates. [14]. We have marked all morphospecies that are not identifiable. Thus, we conclude that the Viličić et al. 2002 species list contains 51 morphotypes of coccolithophores from the revised eastern part of the Adriatic for which we are confident to give the currently accepted name.

3.2. Recent Studies on Coccolithophores in the Northern Adriatic

In the last 15 years, the coccolithophore assemblage has been studied in several publications covering the northern, central and southern Adriatic [24,25,26,27,28,29,30,31,32,33]. These publications reveal a great diversity of coccolithophores not only in the open sea, but also in coastal and estuarine waters.
Table 4 contains a list of 141 coccolithophores morphotypes published during last 15 years in the previously mentioned publications (combination coccospheres were not considered). Of these 141 morphotypes, 84 were recorded in the HET—diploid phase and the remaining 58 morphotypes in the haploid phase (including HOL, POL, NAN). Seventeen species were confirmed in both phases (HET-HOL), marked in bold in Table 4. The available images of coccolithophores from the northern and middle Adriatic Sea are listed in Figure 1, Figure 2, Figure 3, Figure 4 and Figure 5.
The most diverse area was the middle Adriatic with 124 morphotypes, but this was probably the result of more intensive research efforts (six publications) compared to the northern (three) and southern parts (one). In addition, 54 morphotypes were found from the northern part and 48 from the southern part of the Adriatic.
The most frequently observed species is E. huxleyi throughout the Adriatic Sea [25,30,33]. However, the highest abundance of coccolithophores was observed for Syracosphaera halldalii, reaching 2.51 × 106 cells L−1 in the middle Adriatic Sea [31].
Records of combination coccospheres (with HET-HOL coccoliths) are important for clarifying taxonomic questions due to the different morphological life cycle stages of coccolithophores. A review of the literature reveals that 11 combination coccospheres were reported in four publications: Acanthoica quattrospina, Alisphaera unicornis, Helicosphaera pavimentum, Syracosphaera arethusae, Syracosphaera bannockii, Syracosphaera histrica, Syracosphaera marginaporata, Syracosphaera mediterranea, Syracosphaera molischii, Syracosphaera pulchra, Syracosphaera strigilis [28,29,30,32].
In summary, we can state that in the past twenty years there has been a notable increase in the number of recorded species in the Adriatic. This increase is most likely due to the use of SEM, which allows a more detailed observation of species characteristics and thus more certain identification. The species lists by Revelante in 1985 [23] and by Viličić et al. in 2002 [13] were largely based on revisions of the older literature, with some species descriptions and associated drawings not being detailed enough to allow unambiguous identification, resulting in duplicate species records. Therefore, we would like to emphasize the importance of including microscopic photographs with species lists, as this allows unambiguous recording of species.

Author Contributions

Conceptualization and writing—original draft preparation: J.G.; formal analysis: J.G., J.A. and S.S.; visualization: M.B.; data curation: J.G., S.S. and J.A.; writing—review and editing, M.B., S.S. and J.A.; funding acquisition, J.G., S.S. and J.A. All authors have read and agreed to the published version of the manuscript.

Funding

This research was funded by the Croatian National Monitoring Program (Project “Jadran”) and by grants from the Croatian Science Foundation under projects UIP-2020-02-3249—Ecology and toxicity of the genus Pseudo-nitzschia in the coastal waters of the central and southern Adriatic Sea (Pseudotox).

Institutional Review Board Statement

Not applicable.

Informed Consent Statement

Not applicable.

Data Availability Statement

Not applicable.

Acknowledgments

We would like to thank everyone who participated in the collection of data sets used in this manuscript. We acknowledge the useful comments of anonymous reviewers.

Conflicts of Interest

The authors declare no conflict of interest.

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Figure 1. SEM images of coccolithophore morphotypes: (a) Acanthoica quattrospina; (b) Acanthoica quattrospina HOL; (c) Algirosphaera robusta; (d) Algirosphaera robusta HOL; (e) Alisphaera capulata; (f) Alisphaera extenta; (g) Alisphaera gaudii; (h) Alisphaera gaudii POL; (i) Alisphaera unicornis; (j) Alisphaera unicornis POL; (k) Calcidiscus leptoporus subsp. quadriperforatus; (l) Calcidiscus leptoporus subsp. quadriperforatus HOL; (m) Calciopapus rigidus; (n) Calciosolenia brasiliensis; (o) Calciosolenia corsellii; (p) Calciosolenia murrayi; (q) Calicasphaera blokii; (r) Calyptrosphaera heimdaliae; (s) Calyptrosphaera sphaeroidea; (t) Corisphaera gracilis; (scale bars: aj,q,r,t = 2 µm; s = 4 µm; k,l = 5 µm; m = 10 µm; np = 20 µm).
Figure 1. SEM images of coccolithophore morphotypes: (a) Acanthoica quattrospina; (b) Acanthoica quattrospina HOL; (c) Algirosphaera robusta; (d) Algirosphaera robusta HOL; (e) Alisphaera capulata; (f) Alisphaera extenta; (g) Alisphaera gaudii; (h) Alisphaera gaudii POL; (i) Alisphaera unicornis; (j) Alisphaera unicornis POL; (k) Calcidiscus leptoporus subsp. quadriperforatus; (l) Calcidiscus leptoporus subsp. quadriperforatus HOL; (m) Calciopapus rigidus; (n) Calciosolenia brasiliensis; (o) Calciosolenia corsellii; (p) Calciosolenia murrayi; (q) Calicasphaera blokii; (r) Calyptrosphaera heimdaliae; (s) Calyptrosphaera sphaeroidea; (t) Corisphaera gracilis; (scale bars: aj,q,r,t = 2 µm; s = 4 µm; k,l = 5 µm; m = 10 µm; np = 20 µm).
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Figure 2. SEM images of coccolithophore morphotypes: (a) Cyrtosphaera aculeata; (b) Discosphaera tubifera; (c) Emiliania huxleyi; (d) Florisphaera profunda; (e) Gladiolithus flabellatus; (f) Gliscolithus amitakareniae; (g) Helicosphaera carteri; (h) Helicosphaera hyalina; (i) Helicosphaera pavimentum; (j) Helicosphaera pavimentum HOL; (k) Helicosphaera HOL catilliferus type; (l) Helicosphaera HOL confusus type; (m) Homozygosphaera spinosa; (n) Navilithus altivelum; (o) Ophiaster formosus; (p) Ophiaster hydroideus; (q) Palusphaera crosiae; (r) Palusphaera vandelii; (s) Pappomonas sp. type 1; (t) Pappomonas sp. type 3; (scale bars: a,c,d,i,j,lo,q,s,t = 2 µm; b,eh,k,p,r = 5 µm).
Figure 2. SEM images of coccolithophore morphotypes: (a) Cyrtosphaera aculeata; (b) Discosphaera tubifera; (c) Emiliania huxleyi; (d) Florisphaera profunda; (e) Gladiolithus flabellatus; (f) Gliscolithus amitakareniae; (g) Helicosphaera carteri; (h) Helicosphaera hyalina; (i) Helicosphaera pavimentum; (j) Helicosphaera pavimentum HOL; (k) Helicosphaera HOL catilliferus type; (l) Helicosphaera HOL confusus type; (m) Homozygosphaera spinosa; (n) Navilithus altivelum; (o) Ophiaster formosus; (p) Ophiaster hydroideus; (q) Palusphaera crosiae; (r) Palusphaera vandelii; (s) Pappomonas sp. type 1; (t) Pappomonas sp. type 3; (scale bars: a,c,d,i,j,lo,q,s,t = 2 µm; b,eh,k,p,r = 5 µm).
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Figure 3. SEM images of coccolithophore morphotypes: (a) Papposphaera lepida; (b) Pileolosphaera longistirpes; (c) Poritectolithus poritectus; (d) Rhabdosphaera clavigera var. stylifera; (e) Rhabdosphaera xiphos; (f) Sphaerocalyptra sp. 1 HOL; (g) Syracosphaera ampliora; (h) Syracosphaera anthos; (i) Syracosphaera anthos HOL; (j) Syracosphaera arethusae; (k) Syracosphaera arethusae HOL; (l) Syracosphaera aurisinae HOL; (m) Syracosphaera azureaplaneta; (n) Syracosphaera bannockii; (o) Syracosphaera bannockii HOL; (p) Syracosphaera corolla; (q) Syracosphaera dilatata; (r) Syracosphaera elevata HOL; (s) Syracosphaera gaarderae HOL; (t) Syracosphaera halldalii; (scale bars: ac,es = 2 µm; d,t = 5 µm).
Figure 3. SEM images of coccolithophore morphotypes: (a) Papposphaera lepida; (b) Pileolosphaera longistirpes; (c) Poritectolithus poritectus; (d) Rhabdosphaera clavigera var. stylifera; (e) Rhabdosphaera xiphos; (f) Sphaerocalyptra sp. 1 HOL; (g) Syracosphaera ampliora; (h) Syracosphaera anthos; (i) Syracosphaera anthos HOL; (j) Syracosphaera arethusae; (k) Syracosphaera arethusae HOL; (l) Syracosphaera aurisinae HOL; (m) Syracosphaera azureaplaneta; (n) Syracosphaera bannockii; (o) Syracosphaera bannockii HOL; (p) Syracosphaera corolla; (q) Syracosphaera dilatata; (r) Syracosphaera elevata HOL; (s) Syracosphaera gaarderae HOL; (t) Syracosphaera halldalii; (scale bars: ac,es = 2 µm; d,t = 5 µm).
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Figure 4. SEM images of coccolithophore morphotypes: (a) Syracosphaera halldalii HOL; (b) Syracosphaera histrica; (c) Syracosphaera histrica HOL; (d) Syracosphaera lamina; (e) Syracosphaera marginaporata; (f) Syracosphaera mediterranea; (g) Syracosphaera mediterranea HOL gracillima type; (h) Syracosphaera meditteranea HOL hellenica type; (i) Syracosphaera mediterranea HOL wettsteini type; (j) Syracosphaera molischii type 1; (k) Syracosphaera molischii type 2; (l) Syracosphaera mollischi type 3; (m) Syracosphaera mollischi type 4; (n) Syracosphaera mollischi HOL; (o) Syracosphaera nana; (p) Syracosphaera nana HOL; (q) Syracosphaera nodosa; (r) Syracosphaera orbiculus; (s) Syracosphaera ossa type 1; (t) Syracosphaera ossa type 2; (scale bars: ac,e,f,h,i,kt = 2 µm; g,j = 5 µm; d = 10 µm).
Figure 4. SEM images of coccolithophore morphotypes: (a) Syracosphaera halldalii HOL; (b) Syracosphaera histrica; (c) Syracosphaera histrica HOL; (d) Syracosphaera lamina; (e) Syracosphaera marginaporata; (f) Syracosphaera mediterranea; (g) Syracosphaera mediterranea HOL gracillima type; (h) Syracosphaera meditteranea HOL hellenica type; (i) Syracosphaera mediterranea HOL wettsteini type; (j) Syracosphaera molischii type 1; (k) Syracosphaera molischii type 2; (l) Syracosphaera mollischi type 3; (m) Syracosphaera mollischi type 4; (n) Syracosphaera mollischi HOL; (o) Syracosphaera nana; (p) Syracosphaera nana HOL; (q) Syracosphaera nodosa; (r) Syracosphaera orbiculus; (s) Syracosphaera ossa type 1; (t) Syracosphaera ossa type 2; (scale bars: ac,e,f,h,i,kt = 2 µm; g,j = 5 µm; d = 10 µm).
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Figure 5. SEM images of coccolithophore morphotypes: (a) Syracosphaera periperforata var. periperforata; (b) Syracosphaera prolongata; (c) Syracosphaera protrudens; (d) Syracosphaera pulchra; (e) Syracosphaera pulchra HOL oblonga type; (f) Syracosphaera pulchra HOL pirus type; (g) Syracosphaera rotula; (h) Syracosphaera strigilis; (i) Syracosphaera tumularis; (j) Tergestiella adriatica; (k) Umbellosphaera tenuis; (l) Umbilicosphaera sibogae var. sibogae; (scale bars: ac,g,h,j,k = 2 µm; df,i = 5 µm; l = 10 µm).
Figure 5. SEM images of coccolithophore morphotypes: (a) Syracosphaera periperforata var. periperforata; (b) Syracosphaera prolongata; (c) Syracosphaera protrudens; (d) Syracosphaera pulchra; (e) Syracosphaera pulchra HOL oblonga type; (f) Syracosphaera pulchra HOL pirus type; (g) Syracosphaera rotula; (h) Syracosphaera strigilis; (i) Syracosphaera tumularis; (j) Tergestiella adriatica; (k) Umbellosphaera tenuis; (l) Umbilicosphaera sibogae var. sibogae; (scale bars: ac,g,h,j,k = 2 µm; df,i = 5 µm; l = 10 µm).
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Table 1. List of morphospecies published by Erwin Kamptner (1941) (Die Coccolithineen der Südwestküste von Istrien. Ann. Naturhist. Mus. Wien. 51: 54–149). HET and HOL indicate the life cycle stage, HET diploid and HOL haploid, and ‘-‘ indicates an unidentifiable life cycle stage. Table lists coccolithophore species along the coast of the Istrian peninsula, northern Adriatic Sea, including 16 new species for science. Accepted names were updated following the recommendations on Young, J.R., Bown P.R., Lees J.A. (2022) Nannotax3 website. International Nannoplankton Association. Accessed 14 December 2022. URL: www.mikrotax.org/Nannotax3. Comments are indicated with *.
Table 1. List of morphospecies published by Erwin Kamptner (1941) (Die Coccolithineen der Südwestküste von Istrien. Ann. Naturhist. Mus. Wien. 51: 54–149). HET and HOL indicate the life cycle stage, HET diploid and HOL haploid, and ‘-‘ indicates an unidentifiable life cycle stage. Table lists coccolithophore species along the coast of the Istrian peninsula, northern Adriatic Sea, including 16 new species for science. Accepted names were updated following the recommendations on Young, J.R., Bown P.R., Lees J.A. (2022) Nannotax3 website. International Nannoplankton Association. Accessed 14 December 2022. URL: www.mikrotax.org/Nannotax3. Comments are indicated with *.
NTypeSpecies Name (Kamptner List)Accepted Name with Comments (Indicated with *)
1HETAcanthoica aculeata nov. spec. Cyrtosphaera aculeata (Kamptner) Kleijne, 1992
2HETAcanthoica ordinata nov. spec.Alisphaera ordinata (Kamptner) Heimdal, 1973
3HETAcanthoica quattrospina Lohm.Acanthoica quatrospina Lohmann, 1903
4-Acanthoica rubus nov. spec.* ? line drawing of coccosphere inadequate to determine species
5HETAnthosphaera robusta (Lohm).Syracosphaera molischii Schiller, 1925 HOL
6HOLAnthosphqera fragaria Kampt. Algirosphaera robusta (Lohmann 1902) Norris, 1984
7HETCalciosolenia grani Schill, var. cylindrothecaeformis Schill.Calciosolenia brasiliensis (Lohmann) Young, in Young et al., 2003
8HOLCalyptrosphaera gracillima nov. spec.Syracosphaera mediterranea Lohmann 1902 HOL gracillima type, sensu Geisen et al., 2002
9HOLCalyptrosphaera pirus Kampt. Syracosphaera pulchra Lohmann, 1902 HOL pirus type, sensu Young et al., 2003
10HOLCalyptrosphaera quatridentata Schill.Sphaerocalyptra quadridentata (Schiller 1913) Deflandre, 1952
11HOLCalyptrosphaera tholifera nov. spec.Sphaerocalyptra quadridentata (Schiller 1913) Deflandre, 1952
12HETCoccolithus carteri (Wall.)Helicosphaera carteri (Wallich 1877) Kamptner, 1954
13HETCoccolithus leptoporus (Murr. & Blackm.)Calcidiscus leptoporus (Murray & Blackman 1898) Loeblich & Tappan, 1978
14HOLCorisphaera arethusae nov. spec.Syracosphaera arethusae (Kamptner 1941) Triantaphyllou et al. 2016 HOL
15HOLCorisphaera corona nov. spec.* Syracosphaera Lohmann 1902 HOL (Anthosphaera type); cf. Syracosphaera periperforata Kleijne 1991 var. periperforata HOL
16HOLCorisphaera gracilis Kampt.Corisphaera gracilis Kamptner, 1937
17HOLCorisphaera ponticulifera nov. spec.* Helicosphaera HOL ponticuliferus type informal
18HOLCorisphaera spinosa nov. spec.Homozygosphaera spinosa (Kamptner 1941) Deflandre, 1952
19HOLHelladosphaera aurisinae nov. spec.Syracosphaera aurisinae HOL (Kamptner 1941) Keuter et al., 2021
20HOLHelladosphaera cornifera (Schill.)Helladosphaera cornifera (Schiller 1913) Kamptner, 1937
21HOLPeriphyllophora mirabilis (Schill.).Syracosphaera anthos (Lohmann 1912) Janin, 1987 HOL
22HETPontosphaera huxleyi Lohm. Emiliania huxleyi (Lohmann 1902) Hay & Mohler, in Hay et al. 1967
23HETPontosphaera nana nov. spec.Syracosphaera nana (Kamptner 1941) Okada & McIntyre, 1977
24HOLPontosphaera pietschmanni Kampt.* ? clearly a holococcolith but holotype is drawing with too little detail to allow identification
25HETPontosphaera steueri Kampt.* ? cf. Hymenomonas globosa (Magne 1954) Gayral & Fresnel, 1976
26-Rhabdosphaera erinaceus Kampt.* not a coccolithophore: Scrippsiella acuminata (Ehrenberg) Kretschmann, Elbrächter, Zinssmeister, S.Soehner, Kirsch, Kusber & Gottschling 2015
27-Rhabdosphaera nigra Schill. * not a coccolithophore: Scrippsiella acuminata (Ehrenberg) Kretschmann, Elbrächter, Zinssmeister, S.Soehner, Kirsch, Kusber & Gottschling 2015
28HETRhabdosphaera stylifer Lohm.Rhabdosphaera clavigera Murray & Blackman, 1898
29HETSyracosphaera binodata Kampt.Syracosphaera mediterranea Lohmann 1902
30HOLSyracosphaera catillifera Kampt.Helicosphaera HOL catilliferus type Young & Bown (2014)
31HETSyracosphaera corii Schill.Michaelsarsia adriaticus (Schiller) Manton, Bremer & Oates, 1984
32HOLSyracosphaera cornus nov. spec.Helicosphaera HOL catilliferus type Young & Bown (2014)
33HOLSyracosphaera dalmatica Kampt.Syracosphaera mediterranea Lohmann 1902 HOL wettsteinii type, sensu Geisen et al., 2002
34HETSyracosphaera histrica nov. spec.Syracosphaera histrica Kamptner, 1941
35HETSyracosphaera molischi Schill.Syracosphaera molischii Schiller, 1925
36HETSyracosphaera nodosa nov. spec.Syracosphaera nodosa Kamptner, 1941
37HETSyracosphaera pulchra Lohm.Syracosphaera pulchra Lohmann, 1902
38HOLSyracosphaera quadriperforata Kampt.Calcidiscus leptoporus subsp. quadriperforatus (Kamptner 1937) Geisen et al., 2002 HOL
39HETSyracosphaera schilleri Kampt.Scyphosphaera apsteinii Lohmann, 1902 HOL
40HETSyracosphaera tuberculata Kampt.Syracosphaera mediterranea Lohmann 1902
41HETTergestiella adriatica nov. spec.Tergestiella adriatica Kamptner, 1941
42HETThalassopapus pellucidus Kampt.* ? cf. Calciopappus caudatus Gaarder & Ramsfjell, 1954
43HOLZygosphaera debilis nov. spec.Zygosphaera amoena Kamptner, 1937
44HOLZygosphaera hellenica Kampt.Syracosphaera mediterranea Lohmann 1902 HOL hellenica type, sensu Cros et al., 2000
45HOLZygosphaera wettsteini Kampt.Syracosphaera mediterranea Lohmann 1902 HOL wettsteinii type, sensu Geisen et al., 2002
Table 2. List of morphospecies compiled by Noelia Revelante (1985) (A catalogue of phytoplankton reported for the Rovinj area of the northern Adriatic. Thalass. Jugoslav. 21/22, 139–169). HET and HOL indicate the life cycle stage, HET diploid and HOL haploid, and ‘-‘ indicates an unidentifiable life cycle stage. This paper reports morphospecies recorded in the northern Adriatic Sea. Accepted names were updated following the recommendations of Young, J.R., Bown P.R., Lees J.A. (2022) Nannotax3 website. International Nannoplankton Association. Accessed 14 December 2022. URL: www.mikrotax.org/Nannotax3. Comments are indicated with *.
Table 2. List of morphospecies compiled by Noelia Revelante (1985) (A catalogue of phytoplankton reported for the Rovinj area of the northern Adriatic. Thalass. Jugoslav. 21/22, 139–169). HET and HOL indicate the life cycle stage, HET diploid and HOL haploid, and ‘-‘ indicates an unidentifiable life cycle stage. This paper reports morphospecies recorded in the northern Adriatic Sea. Accepted names were updated following the recommendations of Young, J.R., Bown P.R., Lees J.A. (2022) Nannotax3 website. International Nannoplankton Association. Accessed 14 December 2022. URL: www.mikrotax.org/Nannotax3. Comments are indicated with *.
NTypeSpecies Name (Revelante List)Accepted Name with Comments (Indicated with *)
1HETAcanthoica acanthos Schiller, 1925Acanthoica acanthos Schiller, 1925
2HETAcanthoica aculeata Kamptner 1941Cyrtosphaera aculeata (Kamptner) Kleijne, 1992
3HETAcanthoica monospina Schiller, 1925Acanthoica acanthifera Lohmann 1912 ex Lohmann, 1913
4HETAcanthoica ordinata Kamptner, 1941Alisphaera ordinata (Kamptner) Heimdal, 1973
5HETAcanthoica quattrospina Lohmann, 1903Acanthoica quattrospina Lohmann, 1903
6-Acantoica rubus Kamptner, 1941* ? line drawing of coccosphere inadequate to determine species
7HOLAnthosphaera fragaria Kamptner, 1937 Syracosphaera molischii Schiller, 1925 HOL
8HETAnthosphaera robusta (Lohmann) Kamptner, 1941Algirosphaera robusta (Lohmann 1902) Norris, 1984
9HETCalciosolenia granii Schiller, 1925Calciosolenia brasiliensis (Lohmann) Young, in Young et al. 2003
10HETCalciosolenia granii var cylindrothecaeformis Schiller, 1925 emend Kamptner 1936Calciosolenia brasiliensis (Lohmann) Young, in Young et al. 2003
11HOLCalyptrosphaera aff. globosa Lohmann in Borsetti & Cati, 1976Calyptrosphaera sphaeroidea Schiller, 1913 HOL
12HOLCalyptrosphaera gracillima Kamptner, 1941Syracosphaera mediterranea Lohmann 1902 HOL gracillima type, sensu Geisen et al., 2002
13HOLCalyptrosphaera incisa Schiller 1913Calyptrosphaera sphaeroidea Schiller, 1913 HOL
14HOLCalyptrosphaera oblonga Lohmann, 1902Syracosphaera pulchra Lohmann, 1902 HOL oblonga type, sensu Young et al., 2003
15HOLCalyptrosphaera pirus Kamptner, 1941Syracosphaera pulchra Lohmann, 1902 HOL pirus type, sensu Young et al., 2003
16HOLCalyptrosphaera pyriformis Schiller 1913Calyptrosphaera sphaeroidea Schiller, 1913 HOL
17HOLCalyptrosphaera quadridentata Schiller, 1913Sphaerocalyptra quadridentata (Schiller 1913) Deflandre, 1952
18HOLCalyptrosphaera sphaeroidea Schiller 1913Calyptrosphaera sphaeroidea Schiller 1913
19HOLCalyptrosphaera tholifera Kamptner, 1941Sphaerocalyptra quadridentata (Schiller 1913) Deflandre, 1952
20HETCoccolithus carteri (Wallich) Kamptner, 1941Helicosphaera carteri (Wallich 1877) Kamptner, 1954
21HETCoccolithus leptoporus (Murray & Blackman) Schiller, 1930Calcidiscus leptoporus (Murray & Blackman 1898) Loeblich & Tappan, 1978
22HETCoccolithus pelagicus var. carteri Kamptner, 1927Helicosphaera carteri (Wallich, 1877) Kamptner, 1954
23HETCoccolithus wallichii (Lohmann) Schiller, 1925Helicosphaera wallichii (Lohmann 1902) Okada & McIntyre, 1977
24HOLCorisphaera arethusae Kamptner, 1941Syracosphaera arethusae (Kamptner 1941) Triantaphyllou et al. 2016 HOL
25HOLCorisphaera corona Kamptner, 1941Syracosphaera sp. HOL (Anthosphaera type)
26HOLCorisphaera gracilis Kamptner, 1937Corisphaera gracilis Kamptner, 1937
27HOLCorisphaera ponticulifera Kamptner, 1941Helicosphaera HOL ponticuliferus type informal
28HOLCorisphaera spinosa Kamptner 1941Homozygosphaera spinosa (Kamptner 1941) Deflandre, 1952
29HOLHelladosphaera aurisinae Kamptner, 1941Syracosphaera aurisinae HOL (Kamptner 1941) Keuter et al., 2021
30HOLHelladosphaera cornifera (Schiller) Kamptner, 1937Helladosphaera cornifera (Schiller 1913) Kamptner, 1937
31HOLLohmanosphaera adriatica Schiller, 1913? cf. Sphaerocalyptra quadridentata (Schiller 1913) Deflandre, 1952
32-Najadea gloriosa Schiller, 1913* ? holotype is drawing with too little detail to allow identification
33HOLOphiaster formosus Gran, 1912Ophiaster formosus Gran, 1912
34HOLPeriphyllophora mirabilis (Schiller) Kamptner, 1937Syracosphaera anthos (Lohmann 1912) Janin, 1987 HOL
35-Pontosphaera echinofera Schiller (1913)* ? holotype is drawing with too little detail to allow identification
36-Pontosphaera haeckelii Schiller, 1913* ? holotype is drawing with too little detail to allow identification
37HETPontosphaera huxleyi Lohmann, 1902Emiliania huxleyi (Lohmann 1902) Hay & Mohler, in Hay et al. 1967
38-Pontosphaera inermis Schiller, 1913* ? holotype is drawing with too little detail to allow identification
39HETPontosphaera nana Kamptner, 1941Syracosphaera nana (Kamptner 1941) Okada & McIntyre, 1977
40-Pontosphaera ovalis Schiller, 1913* ? holotype is drawing with too little detail to allow identification
41HOLPontosphaera pietschmanni Kamptner, 1937* ? clearly a holococcolith but holotype is drawing with too little detail to allow identification
42HETPontosphaera steueri Kamptner, 1937* ? cf. Hymenomonas globosa (Magne 1954) Gayral & Fresnel, 1976
43HETRhabdosphaera clavigera Murray & Blackman, 1898Rhabdosphaera clavigera Murray & Blackman, 1898
44-Rhabdosphaera erinaceus Kamptner, 1937* not a coccolithophore: Scrippsiella acuminate (Ehrenberg 1836) Kretschmann et al., 2015
45HETRhabdosphaera longistylis Schiller, 1925* cf. Palusphaera vandelii Lecal, 1965
46-Rhabdosphaera nigra Schiller, 1913* not a coccolithophore: Scrippsiella acuminate (Ehrenberg 1836) Kretschmann et al. 2015
47HETRhabdosphaera stylifera Lohmann, 1902Rhabdosphaera clavigera Murray & Blackman, 1898
48HETRhabdosphaera tignifer Schiller, 1913Rhabdosphaera clavigera Murray & Blackman, 1898
49-Rhabdosphaera hispida Lohmann, 1912* ?
50-Syracosphaera adriatica Schiller, 1913 * ? line drawing of coccosphere inadequate to determine species
51-Syracosphaera bifenestrata Schiller, 1913* ? line drawing of coccosphere inadequate to determine species
52HETSyracosphaera binodata (Kamptner) Kamptner, 1937Syracosphaera mediterranea Lohmann, 1902
53HOLSyracosphaera catillifera Kamptner, 1937Helicosphaera HOL catilliferus type Young & Bown (2014)
54HETSyracosphaera cori Schiller in Heimdal & Gaarder, 1981Michaelsarsia adriaticus (Schiller) Manton, Bremer & Oates, 1984
55HOLSyracosphaera cornifera Schiller 1913Helladosphaera cornifera (Schiller 1913) Kamptner, 1937
56HOLSyracosphaera cornus Kamptner, 1941Helicosphaera HOL catilliferus type Young & Bown (2014)
57-Syracosphaera coronata Schiller, 1913* ? line drawing of coccosphere inadequate to determine species
58-Syracosphaera cupulifera Schiller, 1913* ? line drawing of coccosphere inadequate to determine species
59HOLSyracosphaera dalmatica Kamptner, 1927Syracosphaera mediterranea Lohmann 1902 HOL wettsteinii type, sensu Geisen et al., 2002
60-Syracosphaera dentata Schiller, 1913* ? line drawing of coccosphere inadequate to determine species
61-Syracosphaera grundii Schiller, 1913* ? line drawing of coccosphere inadequate to determine species
62HETSyracosphaera histrica Kamptner, 1941Syracosphaera histrica Kamptner, 1941
63-Syracosphaera lohmannii Brunnthaler, 1910* ? line drawing of coccosphere inadequate to determine species
64HETSyracosphaera mediterranea Lohmann, 1902Syracosphaera mediterranea Lohmann 1902
65HETSyracosphaera mediterranea var. biondada (Lohmann) Kamptner, 1927Syracosphaera mediterranea Lohmann 1902
66HETSyracosphaera molischii Schiller, 1925Syracosphaera molischii Schiller, 1925
67HETSyracosphaera nodosa Kamptner, 1941Syracosphaera nodosa Kamptner, 1941
68-Syracosphaera ovata Schiller, 1925* ? line drawing of coccosphere inadequate to determine species
69HETSyracosphaera pulchra Lohmann, 1902Syracosphaera pulchra Lohmann, 1902
70HOLSyracosphaera quadricornus Schiller, 1925Algirosphaera robusta (Lohmann 1902) Norris, 1984
71HOLSyracosphaera quadriperforata Kamptner, 1937 Calcidiscus leptoporus subsp. quadriperforatus (Kamptner 1937) Geisen et al., 2002 HOL
72HETSyracosphaera robusta Lohmann, 1902Algirosphaera robusta (Lohmann 1902) Norris, 1984
73HETSyracosphaera schilleri Kamptner, 1927Scyphosphaera apsteinii Lohmann, 1902 HOL
74-Syracosphaera spinosa Lohmann, 1902* ? line drawing of coccosphere inadequate to determine species
75-Syracosphaera tenuis Lohmann, 1902* ? line drawing of coccosphere inadequate to determine species
76HETSyracosphaera tuberculata Kamptner 1937Syracosphaera mediterranea Lohmann 1902
77HETTergestiella adriatica Kamptner, 1941Tergestiella adriatica Kamptner, 1941
78HETThalassopapus pellucidus Kamptner, 1941* ? cf. Calciopappus caudatus Gaarder & Ramsfjell, 1954
79HOLZygosphaera debilis Kamptner, 1941Zygosphaera amoena Kamptner, 1937
80HOLZygosphaera hellenica Kamptner, 1937Syracosphaera mediterranea Lohmann 1902 HOL hellenica type, sensu Cros et al., 2000
81HOLZygosphaera wettsteinii Kamptner, 1937Syracosphaera mediterranea Lohmann 1902 HOL wettsteinii type, sensu Geisen et al., 2002
Table 3. List of morphospecies compiled by Viličić et al. (2002) (Checklist of phytoplankton in the eastern Adriatic Sea. Acta. Bot. Croat. 61 (1), 57–91). HET and HOL indicate the life cycle stage, HET diploid and HOL haploid, and ‘-‘ indicates an unidentifiable life cycle stage. Only one name per species is listed (additional synonyms are listed in Viličić et al. 2002). This paper lists morphospecies reported for the eastern Adriatic Sea, distribution is noted as: N—Northern, M—Middle, S—Southern, and ˟ registered by Revelante (1985), reportedly not found afterwards. Accepted names were updated following the recommendations of Young, J.R., Bown P.R., Lees J.A. (2022) Nannotax3 website. International Nannoplankton Association. Accessed 14 December 2022. URL: www.mikrotax.org/Nannotax3. Comments are indicated with *.
Table 3. List of morphospecies compiled by Viličić et al. (2002) (Checklist of phytoplankton in the eastern Adriatic Sea. Acta. Bot. Croat. 61 (1), 57–91). HET and HOL indicate the life cycle stage, HET diploid and HOL haploid, and ‘-‘ indicates an unidentifiable life cycle stage. Only one name per species is listed (additional synonyms are listed in Viličić et al. 2002). This paper lists morphospecies reported for the eastern Adriatic Sea, distribution is noted as: N—Northern, M—Middle, S—Southern, and ˟ registered by Revelante (1985), reportedly not found afterwards. Accepted names were updated following the recommendations of Young, J.R., Bown P.R., Lees J.A. (2022) Nannotax3 website. International Nannoplankton Association. Accessed 14 December 2022. URL: www.mikrotax.org/Nannotax3. Comments are indicated with *.
NTypeSpecies Name (Viličić et al. 2002 List)Accepted Name with Comments (Indicated with *)Distribution
1HETAcanthoica acanthos (Schiller) DeflandreAcanthoica acanthos Schiller, 1925N
2HETAcanthoica monospina SchillerAcanthoica acanthifera Lohmann, 1912 ex Lohmann, 1913N
3HETAcanthoica quattrospina Lohmann Acanthoica quattrospina Lohmann, 1903N, M, S
4-Acanthoica rubus Kamptner* ? line drawing of coccosphere inadequate to determine speciesN ˟
5HOLAlgirosphaera quadricorni (Schiller) NorrisAlgirosphaera robusta (Lohmann 1902) Norris, 1984N, M, S
6HETAlgirosphaera robusta (Lohmann) NorrisAlgirosphaera robusta (Lohmann 1902) Norris, 1984N ˟
7HETAlisphaera ordinata (Kamptner) HeimdalAlisphaera ordinata (Kamptner) Heimdal, 1973N ˟
8HETAnoplosolenia brasiliensis (Lohmann) Deflandre Calciosolenia brasiliensis (Lohmann) Young, in Young et al. 2003N, S
9HOLAnthosphaera fragaria KamptnerSyracosphaera molischii Schiller, 1925 HOLN ˟
10HETCalcidiscus leptoporus (Murray & Blachman) Loeblich Jr. & TappanCalcidiscus leptoporus (Murray & Blackman, 1898) Loeblich & Tappan, 1978N ˟
11HETCalciopappus caudatus Gaarder & Ramsfjell Calciopappus caudatus Gaarder & Ramsfjell, 1954M, S
12HETCalciopappus rigidus Heimdal Calciopappus rigidus Heimdal in Heimdal & Gaarder, 1981N, M
13HETCalciosolenia murrayi Gran Calciosolenia murrayi Gran, 1912not reported
14HOLCalyptrolithina wettsteinii (Kamptner) Kleijne Syracosphaera mediterranea Lohmann 1902 HOL wettsteinii type, sensu Geisen et al., 2002N ˟
15-Calyptrosphaera dalmatica Schiller* ?N ˟
16HOLCalyptrosphaera globosa LohmannCalyptrosphaera sphaeroidea Schiller, 1913 HOLN ˟, M
17HOLCalyptrosphaera incisa Schiller Calyptrosphaera sphaeroidea Schiller, 1913 HOLN ˟
18HOLCalyptrosphaera oblonga Lohmann Syracosphaera pulchra Lohmann, 1902 HOL oblonga type, sensu Young et al., 2003N, M, S
19HOLCalyptrosphaera pirus Kamptner Syracosphaera pulchra Lohmann, 1902 HOL pirus type, sensu Young et al., 2003N ˟
20HOLCalyptrosphaera pyriformis Schiller Calyptrosphaera sphaeroidea Schiller, 1913 HOLN ˟
21HOLCalyptrosphaera quadridentata SchillerSphaerocalyptra quadridentata (Schiller 1913) Deflandre, 1952N ˟
22HOLCalyptrosphaera sphaeroidea SchillerCalyptrosphaera sphaeroidea Schiller 1913 HOLN ˟
23-Calyptrosphaera uvella Schiller * ?N ˟
24HETCoccolithus pelagicus (Wallich) Lohmann Helicosphaera carteri (Wallich) Kamptner, 1954N ˟
25HETCoccolithus wallichii (Lohmann) SchillerHelicosphaera wallichii (Lohmann 1902) Okada & McIntyre, 1977N ˟
26HOLCorisphaera arethusae Kamptner Syracosphaera arethusae (Kamptner 1941) Triantaphyllou et al., 2016 HOL
27HOLCorisphaera corona Kamptner Syracosphaera Lohmann, 1902 HOL (Anthosphaera type)N ˟
28HOLCorisphaera gracillis Kamptner Corisphaera gracilis Kamptner, 1937N ˟
29HOLCorisphaera spinosa KamptnerHomozygosphaera spinosa (Kamptner 1941) Deflandre, 1952N ˟
30HETCoronosphaera binodata (Kamptner) GaarderSyracosphaera mediterranea Lohmann 1902N ˟
31HETCoronosphaera mediterraneaSyracosphaera mediterranea Lohmann 1902N ˟
32HETCyrtosphaera aculeata (Kamptner) Kleijne Cyrtosphaera aculeata (Kamptner) Kleijne, 1992N ˟
33HETDiscosphaera thomsonii Ostenfeld Discosphaera tubifera (Murray & Blackman 1898) Ostenfeld, 1900S
34HETDiscosphaera tubifera (Murray & Blackman) Ostenfeld Discosphaera tubifera (Murray & Blackman 1898) Ostenfeld, 1900S
35HETEmiliania huxleyi (Lohmann) Hay & Mohler Emiliania huxleyi (Lohmann 1902) Hay & Mohler, in Hay et al. 1967N, M, S
36HETHelicosphaera carteri (Wallich) KamptnerHelicosphaera carteri (Wallich 1877) Kamptner, 1954N ˟
37HOLHelladosphaera cornifera (Schiller) KamptnerHelladosphaera cornifera (Schiller 1913) Kamptner, 1937N ˟
38HOLHomozygosphaera ponticulifera (Kamptner) KamptnerSyracosphaera mediterranea Lohmann 1902 HOL gracillima type, sensu Geisen et al., 2002N ˟
39-Homozygosphaera tholifera (Kamptner) Halldal & MarkaliSphaerocalyptra quadridentata (Schiller 1913) Deflandre, 1952N ˟
40HOLLohmannosphaera adriatica SchillerSphaerocalyptra quadridentata (Schiller 1913) Deflandre, 1952N ˟
41HETLohmannosphaera paucoscyphos SchillerHelicosphaera carteri (Wallich) Kamptner, 1954N ˟
42HETMichaelsarsia adriatica Schiller emend. Manton, Bremer & BatesMichaelsarsia adriaticus (Schiller) Manton, Bremer & Oates, 1984M, S
43HETMichaelsarsia falklandica Lohmann Michaelsarsia elegans Gran, 1912N ˟
44-Najadea gloriosa Schiller* ? holotype is drawing with too little detail to allow identificationN ˟
45HOLOphiaster formosus GranOphiaster formosus Gran, 1912N, S
46HETOphiaster hydroideus (Lohmann) Lohmann +Ophiaster hydroideus (Lohmann 1903) Lohmann, 1913M
47HOLPeriphyllophora mirabilis (Schiller) Kamptner Syracosphaera anthos (Lohmann 1912) Janin, 1987 HOLN ˟
48HETPontosphaera discopora Schiller Pontosphaera discopora Schiller, 1925N ˟
49-Pontosphaera echinophora Schiller * ? holotype is drawing with too little detail to allow identificationN ˟
50-Pontosphaera haeckelii Lohmann* ? holotype is drawing with too little detail to allow identificationN ˟
51-Pontosphaera hartmannii Schiller* ? holotype is drawing with too little detail to allow identificationN ˟
52-Pontosphaera inermis Lohmann * ? holotype is drawing with too little detail to allow identificationN ˟, M
53HETPontosphaera nana Kamptner Syracosphaera nana (Kamptner 1941) Okada & McIntyre, 1977N ˟
54-Pontosphaera nigra Schiller* not a coccolithophore: Scrippsiella acuminate (Ehrenberg 1836) Kretschmann et al. 2015N ˟
55-Pontosphaera ovalis Schiller* ? holotype is drawing with too little detail to allow identificationN ˟, M
56HOLPontosphaera pietschmannii Kamptner * ? clearly a holococcolith but holotype is drawing with too little detail to allow identificationN ˟
57HETPontosphaera steueri Kamptner* ? cf. Hymenomonas globosa (Magne 1954) Gayral & Fresnel, 1976N ˟
58HOLPoricalyptra aurisinae (Kamptner) KleijneSyracosphaera aurisinae HOL (Kamptner 1941) Keuter et al., 2021N ˟
59HETRhabdosphaera clavigera Murray & Blackman Rhabdosphaera clavigera Murray & Blackman, 1898N, M, S
60-Rhabdosphaera erinacea Kamptner * not a coccolithophore: Scrippsiella acuminate (Ehrenberg 1836) Kretschmann et al. 2015N ˟
61HETRhabdosphaera longistylis Schiller* cf. Palusphaera vandelii Lecal, 1965N ˟, M
62-Rhabdosphaera nigra Schiller* not a coccolithophore: Scrippsiella acuminate (Ehrenberg 1836) Kretschmann et al. 2015N ˟
63HETRhabdosphaera tignifer SchillerRhabdosphaera clavigera Murray & Blackman, 1898N, M, S
64-Rhabdosphaera tubulosa Schiller * not a coccolithophore: Scrippsiella acuminate (Ehrenberg 1836) Kretschmann et al. 2015N ˟
65-Rhabdothorax hispidus (Lohmann) Kamptner * ?N, M, S
66HETScyphosphaera apsteinii Lohmann Scyphosphaera apsteinii Lohmann, 1902
67HOLSyracolithus catilliferus (Kamptner) DeflandreHelicosphaera HOL catilliferus type Young & Bown (2014)N ˟
68HOLSyracolithus catilliferus (Kamptner) Deflandre Helicosphaera HOL catilliferus type Young & Bown (2014)N ˟
69HOLSyracolithus dalmaticus (Kamptner) Loeblich & TappanSyracosphaera mediterranea Lohmann 1902 HOL wettsteinii type, sensu Geisen et al., 2002N ˟
70HOLSyracolithus quadriperforatus (Kamptner) GaarderCalcidiscus leptoporus subsp. quadriperforatus (Kamptner 1937) Geisen et al., 2002 HOLN ˟
71HETSyracolithus schilleri (Kamptner) KamptnerScyphosphaera apsteinii Lohmann, 1902 HOLN ˟
72HOLSyracosphaera adriatica Schiller Helicosphaera HOL ponticuliferus type informalN ˟, M
73HOLSyracosphaera anthos (Lohmann) ZaninSyracosphaera aurisinae HOL (Kamptner 1941) Keuter et al., 2021N ˟
74-Syracosphaera bifenestrata Schiller * ? line drawing of coccosphere inadequate to determine speciesN ˟
75HOLSyracosphaera cordiformis Schiller ? cf. Sphaerocalyptra quadridentata (Schiller 1913) Deflandre, 1952M
76HOLSyracosphaera cornifera Schiller Helladosphaera cornifera (Schiller 1913) Kamptner, 1937N ˟
77HOLSyracosphaera cornus Kamptner Helicosphaera HOL catilliferus type Young & Bown (2014)N ˟
78-Syracosphaera coronata Schiller *? line drawing of coccosphere inadequate to determine speciesN ˟
79-Syracosphaera cupulifera Schiller * ? line drawing of coccosphere inadequate to determine speciesN ˟
80-Syracosphaera dentata Lohmann* ? line drawing of coccosphere inadequate to determine speciesN ˟
81-Syracosphaera grundii Schiller * ? line drawing of coccosphere inadequate to determine speciesN ˟
82HETSyracosphaera halldalii Gaarder ex Jordan & Green Syracosphaera halldalii Gaarder, in Gaarder & Hasle 1971 ex Jordan, 1994N, M
83HETSyracosphaera histrica Kamptner Syracosphaera histrica Kamptner, 1941N ˟
84-Syracosphaera lohmannii Brunnthaler * ? line drawing of coccosphere inadequate to determine speciesN ˟
85HETSyracosphaera molischii SchillerSyracosphaera molischii Schiller, 1925N, M
86-Syracosphaera ovata Schiller * ? line drawing of coccosphere inadequate to determine speciesN, M
87HETSyracosphaera pulchra LohmannSyracosphaera pulchra Lohmann, 1902N, M, S
88-Syracosphaera radiata Schiller * ? line drawing of coccosphere inadequate to determine speciesN ˟
89-Syracosphaera spinosa Lohmann* ? line drawing of coccosphere inadequate to determine speciesN ˟
90-Syracosphaera tenuis Lohmann* ? line drawing of coccosphere inadequate to determine speciesN ˟
91HETTergestiella adriatica Kamptner Tergestiella adriatica Kamptner, 1941N ˟
92HETThalassopappus pellucidus Kamptner * ? cf. Calciopappus caudatus Gaarder & Ramsfjell, 1954N ˟
93HETThorosphaera elegans OstenfeldScyphosphaera porosa Kamptner, 1967S
94HOLZygosphaera debilis Kamptner Zygosphaera amoena Kamptner, 1937N ˟
95HOLZygosphaera hellenica Kamptner Syracosphaera mediterranea Lohmann 1902 HOL hellenica type, sensu Cros et al., 2000N ˟
Table 4. List of species compiled from recent publications studying coccolithophores diversity assemblage in the Adriatic Sea. HET, HOL, POL and CER indicate life cycle stages. HET-diploid phase; HOL, NAN, POL haploid phase. Species recorded in both phases (HET and HOL) are marked in bold.
Table 4. List of species compiled from recent publications studying coccolithophores diversity assemblage in the Adriatic Sea. HET, HOL, POL and CER indicate life cycle stages. HET-diploid phase; HOL, NAN, POL haploid phase. Species recorded in both phases (HET and HOL) are marked in bold.
NTypeSpecies NameDistribution
1HETAcanthoica acanthos Schiller, 1925M
2HETAcanthoica acantifera Lochmann, 1912 ex Lochmann,1913S
3HETAcanthoica quattrospinaLohmann, 1903N, M, S
4HOLAcanthoica quattrospina Lohmann, 1903 HOLN, M
5HETAlgirosphaera robusta (Lohmann, 1902) Norris, 1984N, M, S
6HOLAlgirosphaera robusta HOL (Lohman, 1902, Norris, 1984) Dimiza et al. 2008N, M, S
7HETAlisphaera capulata Heimdal in Heimdal & Gaarder, 1981M
8HETAlisphaera extenta Kleijine, 2002N, M
9HETAlisphaera gaudiiKleijine, 2002N, M
10POLAlisphaera gaudii Kleijine, 2002 POLM
11HETAlisphaera quadrilatera Kleijne, 2002M
12HETAlisphaera unicornisOkada & McIntyre, 1977M, S
13POLAlisphaera unicornis Manton & Oates, 1980 POLM
14HOLAnthosphaera type C CrosM
15HETCalcidiscus leptoporus (Murray & Blackman, 1898) Loeblich & Tappan, 1978N, M, S
16HETCalcidiscus leptoporussubsp.quadriperforatus (Kamptner, 1937) Geisen et al., 2002M, S
17HOLCalcidiscus leptoporus subsp. quadriperforatus (Kamptner 1937) Geisen et al., 2002 HOLN, M, S
18HOLCalcidiscus leptoporus subsp. leptoporus (Murray & Blackman 1898) Loeblich & Tappan, 1978 HOLS
19HETCalciopappus caudatus Gaarder & Ramsfjell, 1954 *N
20HETCalciopappus rigidus Heimdal in Heimdal & Gaarder, 1981N, M
21HETCalciosolenia brasiliensis (Lohmann, 1919) Young in Young et al., 2003N, M, S
22HETCalciosolenia corsellii Malinverno, 2004N, M
23HETCalciosolenia murrayi Gran, 1912N, M, S
24HOLCalicasphaera blokii Kleijne, 1991N, M
25HOLCalyptrosphaera dentata Kleijne, 1991M, S
26HOLCalyptrosphaera heimdaliae Norris, 1985M
27HOLCalyptrosphaera sphaeroidea Schiller, 1913 HOLN, M
28CERCeratolithus cristatus (Kamptner 1950) CER cristatus type sensu Young et al., 2003M, S
29HETCoccolithus pelagicus (Wallich 1877) Schiller, 1930 *N
30HOLCorisphaera gracilis Kamptner, 1937N, M
31HETCyrtosphaera aculeata (Kamptner 1941) Kleijne, 1992N, M
32HETDiscosphaera tubifera (Murray & Blackman 1898) Ostenfeld, 1900M, S
33HETEmiliania huxleyi (Lohmann, 1902) Hay & Mohler, in Hay et al. 1967N, M, S
34HETEmiliania huxleyi type A (Lohmann, 1902) sensu Young & Westbroek, 1991M
35NANFlorisphaera profunda Okada & Honjo, 1973M
36NANFlorisphaera profunda var. profunda Okada & Honjo, 1973S
37HETGephyrocapsa oceanica Kamptner 1943S
38HETGladiolithus flabelatus (Halldal & Markali 1955) Jordan & Chamberlain, 1993M, S
39HOLGliscolithus amitakareniae Norris, 1985M
40HETHelicosphaera carteri (Wallich, 1877) Kamptner,1954N, M, S
41HETHelicosphaera hyalina Gaarder,1970M, S
42HETHelicosphaera pavimentumOkada & McIntyre, 1977M, S
43HOLHelicosphaera pavimentum HOL (Kamptner, 1927) sensu Young et al., 2020N, M, S
44HETHelicosphaera wallichii (Lohmann, 1902) Okada & McIntyre, 1977S
45HOLHelicosphaera HOL catiliferus type (Kamptner, 1937) sensu Young & Bown, 2014N, S
46HOLHelicosphaera HOL confusus type (Kleijne, 1991) sensu Geisen et al. 2002N, M, S
47HOLHelladosphaera cornifera (Schiller 1913) Kamptner, 1937N, M
48HOLHomozygosphaera spinosa (Kamptner 1941) Deflandre, 1952N, M
49HETHymenomonas roseola Stein, 1878M
50HETMichaelsarsia adriaticus (Schiller 1914) Manton et al., 1984 *N
51HETMichaelsarsia elegans Gran, 1912N, S
52NANNavilithus altivelum Young & Andruleit, 2006M
53HETOolithotus fragilis (Lohmann 1912) Martini & Müller, 1972S
54HETOphiaster formosus Gran, 1912M
55HETOphiaster hydroideus (Lohmann 1903) Lohmann, 1913N, M, S
56HETOphiaster minimus Manton & Oates, 1983M
57HETPalusphaera crosiae Archontikis & Young 2021M °
58HETPalusphaera vandelii Lecal, 1965 N, M
59HETPappomonas sp. Manton & OatesM
60HETPappomonas sp. type 1 Cros & Fortuno, 2002M
61HETPappomonas sp. type 3 Cros & Fortuno, 2002M
62HETPapposphaera lepida Tangen, 1972M
63HETPapposphaera sp. type 3 Cros & Fortuno, 2002M
64HETPapposphaera thomsenii Norris, 1983M
65HOLPapposphaera sp. HOL M
66HETPicarola margalefii Cros & Estrada, 2004M
67NANPileolosphaera longistirpes Meier, Kinkel & Young 2014N, M *
68POLPolycrater sp. ladle-like Cros & Fortuño, 2002M
69HETPontosphaera syracusana Lohmann, 1902S
70HOLPoritectolithus poritectus (Heimdal in Heimdal & Gaarder 1980) Kleijne, 1991M
71HETReticulofenestra sessilis (Lohmann 1912) Jordan & Young, 1990S
72HETRhabdosphaera clavigera Murray & Blackman, 1898M, S
73HETRhabdosphaera clavigera var. stylifera (Lohmann, 1902) Kleijne & Jordan, 1990N, M
74HETRhabdosphaera xiphos (Deflandre & Fert 1954) Norris, 1984N, M, S
75HETScyphosphaera apsteinii Lohmann, 1902S
76HOLScyphosphaera apsteinii Lohmann, 1902 HOLM
77HOLSphaerocalyptra cf. adenensis of Cros & Fortuño 2003M
78HOLSphaerocalyptra sp. 1 Cros & Fortuño, 2002M
79HOLSphaerocalyptra sp. 2 Cros & Fortuño, 2002M
80HOLSphaerocalyptra sp. 3 Cros & Fortuño, 2002M
81HOLSphaerocalyptra sp. 5 Cros & Fortuño, 2002M
82HOLSphaerocalyptra sp. 6 Cros & Fortuño, 2002M
83HOLSyracolithus sp. type A Kleijne, 1991M
84HETSyracosphaera ampliora Okada & McIntyre, 1977N, M
85HETSyracosphaera anthos (Lohmann, 1912) Janin,1987M, S
86HOLSyracosphaera anthos (Lohmann 1912) Janin, 1987 HOLM, S
87HETSyracosphaera arethusae (Kamptner, 1941) Triantaphyllou et al. 2016N, M
88HOLSyracosphaera arethusae (Kamptner 1941) Triantaphyllou et al. 2016 HOLN, M, S
89HOLSyracosphaera aurisinae HOL (Kamptner 1941) Keuter et al., 2021M
90HETSyracosphaera azureaplaneta Young, 2018M
91HETSyracosphaera bannockii (Borsetti & Cati 1976) Cros et al., 2000N, M, S
92HOLSyracosphaera bannockii (Borsetti & Cati 1976) Cros et al., 2000 HOLM
93HETSyracosphaera corolla Lecal, 1966 emend Young et al. 2018N, M
94HETSyracosphaera dilatata Jordan et al., 1993N, M
95HOLSyracosphaera elevata Archontikis, Young & Cros 2020 HOLM *
96HOLSyracosphaera gaarderae (Borsetti & Cati 1976) Keuter, Young & Frada 2019 HOLM
97HETSyracosphaera halldaliiGaarder, in Gaarder & Hasle 1971 ex Jordan, 1994M, S
98HOLCalyptrolithina divergens (Halldal & Markali 1955) Heimdal, 1982, Triantaphyllou et al. 2004 (Syracosphaera halldalii HOL)N, M
99HETSyracosphaera histricaKamptner, 1941N, M, S
100HOLSyracosphaera histrica Kamptner, 1941 HOLN, M
101HOLSyracosphaera lafourcadii (Lecal 1967) Archontikis et al., 2020 HOLM
102HETSyracosphaera lamina Lecal-Schlauder, 1951M
103HETSyracosphaera marginaporataKnappertsbusch, 1993N, M
104HOLSyracosphaera marginiporata Knappertsbusch 1993 HOLM
105HETSyracosphaera mediterraneaLohmann, 1902N, M, S
106HETSyracosphaera mediterraneavar. binodata (Kamptner, 1927) sensu Triantaphyllou et al. (2016) & Young et al. (2020)S
107HOLSyracosphaera mediterranea Lohmann 1902 HOL gracillima type, sensu Geisen et al., 2002M
108HOLSyracosphaera mediterranea Lohmann 1902 HOL hellenica type, sensu Cros et al., 2000M
109HOLSyracosphaera mediterranea Lohmann 1902 HOL wettsteinii type, sensu Geisen et al., 2002N, M
110HETSyracosphaera molischiiSchiller, 1925N, M, S
111HETSyracosphaera molischii type 1 (Schiller, 1925) sensu Young et al. 2003N
112HETSyracosphaera molischii type 2 (Schiller, 1925) sensu Young et al. 2003N
113HETSyracosphaera molischii type 3 (Schiller, 1925) sensu Young et al. 2003M
114HETSyracosphaera molischii type 4 (Schiller, 1925) sensu Young et al. 2003M
115HOLSyracosphaera molischii Schiller, 1925 HOLN, M
116HETSyracosphaera nana (Kamptner, 1941) Okada & McIntyre, 1977M
117HOLSyracosphaera nana (Kamptner 1941) Okada & McIntyre, 1977 HOLM
118HETSyracosphaera nodosa Kamptner, 1941N, M
119HETSyracosphaera orbiculus Okada & McIntyre, 1977M
120HOLSyracosphaera origami (Cros & McGrane 2014) Archontikis et al. 2020 HOLM
121HETSyracosphaera ossa (Lecal 1966) Loeblich & Tappan, 1968M
122HETSyracosphaera ossa type 1 (Lecal 1966) Loeblich & Tappan, 1968, of Young et al. 2003M
123HETSyracosphaera ossa type 2 (Lecal 1966) Loeblich & Tappan, 1968, of Young et al. 2003N, M
124HOLSyracosphaera periperforata (Kleijne 1991) Archontikis, Young & Cros 2020 HOLM
125HOLSyracosphaera periperforata Kleijne 1991 var. periperforata HOL sensu Archontikis et al., 2020N, M
126HOLSyracosphaera periperforata Klejine 1991 var. cylindrata HOL sensu Archontikis et al., 2020M
127HETSyracosphaera prolongata Gran 1912 ex Lohmann, 1913M, S
128HETSyracosphaera protrudens Okada & McIntyre, 1977N, M
129HETSyracosphaera pulchraLohmann, 1902N, M, S
130HOLSyracosphaera pulchra Lohmann, 1902 HOL oblonga type, sensu Young et al., 2003N, M, S
131HOLSyracosphaera pulchra Lohmann, 1902 HOL pirus type, sensu Young et al., 2003M
132HOLSyracosphaera rotaconica Archontikis, Young & Cros, in Archontikis et al., 2020 HOLM
133HETSyracosphaera rotula Okada & McIntyre, 1977M
134HETSyracosphaera strigilis (Gaarder 1962) Šupraha et al., 2018N, M
135HOLSyracosphaera strigilis (Gaarder, 1962) Supraha et al., 2018 HOLN, M
136HETSyracosphaera tumularis Sánchez-Suárez, 1990M
137HETTergestiella adriatica Kamptner, 1941N
138HETUmbellosphaera tenuis (Kamptner 1937) Paasche in Markali & Paasche, 1955M, S
139HETUmbilicosphaera foliosa (Kamptner 1963) Geisen in Sáez et al., 2003S
140HETUmbilicosphaera sibogae var sibogue (Weber—van Bosse 1901) Gaarder, 1970M, S
141HOLZygosphaera amoena Kamptner, 1937M
°Palusphaera sp. Cros & Fortuño, 2002—published in Skejić et al. 2018. has been confirmed as Palusphaera crosiae Archontikis & Young 2021. * personal observation—unpublished species.
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Godrijan, J.; Arapov, J.; Skejić, S.; Bužančić, M. Back to Basics: Revision of Coccolithophore Species List in the Adriatic Sea. Water 2023, 15, 603. https://doi.org/10.3390/w15030603

AMA Style

Godrijan J, Arapov J, Skejić S, Bužančić M. Back to Basics: Revision of Coccolithophore Species List in the Adriatic Sea. Water. 2023; 15(3):603. https://doi.org/10.3390/w15030603

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Godrijan, Jelena, Jasna Arapov, Sanda Skejić, and Mia Bužančić. 2023. "Back to Basics: Revision of Coccolithophore Species List in the Adriatic Sea" Water 15, no. 3: 603. https://doi.org/10.3390/w15030603

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