Falsolikanella Campanensis (Azéma and Jaffrezo, 1972) Granier, 1987 Revisited on Type Material, Evidence of Polyphysacean Nature (Green Algae)

Falsolikanella campanensis (Azéma and Jaffrezo, 1972) Granier, 1987 revisited on type material, evidence of polyphysacean nature (green algae)

Filippo Barattolo,1 Nicola Carras,2 Marc André Conrad,3 and Rajka Radoičić4

1Dipartimento di Scienze della Terra dell’Ambiente e delle Risorse, Università degli Studi di Napoli “Federico II,” Complesso universitario di Monte Sant’Angelo, via Cintia, 21 - 80126 Napoli, Italy <ﬁ[email protected]> 2I.G.M.E., Spirou Loui 1, Olympic Village, Acharnes, Greece 3Chemin de Planta 71–1223 Cologny, Switzerland 4Kralja Petra 38, 11158 Stari Grad, Beograd, Serbia

Abstract.—The genus Falsolikanella, introduced by Granier (1987), was based on the basal Cretaceous species Likanella campanensis Azéma and Jaffrezo (1972) and assigned to the tribe Diploporeae (Pia, 1920) emend. Güvenc, 1979 within the green alga order Dasycladales. Later, other species were assigned to Falsolikanella. Sections of the type specimens of Likanella campanensis are reviewed. They show that in this species, the arrangement of the laterals is not metaspondyl, but typical of the genus Actinoporella (Gümbel in Alth, 1882) emend Conrad, Praturlon, and Radoičić, 1974, with coronae arising from a single primary lateral. Therefore, the species is assigned to the genus Actinoporella within the tribe Acetabularieae Decaisne, 1842, family Polyphysaceae Kützing, 1843, and the generic attribution of other species previously assigned to Falsolikanella is discussed.

Introduction An alternative structural view of L. campanensis was presented by Granier (1987). The whorls are simple. They consist Located in the eastern Betic Cordillera, southern Spain, the type of clusters (tufts) of three, all similar first-order fertile laterals, level of Actinoporella campanensis new combination is dated by lined up along the axis of the thallus and arising from a proximal Granier (1987) from the Berriasian or basal Valanginian. Other vestibule. Separately, the author noticed the special pattern of cal- specimens in thin section useful to detect the structure of the alga cification, similar to Clypeina jurassica Favre in Favre and Rich- are illustrated by Sokačand Velić(1978a) from the Valanginian ard, 1927.BasedonL. campanensis, the genus Falsolikanella was of Croatia, while other specimens are reported in other areas, introduced with the following diagnosis (translated from French): including Spain, France, Romania, Croatia, Serbia, Slovakia, Turkey, and Switzerland. In the following chronological Alga with a continuous cylindrical main axis bearing sim- account, we will pay special attention to the consecutive struc- ple verticils, more or less spaced out, protruding, composed tural interpretations and taxonomy of this species, which, of ramifications only of first order, grouped in tufts, and although seemingly seldom abundant, is nevertheless easily inserted in small number on a simple vestibule (metaspon- identifiable. dyl type, with a vestibule). (Granier, 1987, p. 208) In 1972, Azéma and Jaffrezo established Likanella campanensis. The diagnosis and the reconstruction refer to a cylin- Falsolikanella was assigned to the tribe Diploporeae by drical axis bearing twin whorls of all equal, pyriform, and implicit analogy with the arrangement of the laterals in the Tri- diverging laterals. The species is assigned to the Permian assic type species of the tribe, Diplopora annulata (Schafhäutl, genus Likanella Milanović, 1966, whose original diagnosis, 1853) Schafhäutl, 1863. potentially subject to various interpretations, is as follows: According to Berger and Kaever (1992, p. 47), the genus Falsolikanella may not belong to the Diploporeae. The thallus is composed of loosely connected cylindrical The structure of the alga is here reexamined. The analysis of segments, which at their lower end possess 3 whorls of specimens in thin section of the type material allows us to pro- branches. The long branches which at the top are open spect a new interpretation of F. campanensis (Azéma and Jaf- and non-ramified have separate calcareous walls. In the frezo, 1972), and consequently the validity of the genus walls of the main stem and of branches there are to be Falsolikanella is discussed. found fine pores. (Milanović, 1966,p.9) Materials and methods Later, Sokačand Velić(1978b) implicitly transferred L. campanensis to the genus Selliporella Sartoni and Crescenti, This study is based on the type material of Likanella campanen- 1963 emend. Barattolo, De Castro, and Radoičić, 1988. sis. It consists of three thin sections labeled as follows. FSL Although not formally emended, Selliporella is newly described 420001: Likanella companensis, lame 3486, holotype (Azéma and assigned to the tribe Diploporeae. and Jaffrezo, 1972,pl.1,fig. 1), Puig Campana (Alicante, 593 Downloaded from https://www.cambridge.org/core. University of Athens, on 06 Oct 2021 at 18:01:03, subject to the Cambridge Core terms of use, available at https://www.cambridge.org/core/terms. https://doi.org/10.1017/jpa.2018.108 594 Journal of Paleontology 93(4):593–611

Spain), Berriasien. FSL 420002: Likanella companensis,lame 3486-3 (Azéma and Jaffrezo, 1972,pl.1,ﬁg. 5), Puig Campana (Alicante, Spain), Berriasien. FSL 420003: Likanella campanensis, lame 3486-4, Puig Campana (Alicante, Spain), Berriasien. Material with Praturlonella danilovae Radoičić, 1975 and Actinoporella kukoci Radoičić, 1974 belongs to the Rajka Radoičićcollection (thin sections labeled RR).

Repositories and institutional abbreviations.—Types examined in this study are deposited in the following institutions: (1) Likanella campanensis is in the collections of the University of Lyon, UCB Lyon 1 (inventory FSL 420001 = slide 3486; FSL 420002 = slide 3486.3; FSL 420003 = slide 3486.4). (2) Praturlonella danilovae Radoičić, 1975 and Actinoporella kukoci Radoičić, 1974 (labeled RR) are housed in R. Radoičićprivate collection at Kralja Petra 38, 11158 Stari Grad, Beograd, Serbia.

Structural constraints

According to Azéma and Jaffrezo (1972), the alga bears spaced whorls, each composed of two rows of 12 laterals set in alterna- tion (here indicated as two-rows model, Fig. 1). The swollen outer part of the laterals is responsible for the downward and upward bending of the laterals in the lower and upper rows, respectively. The two authors are elusive on the significance of the two rows. Possibilities are as follows: (1) the rows are sim- ply due to the heterocline (upward and downward) arrangement of the laterals; however, the whorl is single, making the number of laterals per whorl correspond to the sum of the two rows (i.e., w = 24); and (2) contrariwise, the two rows result from the presence of two separate whorls; in this case the value of w is 12. Figure 1 illustrates the second possibility. Drawing on new material originating from the type locality, Granier (1987) anticipated the presence of a tuft of three vertically aligned pores (Granier, 1987, p. 207, pl. 6, fig. d). For this author, the structure corresponds to a metaspondyl tuft. In the table of biometrical values, the author did not indicate the number of laterals per whorl (Granier, 1987, p. 209). In the reconstruction (Granier, 1987, fig. 1b), the displayed number of vestibules is eight, corresponding to the value of w, each vestibule bearing three vesiculiferous, fertile primary laterals distally arranged chiefly in two rows as usual, such as already Figure 1. The two-rows model: organization of Likanella campanensis sensu interpreted by Azéma and Jaffrezo (1972). This model is here Azéma and Jaffrezo (1972). (1) Axial view with traces of the oblique and tangen- designated as the metaspondyl model (Fig. 2). Considering tial sections; (2) oblique section; (3) tangential section. such a threefold structure, this metaspondyl model implies that the pores are proximally eight in number, distally corresponding to 24 in number. In case of arrangement in two rows, each row Number of proximal pores per whorl (w).—Even taking into should contain 12 vesiculiferous laterals. consideration the metaspondyl model, if a verticil is cut transversally (Fig. 5), the number of innermost pores Number and arrangement of pores per tuft.—The occurrence of corresponds to the value of w. In the type material, the values of the threefold structure put forward by Granier (1987)(Figs. 3, 4) w range from 16 to 22. As to the estimation of w from oblique in the type material is herein confirmed. As illustrated by this sections, several methods are known (Pia, 1920;DeCastro, author, three pores are present, axially aligned close to the 1997). Based on the ‘couple of pores’ method (see central stem (Figs. 3.5, 4.1). As shown by Granier (1987,pl.6, Supplemental Data), the values range from 18 to 25, comparable fig. d), the tufts are laterally very close each other (Fig. 4.2, to 16–22. Likewise, this calculation is confirmed in 4.6–4.8); moreover, they are not completely aligned in a single proximal-tangential sections (e.g., Fig. 4.6, 4.8), resulting in the row, but somewhat scalloped, shifted upward and downward distance between tufts (Δtuft)toaverage76µ(55–100 µ) and the by lateral compression. estimated value of w to come to about 20 (w = d threefold* π/Δtuft).

Downloaded from https://www.cambridge.org/core. University of Athens, on 06 Oct 2021 at 18:01:03, subject to the Cambridge Core terms of use, available at https://www.cambridge.org/core/terms. https://doi.org/10.1017/jpa.2018.108 Barattolo et al.—Revision of Cretaceous dasycladalean algae 595

p. 126–127). Although limited, the type material, allows to estimate the total number of distal pores (wdist)to19–24. A method based on trigonometry aimed at calculating the inferred value of w and wdist from oblique sections is presented in the Supplemental Data. When applied to the holotype (Fig. 3.6), the estimated value of wdist is 22 (see Table 1 in Supplemental Data).

Metaspondility versus euspondility—geometrical constraints

It is usually easy to distinguish metaspondyl algae from euspondyl algae with heterocline laterals. However, under certain conditions this distinction may prove problematic, and some sections can lead to erroneous conclusions if taken alone, out of the context of accompanying sections. Figure 6.1 shows a typical axial section referable to a metaspondyl or euspondyl alga. The calcareous skeleton is poorly calcified, slightly intusannu- lated at the junction of the laterals and the central stem, which typically occurs when the laterals are closely packed. As displayed by the figure, the same axial section can be interpreted as metaspondyl (Fig. 6.2) or euspondyl (Fig. 6.4). The longitudinal section is not the only one to be problematic, also a tangential s. Subtransversal-oblique sections depicted in Figures 1.2 and 2.2 are particularly indicative to discern between the metaspondyl and euspondyl models. Such sections are present in Figure 3.3 and Granier (1987,pl.7,figs. b, c). In the two-rows model (Fig. 1.2) sensu Azéma and Jaffrezo (1972), there is no trace of proximal circular pores (no tuft of three pores vertically aligned), while in the metaspondyl model sensu Granier (1987) and Figure 2.2, the number of proximal pores is far fewer than observed (8 versus 24) while the value of the number of distal pores per row is the same (i.e., 12). If a w = 24 metaspondyl model sensu Granier (1987) is applied (Fig. 7), to correct the proximal pore framework, the number of distal pores (wdist) inevitably increases too much (72 instead of 24) as does the number of rows (at least three). As assessed in the preceding, wprox =16–25 and wdist ≅19– 24, which means that the wdist/wprox ratio is equal to one instead of three, as in the metaspondyl model. Consequently, the two rows of 12 pores arrangement is attained by the upward and downward divergence of the ‘gametangia,’ due to their outward flaring. In this context, the proximal tuft of three pores has no Figure 2. The metaspondyl model: organization of Falsolikanella campanen- fl sis sensu Granier (1987). (1a) Axial view with trace of an oblique section; (1b) in uence on the distal number of laterals. Such a paradox is perspective view of the tufts cut at branching level; (1c) perspective view of the explained considering that two out of the three pores correspond vesiculiferous laterals arranged in two rows; triangles indicate the tufts; (2) to the short, ‘blind’ structures, visible in longitudinal section oblique section. (Fig. 3.5). The corresponding new model is shown in Figures 8 and 9, where it is illustrated in subtransversal-oblique section (Fig. 8.2), in tangential section (Fig. 8.3), and in almost transver- The diameter at the threefold level (d threefold) it is about 1.35 times the inner diameter (d = 0.37 mm in average, then d sal section (Fig. 9.2). threefold = 0.50 mm). To sum up, the number of pores in a whorl (w) ranges within 16–25. Assignment at genus level

‘ ’ Number of distal pores per whorl (wdist).—From the type The two short, blind structures are depicted in Figure 10.An material, the arrangement of the large distal pores prevalently interpretation of transversal, slightly oblique sections is given set in two rows can be conﬁrmed. The number of distal pores in the Systematic paleontology in the section describing Actino- per row is 10–12, according to Azéma and Jaffrezo (1972, porella campanensis, under Remarks (Fertile laterals and

Figure 3. Actinoporella campanensis (Azéma and Jaffrezo, 1972) n. comb. Type material. (1) Subtransversal section; upper coronae appear as elliptic pores between gametophores (white arrow); other equivalent pores show the tendency to split in two circular smaller pores aligned radially (black arrows), possibly corresponding to corona protuberances like in extant polyphysaceans; thin section n. 3486.3 (specimen already figured by Azema and Jaffrezo, 1972, pl. 1, fig. 5); (2) transversal section, thin section n. 3486.3; (3) oblique section of an incomplete specimen, thin section n. 3486.4; (4) oblique section; notice the narrow circular pore (arrow) close to the thin calcification of the central stem corresponding to upper corona, thin section n. 3486 (specimen already figured by Azéma and Jaffrezo, 1972, pl. 1, fig. 4); (5) longitudinal section showing a fertile lateral bending downward (left) and another one directed upward (right); notice the arrows showing the basal part, the upper and lower coronae (left), and the basal part and lower corona (right); thin section n. 3486; (6) oblique section, thin section n. 3486 (specimen already figured by Azéma and Jaffrezo, 1972, pl. 1, fig. 1, the holotype). (1, 2) Scale bar shown in (1) = 0.25 mm; (3–6) scale bar shown in (3) = 0.25 mm.

coronae). Polyphysacean coronae have been interpreted as: (1) the original aragonitic structure changed into blocky calcite, also expansions of the central stem, (2) primary laterals together occurs in other fossil taxa. A generic, if not suprageneric, with the basal part, or (3) secondary laterals (see review in significance of the mineralogy of the calcareous skeleton is Dumais and Harrison, 2000). Here, coronae are considered as assigned by Simmons et al. (1991). But in the literature, a corresponding to secondary laterals, concurring with Valet’s much broader consensus assigns a species-level rank to this (1968,p.62–63) interpretation. character, as shown by various examples found in different From a taxonomic perspective, the occurrence of upper and fossil genera, such as Salpingoporella (S. muhelbergii lower coronae arising from a short basal lateral, and large outer [Lorenz, 1902] Pia, 1917, the type species, originally pores corresponding to fertile laterals serving for reproduc- aragonitic, and S. dinarica Radoičićin Granier and Doloffre, tion (gametophores), make F. campanensis match the genus 1993, fibrous calcite), Clypeina (C. marginoporella Michelin, Actinoporella. Conversely, the presence of heterocline euspon- 1840, aragonitic, and C. sulcata [Alth, 1881] Granier and dyl laterals, with gametophores markedly swollen outward Brun, 1991, fibrous calcite), Pseudoclypeina (P. cirici (pear-like), a calcareous skeleton made of fibrous calcite, and Radoičić, 1975 and P. distomensis Barattolo and Carras, 1990, spaced whorls, make F. campanensis markedly different from fibrous calcite, and P. farinacciae Radoičić, 1975, aragonitic), Actinoporella podolica (Alth, 1878), the type species of the and Falsolikanella (F. campanensis, fibrous calcite, and F. genus. danilovae Granier et al., 1999, aragonitic).

Heterocline laterals.—Several factors, such as the spacing of Spacing of the whorls.—In families Triploporellaceae and the whorls, size of laterals (p), number (w) of laterals per Dasycladaceae, distance between the whorls, close versus whorl, and calcification may condition the isocline versus spaced, is an important character since directly related to the heterocline arrangement. In extant genera Acetabularia and general aspect of the alga, reflecting the organization of the Parvocaulis, both isocline (e.g., A. acetabulum [Linnaeus, 1758] thallus. Close whorls usually occur in taxa provided with an Silva, 1952 and P. parvula [Solms-Laubach, 1895]Bergeretal., external cortex and a strong calcareous skeleton. Close versus 2003) and heterocline (e.g., A. peniculus and P. clavata spaced whorls is one of the characters used to macroscopically [Yamada, 1934] Berger et al., 2003) species exist (e.g., Valet, discern Dasycladus from Batophora (Valet, 1969, p. 578, 1969;Bergeretal.,2003). In our opinion, heteroclinity, taken 584). The problem arises when applied to the Polyphysaceae. solely, is not enough to support the ordering of taxa at the level In this family, most extant species are single whorled at of the genus, especially in family Polyphysaceae. maturity, with umbrella-like thalli, while Halicoryne is the only extant genus where comparisons between the relative Gametophores markedly swollen outward (pear-like).— distance between whorls were carried out. The h/D ratio (h = Sometimes, authors use the shape, more or less swollen, of distance between whorls, D = outer diameter of the thallus) the laterals to differentiate two genera (e.g., Suppiluliumaella was estimated in H. spicata (Kützing, 1863) Solms-Laubach, from Triploporella in Barattolo, 1983). Such a character, taken 1895 by Valet (1968, pl. 20, fig. 4) and in H. wrightii Harvey, as an individual element of differentiation at the level of the 1859 by Valet (1969, pl. 147, fig. 1). In H. spicate, the h/D genus, looks problematic. The shape of gametophores in ratio is relatively large compared to H. wrightii (i.e., 0.40 extant genera Acetabularia and Parvocaulis is highly variable versus 0.20, respectively). The same parameter in (Berger and Kaever, 1992; Berger et al., 2003; Fig. 11). Falsolikanella campanensis (Fig. 10.1) and Actinoporella Species such as Acetabularia acetabulum, A. crenulata podolica (Fig. 10.2) is even more emphasized (0.56 versus Lamouroux, 1816, and A. farlowii Solms-Laubach, 1895 hold 0.15). As a matter of fact, Actinoporella podolica exhibits an slender gametophores, but in Acetabularia peniculus, they are extreme variability in size and distance between whorls, as ‘vesiculiferous’ (Valet, 1969, p. 619). The same happens in shown by the topotype specimens illustrated by Conrad et al. genus Parvocaulis, with Parvocaulis exigua (Solms-Laubach, (1974, fig. 6b), where the h/D ratio is around 0.50, a value 1895) Berger et al., 2003 holding elongated (acrophorous) also found in Actinoporella campanensis. gametophores and P. pusilla (Howe, 1909) Berger et al., 2003 Bearing in mind such a close examination of the taxonomic holding pear-shaped gametophores. The same, but characters, in our opinion the moderately phloiophorous shape, intraspecific, variability is also very high in extant taxa such isocline gametophores, and rather closer whorls in Actinoporella as Acetabularia crenulata (Berger and Kaever, 1992, p. 144). podolica, as opposed to the markedly flared out, heterocline laterals and spaced whorls in F. campanensis, are characters of spe- Calcareous skeleton made of fibrous calcite.—Such a structure cific weight only. At genus level, and for the same reason, there of the calcareous wall (fibrous calcite), even if less common than are no strong arguments to separate Falsolikanella from

Figure 4. Actinoporella campanensis (Azéma and Jaffrezo, 1972) n. comb. Type material. (1) Tangential section, thin section n. 3486; notice the threefold fertile laterals (arrow); (2) oblique section, thin section n. 3486.3; (3) oblique section, thin section n. 3486; (4) oblique-tangential section, thin section n. 3486.3; (5) longitudinal-oblique section, thin section n. 3486 (specimen already figured by Azéma and Jaffrezo, 1972, pl. 1, fig. 9); (6) tangential section; thin section n. 3486 (specimen already figured by Azéma and Jaffrezo, 1972, pl. 1, fig. 10); notice two adjacent pores not separated by calcareous wall (arrow); (7) tangential section, thin section n. 3486 (specimen already figured by Azéma and Jaffrezo, 1972, pl. 1, fig. 8 left); (8) tangential section, thin section n. 3486; (9) tangential section; thin section n. 3486.3. Scale bar = 0.25 mm.

Actinoporella. Consequently, Falsolikanella campanensis is (e.g., Actinoporella and Hamulusella) genera. In extant ascribed to the genus Actinoporella, and the genus Falsolika- acetabulariacean algae, fertile laterals hold further, sterile laterals nella Granier, 1987 is dealt with as a junior synonym of Actino- arising from the corona, but no terminal second and perhaps porella Gümbel, 1882. higher-order laterals, such as in the fossil Pseudoclypeina. Consequently, genera with fertile laterals bearing terminal Systematic paleontology ramifications must be separated from genera with crown-like proximal structures included in tribe Halicoryneae. Certainly, Order Dasycladales Pascher, 1931 the most remarkable character of Halicoryne is the occurrence of Family Polyphysaceae Kützing, 1843 alternating sterile and fertile whorls, but this character is also visible in Acetabularia crenulata (Berger and Kaever, 1992, Systematics at tribe level.—The family Polyphysaceae is p. 144), resulting that the only possibly distinguishing character traditionally subdivided in three tribes: Acetabularieae is the presence of coronae: upper or upper and lower coronae in Decaisne, 1842, Halicoryneae Valet, 1969, and Clypeineae tribe Acetabularieae and only more or less evident lower corona (Elliott, 1968) Bassoullet et al., 1979 (Deloffre, 1988). in the Clypeineae. At this point, these two tribes are herein The tribe Acetabularieae includes genera presenting a more defined as follows: or less calcified thallus, with a disc at top of the axis, made of Tribe Acetabularieae: thallus cylindrical, umbelliform, or reproductive chambers (definition in Deloffre, 1988: “Thalle with spaced-out whorls of fertile rays. Upper corona only, or plus ou moins calcifié. Disque reproducteur au sommet de upper and lower coronae. Genera ascribed: Acetabularia, Parvo- l’axe formé par des loges reproductrices”). According to Valet caulis, Chalmasia, Halicoryne, Orioporella, Actinoporella. (1969, p. 578), the tribe Halicoryneae is defined as follows Tribe Clypeineae: thallus cylindrical, umbelliform, or with (translated from French): “Plants with numerous alternating spaced-out fertile laterals. Possible presence of variously sterile and fertile verticils.” The last tribe, Clypeineae, is defined marked lower corona. Genera ascribed: Clypeina, Hamulusella. by Deloffre (1988, p. 182) with the following diagnosis: “Rami- fications primaires, secondaires possibles et de troisième ordre éventuelles; verticilles espacés saillants; présence éventuelle Tribe Acetabularieae Decaisne, 1842 de corona; cavité axiale cylindrique” (free English translation: Genus Actinoporella Gümbel, 1882, emended Primary branches, possibly secondary branches, tentatively — third-order branches; corona possibly present; axial cavity Type species. Actinoporella podolica (Alth, 1878) cylindrical.) Emended diagnosis.—Thallus simple, unbranched, bearing Remarks.—Even considering that the reproductive function of the spaced-out whorls of short primary laterals. Second-order laterals of fossil taxa is widely speculative, such a definition makes laterals are three in number: one upper and one lower corona, the Clypeineae tribe a catchall taxon since it aggregates both arising from the proximal part of the primary lateral, cladospore (e.g., Pseudoclypeina) and choristospore lato sensu and one large gametophore arising from the tip of the primary lateral.

Taxonomic position at higher rank.—Deloffre (1988) ascribed Actinoporella to the tribe Clypeineae. In this paper, drawing on the preceding discussion showing the presence of lower and upper coronae, compliant with Génot et al. (2008), the genus Actinoporella is assigned to the tribe Acetabularieae.

Remarks.—The paper of Conrad et al. (1974) includes a historical record of Actinoporella, a synonymy (Verticilloporella Raviv and Lorch, 1970 is dealt with as a junior synonym), reproductions of key, historically important illustrations issued by Alth (1878) and Pia (1920), an emended diagnosis, similarities with the extant Polyphysaceae, and a review study of Actinoporella podolica based on Figure 5. Transversal sections across the metaspondyl model (organization of topotype specimens collected at Tanutinska Gora, near Falsolikanella campanensis sensu Granier, 1987). (1) Cut just above the ‘vestibule’;(2) cut in correspondence with the ‘vestibule’;(3) cut just below the ‘ves- Nizniow city, now in Ukraine. The emended diagnosis was as tibule.’ In all cuts, the number of expected proximal pores is the same (i.e., eight). follows: “Cylindrical stalk bearing regularly spaced, disc-like

Figure 6. Metaspondyl versus euspondyl arrangement. (1) Axial section of an ideal whorl, calcification (black); (2) metaspondyl model of (1) in axial view; black = calcification; (3) tangential cut marked by vertical line in (2); (4) euspondyl laterals model of (1); black = calcification; (5) tangential section marked by vertical line in (4). (3) and (5) are very similar to each other, but in the central part of (3), tufts can be still perceived.

whorls of branches. More or less marked outgrowths, or (1993). Basionym: Clypeina durandelgai. Tithonian–Berria- excrescences occur at the branches’ proximal ends. These sian. Spaced-out whorls of laterals bearing in their lower part outgrowths develop upper and/or lower coronas surrounding conspicuous proximal outgrowths evoking a corona. Transferred the stalk” (Conrad et al., 1974, p. 4). to Hamulusella by Barattolo (1998). Compared to the previous diagnosis, somewhat outdated ?Actinoporella geredeensis Farinacci and Radoičić, 1991, terms such as ‘branches’ and ‘outgrowths’ are respectively Berriasian. Upper and lower proximal outgrowths. Laterals replaced by ‘laterals’ and ‘corona,’ the major difference calling strongly tilted. The questionable assignment to Actinoporella for the presence of coronae arising from and located above as is original. well as below the primary lateral, a character considered at the Actinoporella guembeli Alth, 1882, a junior synonym of level of the genus. Other characters, such as expansions (vesti- Actinoporella podolica according to Pia (1920) and Conrad bules) of the axis of the plant, whorls made of iso- or heterocline et al. (1974). laterals, dimensions, and pattern of calcification, are less import- Actinoporella fragilis (Conrad, 1970) Granier, 1994, late ant for the systematics. Hauterivian–early Aptian. Laterals closely packed, strongly In this paper, three species, A. podolica, A. campanensis tilted, bearing a single, upper proximal feature suggesting a n. comb., and A. kukoci are assigned to Actinoporella, but in corona. In this paper, the original name, Pseudoactinoporella the literature, no less than 15 other species were assigned, fragilis (Conrad, 1970) Conrad and Peybernés, 1976,is although sometimes with doubt, to this genus. A comprehen- upheld. sive review of all of them is beyond the scope of this paper. Actinoporella israelensis Raviv and Lorch, 1992, Callo- Updating a list issued by Conrad et al. (1974), an alphabetical vian. A nomen nudum, close if not a synonym of Salpingopor- list of species assigned to Actinoporella in the literature ella grudii Radoičić, 1975. follows. As for the genus Pseudoactinoporella Conrad, Actinoporella jaffrezoi Granier, 1994, Berriasian–Hauteri- 1970, reference is made to a review issued by Bucur et al. vian. Tentative assignment to Praturlonella, pending a revision. (2012). Actinoporella krymensis Maslov, 1958, nom. nud., Late Actinoporella durandelgai Jaffrezo and Fourcade ex Jaf- Jurassic. Possibly a Clypeina according to Conrad et al. frezo in Bassoullet et al., 1978, name fide Granier and Deloffre (1974). Not properly typified.

mantle and probably are not genuine coronae. The attribution to Actinoporella remains open. Actinoporella podolica (Alth, 1878) Alth, 1882, type species of Actinoporella. Basionym: Gyroporella podolica. Titho- nian–early Barremian. See Conrad et al. (1974) for details. Actinoporella sadalanensis (Elliott, 1977) Granier, 1994. Hamulusella sadalanensis, the basionym, is upheld in this paper. Paleocene. Widely spaced whorls of laterals showing a short proximal portion below the connection with the axial stem. See Barattolo (1998) for a discussion. Actinoporella? silvaeregis—Basionym: Pseudoactinopor- ella silvaeregis Bucur, 1981.Bucur(1992): questionable assignment to Pseudoactinoporella. Bucur (1999): questionable assignment to Falsolikanella.Granier(1994): questionable assignment to Actinoporella.Barremian–early Aptian. Notice: no upper and/or proximal features evoking coronae occur, but horizontal (transversal) proximal swellings of the laterals. Actinoporella sulcata Alth, 1882. Basionym: Clypeina sulcata. Quoted by some authors as a senior synonym of Clypeina jurassica Favre in Favre and Richard, 1927. The transfer to Cly- peina, ﬁrst made by Bassoullet et al. (1978, p. 32), is debatable. See also Conrad et al. (1974). Based on the preceding analysis of taxonomic criteria (assignment at genus level), F. campanensis is revised and transferred to the genus Actinoporella, whose systematics is emended. Currently, the genus comprises three species, formally valid, A. podolica, A. kukoci, and A. campanensis. Actinoporella kukoci is the youngest species (Eocene). In Figure 12, some specimens from the type material are supplied, showing the typical characters of the genus.

Actinoporella campanensis (Azéma and Jaffrezo, 1972) new combination Figures 3, 4

1972 Likanella campanensis Azéma and Jaffrezo, p. 126, pl. 1, figs. 1–12. 1972 Likanella campanensis; Fourcade et al., p. 235, pl. 2, fig. 9. 1974 Likanella campanensis; Jaffrezo, p. 28, pl. 2, figs. 13, 14. 1977 Likanella campanensis; Conrad, p. 199. Figure 7. The metaspondyl model with w = 22. (1) Axial view with three rows of 22 laterals; (2) oblique section corresponding to oblique line in (1); (3) tangen- 1978 Likanella? campanensis; Barattolo, p. 9. tial section marked by vertical line in (1). 1978 Likanella campanensis; Bassoullet et al., p. 141, pl. 16, figs. 4–6, bibliographic review. 1978 Likanella campanensis; Radoičić, p. 189, pl. 1, 2. Actinoporella kukoci Radoičić, 1974, Eocene. Similar to A. 1978a Selliporella campanensis; Sokačand Velić, pl. 5. podolica but usually larger, with a larger number of laterals per 1979 Likanella campanensis; Canérot, p. 505. whorl (w value) (see also Radoičić, 1995). 1979 Likanella campanensis; Peybernès, p. 735. Actinoporella lucasi (Emberger, 1955) Granier, 1992, 1987 Falsolikanella campanensis; Granier, p. 208, pl. 1, Valanginian. Basionym: Clypeina lucasi Emberger, 1956.A figs. a–d, g, h, l, m, bibliography. synonym of A. podolica according to Conrad et al. (1974). 1988 Falsolikanella campanensis; Granier, p. 52, pl. 6, 7, Actinoporella maslovi Praturlon, 1964, Late Jurassic to figs. a–c, f, i. basal Neocomian. Transferred to Clypeina by Radoičić(1986). 1991 Falsolikanella campanensis; Farinacci and Radoičić, Actinoporella nigra (Conrad and Peybernès, 1978) Granier, p. 138, pl. 7, figs. 1, 2, 4, 5. 1994, late Hauterivian–late Barremian. Basionym: Clypeina 1993 Falsolikanella campanensis; Granier and Deloffre, nigra Conrad and Peybernés, 1978. The ‘excrescences’ of Gran- p. 30. ier (1994, pl. 3, fig. 1, arrow), next to the axis of the plant, denote 1993a Falsolikanella campanensis;Masse,p.315,pl.1,fig. 6. solid (closed) thickenings of the dark, microgranular calcareous 1993b Falsolikanella campanensis; Masse, p. 366.

Figure 9. Actinoporella campanensis (Azéma and Jaffrezo, 1972) n. comb. (1) Polyphysacean model in axial view with w = 22 and two rows of 11 laterals; (2) subtransversal section marked by almost horizontal line in (1). Dark gray circles (inner upper part) correspond to the upper corona of fertile laterals directed Figure 8. Actinoporella campanensis (Azéma and Jaffrezo, 1972) n. comb. (1) downward (dark colored); light gray ellipses (inner lower part) intersect the Polyphysacean model in axial view with w = 22 and two rows of 11 laterals; (2) lower corona of the fertile laterals inclined upward. Dark and light gray ellipses oblique section marked by oblique line in (1); note that transversally cut coronae (inner left and right sides) represent the basal parts of fertile laterals. (3) Same are limited to the lower half of the section; (3) tangential section marked by ver- specimen as Figure 3.1, coherent with (2). tical line in (1). (4) Same specimen as Figure 3.3, coherent with (2).

Holotype.—Oblique section with three whorls (University of Lyon collection, FSL 420001), thin section n. 3486, from Puig Campana (Alicante, Spain), Berriasian, (Azéma and Jaffrezo, 1972,pl.1,ﬁg. l), here shown in Figure 3.6.

Emended diagnosis.—Central stem cylindrical, bearing spaced whorls of heterocline laterals distally arranged in two rows, less frequently more than two rows. Each proximal pore corresponds to an initial short tract, corresponding to a primary lateral. Three vertically aligned elements arise from the primary lateral; the central element, arranged in distal continuity with the primary lateral, markedly flares outward, serving as fertile lateral (gametophore). Conversely, the upper and lower elements are rather short and slender, homologous to upper and lower coronae, as in Acetabularia. Calcification consists of a thin fibrous layer coating the primary laterals and the coronae. The layer extends outward, coating the gametophores for a large portion of their length. A thinner, continuous layer coats the central stem, leaving the space between whorls uncalcified.

Remarks.—Topotype specimens show the calcareous skeleton to be articulated, made of a thin, yellowish, fibrous layer coating the fertile laterals and the central stem, upward and downward, but excluding the bulk of the space between adjacent whorls. The thin, calcified layer follows the outline of the central stem and fertile laterals. Despite the primary fibrous mineralization, tangential sections sometimes suggest a lateral ‘fusion’ of the proximal pores (Fig. 4.6, 4.8), misleadingly calling for two gametangia to arise from a single proximal point. In well-preserved specimens, a darker rim, tentatively of primary origin, marks the outer side of the layer; it is particularly visible at the boundary between adjacent fertile laterals and the proximal contact between the corona and the fertile lateral (Figs. 3.6, 4.2, 4.6, 4.8). Instead, only a single layer is visible at the contact between the laterals, in the inner portion of the whorl (Fig. 3.1, 3.2). Toward the exterior, the calcareous layer distally decreases in thickness and is missing at the tip of the fertile lateral (Fig. 3.3). Along the central stem the calcareous layer also becomes thinner and thinner, coating the axis downward and upward from each whorl (Fig. 3.5), leaving the central stem mid part uncalcified, thus triggering the postmortem disarticula- Figure 10. (1) Actinoporella campanensis (Azéma and Jaffrezo, 1972) n. comb. (2) Actinoporella podolica (Alth, 1878). (1) Components of the fertile tion of the alga. lateral and calcification (fibrous calcite) colored black. The basal part corre- Fertile laterals are 16 to 25, commonly 20–24 per whorl. In sponds to the primary (first-order) lateral arising from the axis of the plant. (2) Notice the same structural elements in Actinoporella podolica. Compared with axial section, they arise from the tip of a short tract (e.g., Actinoporella campanensis, laterals appear slenderer with closer whorls; calcifi- Fig. 3.5) bearing three elements of different length. The longest cation (aragonite) is drawn in dark gray. and largest, phloiophorous and pear shaped, is set in prolonga- tion of the first tract (Fig. 3.5). The upper element (upper corona), cylindrical and rather elongated, arises perpendicularly 1993 Falsolikanella campanensis; Soták and Mišik, pl. 8, to the first tract and bends slightly forward, as some sections sug- figs. 3, 4. gest (Fig. 4.1, 4.9). In transversal section (Fig. 3.1) the upper 1994 Falsolikanella campanensis; Schindler and Conrad, coronae appear as elliptic pores between gametophores (white p. 75. arrow), but aside some equivalent pores tend to split in two cir- 1994 Falsolikanella campanensis; Granier and Berthou, cular smaller pores aligned radially (black arrow). Available p. 115, fig. 3. material does not allow better documentation and explanation Non Falsolikanella cf. F. campanensis; Bucur et al., of this character. Possibly, the couple of pores shown by the 2000 p. 440, pl. 5, figs. 11–13. black arrows correspond to corona protuberances visible on ?2014 Falsolikanella campanensis; Granier et al., p. 27, pl. extant polyphysacean algae (Dumais and Harrison, 2000, fig. 6, fig. d. 4h). The lower element (lower corona), shorter than the upper

Figure 11. Shape of gametophores in the extant genera Acetabularia and Parvocaulis.(1) Acetabularia farlowii Solms-Laubach, 1895;(2) Acetabularia peniculus (Brown, 1819) Solms-Laubach, 1895;(3) Parvocaulis exigua (Solms-Laubach, 1895) Berger et al., 2003;(4) Parvocaulis pusilla (Howe, 1909) Berger et al., 2003. Notice the absence of corona inferior in Parvocaulis and the slender to ‘pear-shaped’ morphology of the gametophores in the two genera. Drawings based on Berger and Kaever (1992) and Berger et al. (2003).

Figure 12. Actinoporella kukoci Radoičić, 1974. Type material. (1) Tangential section cutting a whorl proximally; the two rows of aligned rounded pores correspond to gametophores (arrow g) and one of two coronae (arrow c1); the pore pointed to by arrow c2 probably corresponds to the other corona; thin section n. RR2808; (2) longitudinal oblique section cutting a single whorl; the upper and lower protuberances set proximally aside each pore can be interpreted as calciﬁcation around recrystallized coronae; original illustration by Radoičić(1974, pl. 1, ﬁg. c); (3) tangential section of a whorl; one of two superimposed pores (arrow) probably corresponding to a corona; thin section n. RR2808. Scale bar = 0.50 mm.

one, is gently leaning and tapered forward. Depending on whether it belongs to fertile laterals, it bends upward or downward (Figs. 3.5, 4.5). The first tract, halfway in the section from the upper and lower coronae, forms the continuation of the short primary lateral, whereas its distal phloiophorous exten- sion corresponds to the gametophore (Fig. 10). In each whorl, pores corresponding to the fertile laterals are roughly aligned on a single horizontal plane. Proximally, however, at level of the corona, they diverge upward and downward; thus, in the proximal tangential section, adjacent tufts of three pores look slightly twisting (Fig. 4.6, 4.7). Distally, the fertile laterals are arranged in a single plane made of two rows (Fig. 4.7, 4.9), but noticeably, more than two tilts (probably three) occur in some specimens (Fig. 3.6). Transversal-oblique sections allow one to pursue the transi- tion among the elements forming the fertile laterals. In this kind of section, the coronae are cut transversally and appear as circles (Fig. 3.1–3.4). In Figure 8.2, the dark-gray circles correspond to the lower corona of downward-bending gametophores, being in continuity with them, whereas the light-gray ellipses match the lower corona of those gametophores directed upward. Note that in Figure 3.3 and 3.4 (last circular pore at right), the pores occur only in correspondence with the whorl from which they are originating. This means that such upright structures are short, not forming elongated laterals. Otherwise, laterals cut transversally would have all been visible, forming circles/ellipses (e.g., model shown in Fig. 7.2; see also De Castro, 1997, figs. 17, 18). Figure 8.2 explicates the elements cut in subtransversal section. In the middle part of the figure, the dark-gray circles correspond to the lower corona of downward-bending fertile laterals, while the two light-gray ellipses correspond to the lower corona of upward-bending fertile laterals. The two small, dark- gray elongated ellipses correspond to basal parts of downward- bending fertile laterals. In tangential section (e.g., Fig. 4.6, 4.8) the upper and lower coronae are ‘fused’ with the basal part forming an elliptic vertical pore (see also Fig. 8.3). Noticeably, in this kind of section, the threefold tuft (three vertically aligned pores) passes sideways to a couple of large pores (the gametophores, Fig. 4.1, 4.9)in agreement with wdist/wprox ratio equal to 1. No trace of reproductive organs (gametangia) was observed in the large pear-shaped gametophores or in the calcareous skeleton, making that the kind of reproduction that is supposedly homologous to that of Acetabularia. The reconstructions of the thallus (Figs. 13, 14) depict five whorls in longitudinal view/section and one in upper view. Only the top and the bottom of the thallus are not recorded, probably uncalcified. Tentatively, we assume that the shape of these uncalcified parts was like in the extant genus Halicoryne (Berger and Kaever, 1992). In fact, Actinoporella and Halicoryne belong to the same family, with a cylindrical thallus bearing spaced whorls of fertile laterals. Thus, in A. campanensis, the fertile laterals were probably decreasing gradually in size and inclination toward the apex. Moreover, because known specimens consist of whorls of fertile laterals, the calcified thallus Figure 13. Actinoporella campanensis (Azéma and Jaffrezo, 1972)n.comb. most likely corresponds to the mature stage of the alga while Reconstruction of the alga in axial view; (1) three consecutive whorls, removing fi the calcification; (1a) deprived of frontal fertile laterals; (1b) frontal fertile laterals the juvenile, sterile stages were probably not calci ed and truncated to show the coronae; (1c) whorl shown in perspective view; (2)softparts were lost during the algal growth. and calcareous skeleton of a whorl; (3) calcareous skeleton (black) of a whorl.

Systematics of species previously assigned to the genus Praturlonella danilovae (Radoičić, 1975) Falsolikanella Figure 15

Several species have been ascribed to the genus Falsolika- 1975 Likanella danilovae Radoičić, p. 151, pls. 1, 2. nella Granier, 1987; the status is here discussed for some of 1978 Praturlonella danilovae; Barattolo, p. 9, 31. them. 1999 Falsolikanella danilovae; Granier et al., p. 58, pl. 3, fig. 8, with synonymy. Pseudoactinoporella? silvaeregis (Bucur, 1981) Holotype.—Oblique section designed by Radoičić(1975), thin 1981 Pseudoactinoporella silvaeregis Bucur, p. 151, pls. 1, 2. section RR4243 of the Radoičićcollection (Fig. 15.7), 1992 Pseudoactinoporella? silvaeregis; Bucur, p. 450, pl. 3. corresponding to Radoičić(1969, pl. 1, fig. 1). 1999 Falsolikanella(?) silvaeregis; Bucur, p. 58, pl. 3, fig. 8, with synonymy. Remarks.—The late Hauterivian to early Barremian Likanella danilovae Radoičić, 1975 is transferred by Granier et al. (1999) to the genus Falsolikanella. Quoting Granier et al. (1999, Holotype.—Perpendicular–oblique section, sample 48E, thin p. 177), the “thallus bears single whorls consisting of ‘bundles’ section N.I. 1051, Carnet - Valea Poienii, Romania (Bucur, of branches, and that each bundle consists of a few secondary 1981, pl. 1, fig. 1). branches on top of a short vestibular part. Vestibules were first regarded as expansions of the main axis; they are presently Remarks.—Although with doubt, the Barremian to early Aptian considered as equivalents to primary branches.” The authors do Pseudoactinoporella silvaeregis Bucur, 1981 is assigned by not specify the number of secondary laterals per tuft. Bucur (1999) to the genus Falsolikanella. Quoting this author, Figure 16, based on the holotype (Fig. 15.7)ofP. danilo- the specimen in longitudinal section illustrated by Bucur vae, compares the metaspondyl model (Fig. 16.1a, 16.2a)to (1992, p. 451, pl. 3, fig. 7; reillustrated in Bucur 1999, pl. 3, the heterocline model (Fig. 16.1b, 16.2b) applicable to this spe- fig. 8) “exhibits a short globulous primary branch … splitting cies. In the metaspondyl model, the distal number of laterals in two phloiophorous branches arranged in a vertical plane.” (wdist) is estimated to be 80, while the number of secondary lat- The laterals start with a short peduncle, then those verging erals per tuft is supposed to be six. Also in the metaspondyl upward (e.g., second, third, fifth, and sixth whorls on the model, some 13 vestibules about 86 µ from each other are right) exhibit a sort of upward-verging ‘triangular’ expansion. expected to be present. In the heterocline model, 80 laterals Those slightly verging downward (second whorl on the left, are strictly and regularly aligned in a single whorl. Sections fourth and seventh on the right) show the similar ‘triangular’ of Likanella danilovae belonging to the type material are expansion to be set downward, resulting in all laterals shown in Figure 15. Although tangential sections (Fig. 15.5, widening regularly outward. The lack of proximal tangential 15.6) misleadingly show an arrangement in tufts (see Fig. 6 sections makes it difficult to understand whether the short and relative discussion), transversal and oblique sections proximal peduncle corresponds to a primary lateral. Therefore, (Fig. 15.1, 15.3, 15.4) display regularly crowded heterocline in this paper, the species is, provisionally at least, left in the laterals. The metaspondyl arrangement of P. danilovae is genus Pseudoactinoporella. therefore not proved, and in this paper, the assignment of the species to the genus Praturlonella, as proposed by Barattolo (1978), is accepted.

Praturlonella(?) teakolarae (Radoičić, Jurkovšek, and Jova- nović, 2011) new combination

2011 Clypeina teakolarae Radoičić, Jurkovšek, and Jovanović, p. 13, pl. 1–6. 2012 Falsolikanella? teakolarae; Sokačet al., p. 174, pl. 9–11, 18 (ﬁgs. 1–4).

Holotype.—Subaxial-tangential section from Alveolinid- Nummulitid Limestone of Kras Group, thin section BJ2796-27, Bogdan Jurkovšek collection housed in the Geological Survey of Slovenia, Ljubljana. Figured in Radoičić et al. (2011, pl. 1, ﬁg. 2).

Remarks.—Sokačet al. (2012) tentatively assigned the Eocene species Clypeina teakolarae Radoičić, Jurkovšek, and Jovanović, 2011 to Falsolikanella. Like in Clypeina jurassica, Figure 14. Actinoporella campanensis (Azéma and Jaffrezo, 1972) n. comb. Reconstruction of the alga in transversal view, removing calciﬁcation. the specimens assigned by Sokač et al. (2012)to

Figure 15. Praturlonella danilovae (Radoičić, 1975). (1–6) Type material; (7) Bosnia, Kljuc sheet; (1) transversal section, thin section n. RR1865; (2) oblique section, thin section n. RR1864; (3) subtransversal section, thin section n. RR1822; (4) oblique section, thin section n. RR1865; (5) tangential section, thin section n. RR1866; (6) tangential section, thin section n. RR1893; (7) longitudinal section, thin section n. RR4243. Scale bar = 0.5 mm.

Falsolikanella? teakolarae, including the species original Sokačet al., 2012, pl. 9, fig. 2, pl. 11, fig. 8). Better evidence material, exhibit a yellowish, fibrous calcareous wall. Whorls of metaspondyl arrangement could be shown in proximal cut longitudinally show diverging laterals forming tufts (e.g., tangential sections, but unfortunately, despite the abundant

laterals sometimes are proximally well calcified and arranged in two rows of alternating pores (Sokačet al., 2012, pl. 13, fig. 5, second whorl from the bottom), while sometimes they are less calcified, looking like pores fused together (e.g., Sokačet al., 2012, pl. 13, fig. 7). In our opinion, the metaspondyl versus the euspondyl heterocline arrangement of Falsolikanella? teakolarae is not demonstrated. Moreover, as shown by Sokačet al. (2012, pl. 11, figs. 5, 6), laterals distally widen abruptly, while conversely, the calcareous wall becomes very thin (e.g., Sokačet al., 2012, pl. 11, fig. 6, pl. 10, fig. 5, lowest pore). In fact, it is uncertain whether the laterals are distally closed, forming a cortex such as in Clypeina, or continue, forming a swollen vesicle such as in Praturlonella or Humiella. In this paper, the species is tentatively assigned to Praturlonella, introducing Praturlonella? teakolarae n. comb., noting however that among the so far nine species originally or later, either validly or not, assigned to the Tithonian–Eocene genus Praturlonella, none displays the aforementioned special pattern of calcification typical of Clypeina jurassica.

Humiella(?) macropora (Sokačet al., 2012) new combination

2012 Falsolikanella? macropora Sokačet al., p. 177, 180, pl. 12, figs. 1, 2, 3a, 4, 5, 6a, 7–9, pl. 13–17, 18, figs. 5–7. 2012 Praturlonella salernitana;Sokačet al., pl. 6–8, 12, fig. 3b.

Holotype.—Oblique section labeled RAK–40A/11 northern foothills of Mt. Biokovo, near the village of Kozica (Croatia), thin section RAK–40A, ﬁgured in Sokačet al. (2012, pl. 13, ﬁg. 5).

Remarks.—Sokač et al. (2012) tentatively assigned to Falsolikanella their new Eocene species Falsolikanella? macropora. As proposed by these authors (Sokačet al., 2012, p. 178), the occurrence of phloiophorous laterals differentiated into a stem and a clearly swollen outer part is consistent with a cladospore reproductive function. The fact that the inflated part is markedly elongated (Sokačet al., 2012, e.g., pl. 14, fig. 3) and distally calcified (Sokačet al., 2012, e.g., pl. 17, figs. 2, 5) apparently supports the hypothesis put forward by these authors, assuming the alga is metaspondyl, with six to eight tufts per whorl and each tuft made of five to seven laterals. However, transversal sections cut at the level of a whorl (Sokačet al., 2012, e.g., pl. 17, fig. 10) should display the arrangement in tufts, while conversely, a regular, radiated distribution all around the whorl is visible. Moreover, the size Figure 16. Organization of Likanella danilovae Radoičićin (1) longitudinal of the alga varies considerably, proportionally one to four and (2) transversal view. (1a, 2a) Sensu Granier et al. (1999); (1b, 2b) sensu Bar- (Sokačet al., 2012, compare pl. 17, figs. 5, 6). In the attolo (1978). Dasycladales, the number of pores per whorl proportionally increases with size. In F. macropora, small specimens show the pores cut initially to be roughly arranged on a single line and well-preserved material, none have been supplied. The ‘tuft and distally diverging in two rows (Sokačet al., 2012, e.g., pl. effect’ therefore may result from high values of w in the whorl, 17, fig. 6). Therefore, the alga must be considered euspondyl. combined with the heterocline arrangement. Calcification is As to the forms assigned by Sokačet al. (2012)toClypeina frequently poor or missing close to the insertion of the laterals lucana and Praturlonella salernitana, in our opinion they are on the central stem. In the author’snewFalsolikanella small specimens of a taxon to be assigned, at first glance, to macropora new combination (see the following), for example, Falsolikanella? macropora. In this paper, the heterocline

arrangement of the laterals is thought to be due to their high species, formally valid, A. podolica, A. kukoci, and A. campa- number per whorl, combined with their vesiculiferous shape, nensis. A discussion on the species previously assigned to the and therefore secondary in importance compared to the genus Falsolikanella is supplied, particularly on Falsolikanella euspondyl arrangement, specifically the reproductive function danilovae, whose transfer to the genus Praturlonella is of the laterals. In this paper, F. macropora is therefore confirmed. tentatively assigned to the genus Humiella Sokačand Velić, 1981. Acknowledgments Conclusions The authors thank E. Robert, UCBL-FSL (Collection de l’Uni- In 1972, Azéma and Jaffrezo described Likanella campanensis, versité Lyon 1), for the loan of the type specimens of Likanella a new species from the early Cretaceous (upper Berriasian or campanensis. We are grateful for the comments and suggestions basal Valanginian) of the eastern Betic Cordillera, southern received from S. Lo Duca and from reviewers B. Granier, J.-P Spain. The diagnosis and reconstruction referred to a cylindrical Masse, and F. Schlagintweit. A sincere thank you also goes to axis bearing twin whorls of all equal, pyriform (vesiculiferous), N. Scafetta, Dipartimento di Scienze della Terra dell’Ambiente and diverging laterals. A few years later, L. campanensis was e delle Risorse, Università di Napoli Federico II, and implicitly transferred by Sokačand Velić(1978b) to the genus L. Carbone, Dipartimento di Matematica, Università di Napoli Selliporella Sartoni and Crescenti, 1963 emend. Barattolo, De Federico II, for their useful suggestions and remarks concerning Castro, and Radoičić, 1988. Although not formally emended, the mathematical part (Supplemental Data). We thank H. Gollop Selliporella is newly described and assigned to the tribe Diplo- for the English review. poreae. Later, Granier (1987) supplied an alternative interpretation of L. campanensis, calling for the whorls to be simple, Accessibility of supplemental data made of clusters (tufts) of three, all similar first-order fertile laterals, lined up along the axis of the thallus and arising from a Data available from the Dryad Digital Repository: https://doi. proximal vestibule. Separately, the author noticed the special org/10.5061/dryad.664h2vf. pattern of calcification, comparable to that of Clypeina jurassica. Based on L. campanensis as type species, the genus Falso- likanella was introduced. Our observations largely agree with References those made by Granier (1987). We acknowledge the presence of three laterals (‘pores,’‘branches,’‘laterals,’‘ramifications,’ Alth, A. von, 1878, O Galicyjskich gatunkach skamienialych otwornic rodzaju Gyroporella Gümb: Rosprawy i Sprawozdania z Posiedzen Wydzialu and ‘rays’ are corresponding terms often used in the specialized Matematyczno-Przyrodniczego Akademii Umiejetrosciw Krakowie, v. 5, literature dealing with the fossil record) aligned axially, close to p. 71–112. Alth, A. von, 1881, WapieńNizniowski´ i jego skamieliny: Rozprawy i Spra- the central stem. The three laterals originate from a single, short wozdania z Posiedzen Wydzialu Matematyczno-Przyrodniczego, III Akade- proximal lateral, in this paper interpreted as the basal, first-order mii Umiejetnosciw Krakowie, v. 6, 160 p. origin of a tuft of second-order laterals, not a vestibule. As Alth, A. von, 1882, Die Versteinerungen des Nizniower´ Kalksteines: Beiträge zur Paläontologie Österreich-Ungarns und des Orients, v. 1, p. 183–332. shown by Granier (1987), the tufts are transversally very close Azéma, J., and Jaffrezo, M., 1972, Description de Likanella n. sp. algue dasy- to each other but not completely aligned in a single horizontal cladacée du Portlandien ou du Berriasien du Puig Campana (Province d’Ali- row, shifting upward and downward by lateral compression. cante, Espagne): Revista Española de Micropaleontologia, numero extraordinario XXX Aniversario E.N. Adaro, p. 125–129. Due to the vertical arrangement of the three second-order laterals Barattolo, F., 1978, Su di una nuova dasicladacea (alghe verdi) nel Paleocene forming a tuft, each of the close-set laterals counted in transver- dell’Appennino meridionale: Bollettino della Società dei Naturalisti in sal section at whorl level corresponds to a single tuft, thus enab- Napoli, v. 87, p. 83–157. fi Barattolo, F., 1983, Osservazioni su Triploporella steinmannii n. sp. (alghe ling estimation of the number of proximal, rst-order laterals per verdi, Dasicladali) del Cretacico del Messico: Bollettino della Società dei – whorl to a value of wprox = 16–25. Naturalisti in Napoli, v. 91, p. 89 123. The second-order, pear-shaped laterals are distally arranged Barattolo, F., 1998, Dasycladacean green algae and microproblematica of the uppermost Cretaceous–Paleocene in the Karst area (NE Italy and Slovenia): in two rows. Their heterocline arrangement makes the assess- Dela-Opera SAZU 4 razr., v. 34, p. 65–127. ment of the number of pear-shaped laterals more complicated Barattolo, F., and Carras, N., 1990, Pseudoclypeina distomensis n. sp. (green in thin section. It is however possible to guess the total, distal algae, Dasycladales) from the Malm of Greece: Bollettino della Società Paleontologica Italiana, v. 29, p. 145–162. number of laterals pertaining to a single whorl by doubling the Barattolo, F., De Castro, P., and Radoičić, R., 1988, Notes on the genus Sellipor- distal number of large distal laterals obtained in each row from ella Sartoni & Crescenti 1963 (Chlorophyta, Dasycladales): Società Geolo- gica Italiana - Atti del 74° Congresso Nazionale. Sorrento, Italy, Società transversal sections. In addition, a trigonometric method applied Geologica Italiana, p. A35–A38. to oblique sections is proposed (see Supplemental Data), result- Bassoullet, J.P., Bernier, P., Conrad, M.A., Deloffre, R., and Jaffrezo, M., 1978, Les algues Dasycladales du Jurassique et du Crétacé. Révision critique: ing in a value of wdist = 19–24. Therefore, the wdist/wprox ratio is Geobios, mémoire spécial 2, p. 1–330. one, showing that the presence of tufts of three second-order lat- Bassoullet, J.P., Bernier, P., Deloffre, R., Génot, P., Jaffrezo, M., and erals has no influence on the distal number of laterals. Such a Vachard, D., 1979, Essai de classification des Dasycladales en tribus: Bul- paradox is explained considering that two out of the three letin Centre de Recherches Exploration-Production Elf-Aquitaine, v. 3, ‘ ’ p. 429–442. pores primally found correspond to short, blind structures, cor- Berger, S., and Kaever, M., 1992, Dasycladales. An Illustrated Monograph of a onae, likewise found in the extant genus Acetabularia. From this Fascinating Algal Order: Stuttgart, Georg Thieme Verlag, 247 p. clarification, F. campanensis is transferred from family Diplo- Berger, S., Fettweiss, U., Gleissberg, S., Liddle, L., Richter, U., Sawitzky, H., and Zuccarello, G., 2003, 18S rDNA phylogeny and evolution of cap devel- poraceae to family Polyphysaceae. The genus Actinoporella, opment in Polyphysaceae (formerly Acetabulariaceae; Dasycladales, whose systematics is emended, currently comprises three Chlorophyta): Phycologia, v. 42, p. 506–561.

Brown, R., 1819, Fucus Peniculus, in Turner, D., Fuci or Coloured Figures and Granier, B., 1994, The genus Actinoporella (Gümbel in Alth, 1881) and its Descriptions of the Plants Referred by Botanists to the Genus Fucus, Vol- representatives. A review: Beiträge zur Paläontologie, v. 19, p. 113–127. ume 4: London, McCreery, 153 p. Granier, B., and Berthou, P.Y., 1994, Description de Milanovicella momciliana Bucur, I.I., 1981, Algues calcaires du Crétacé inférieur des Monts Padurea n. gen. n. sp., algue Dasycladale du Portlandien de l’Algarve central (Portu- Craiului: Nymphaea - Folia naturae Bihariae, v. 8–9, p. 53–68. gal), et validation de quelques taxons affins: Revue de Micropaléontologie, Bucur, I.I., 1992, Revised description of some dasyclad species from the Roma- v. 37, p. 113–121. nian Lower Cretaceous: Revue de Paléobiologie, v. 11, p. 447–461. Granier, B., and Brun, R., 1991, Cylindoporella cruciformis et Holosporella Bucur, I.I., 1999, Lower Cretaceous dasyclad algae from the Padurea Craiului arabica, deux Dasycladacées nouvelles du Groupe Thamama, (? Massif (Northern Apuseni Mountains, Romania): Acta Palaeontologica Portlandien-) Berriasien-Aptien d’Abu Dhabi, Emirats Arabes Unis: Cret- Romaniae, v. 2, p. 53–72. aceous Research, v. 12, p. 403–410. Bucur, I.I., Koch, R., Kirmaci, Z.M., and Tatli, K., 2000, Les algues Dasycla- Granier, B., and Deloffre, R., 1993, Inventaire critique des algues Dasycladales dales du Crétacé inférieur (Calcaire de Berdiga) de Kircaova (région de fossiles - II partie - Les algues Dasycladales du Jurassique et du Crétacé: Kale-Gümüshane, NE Turquie): Revue de Paléobiologie, v. 19, p. 435–463. Revue de Paléobiologie, v. 12, p. 19–65. Bucur, I.I., Rashidi, K., and Senowbari-Daryan, B., 2012, Early Cretaceous cal- Granier, B., Bucur, I.I., and Trabold, G., 1999, Falsolikanella danilovae careous algae from Central Iran (Taft formation, south of Aliabad, near Radoicic ex Barattolo 1978, n. comb., a diploporacean alga from the Urgo- Yazd): Facies, v. 58, p. 605–636. nian facies: Acta Palaeontologica Romaniae, v. 2, p. 177–181. Canérot, J., 1979, Les algues et leur environment dans le Malm et le Crétacé Granier, B., Clavel, B., Charollais, J., and Weidmann, M., 2014, Latest Jurassic– inférieur des chaînes ibériques et catalanes (Espagne): Bulletin du Centre Early Cretaceous Dasycladalean Algae (Chlorophyta) from the Morand dril- Recherches Exploration-Production Elf Aquitaine, v. 3, p. 505–518. ling at Montricher (Canton of Vaud, Switzerland): Acta Palaeontologica Conrad, M.A., 1970, Barremian and lower Aptian Dasycladaceae in the area Romaniae, v. 10, p. 25–38. surrounding Geneva (Switzerland): Geologica Romana, v. 9, p. 63–100. Gümbel, C.W., 1882, Genus Actinoporella, in Alth A.v., Die Versteinerungen Conrad, M.A., 1977, The Lower Cretaceous calcareous algae in the area sur- des Nizniower´ Kalksteines: Beiträge zur Paläontologie Österreich-Ungarns rounding Geneva (Switzerland): Biostratigraphy and depositional environ- und des Orients, I, p. 321–322. ments, in Flügel, E., ed., Fossil Algae: Recent Results and Developments: Harvey, W.H., 1859, Characters of new algae, chiefly from Japan and adjacent Berlin, Springer-Verlag, p. 295–300. regions, collected by Charles Wright in the North Pacific Exploring Exped- Conrad, M.A., and Peybernés, B., 1976, Hauterivian Albian Dasycladaceae ition under Captain John Rodgers: Proceedings of the American Academy from the Urgonian limestones in the French and Spanish eastern Pyrenees: of Arts and Sciences, v. 4, p. 327–335. Geologica Romana, v. 15, p. 175–197. Howe, M.A., 1909, Phycological studies-IV. The genus Neomeris and notes on other Conrad, M.A., and Peybernès, B., 1978, Sur quelques Dasycladales de Siphonales: Bulletin of the Torrey Botanical Club, v. 36, p. 75–104. l’Urgo-Aptien du Prébalkan bulgare: Compte Rendu des Séances de la Société Jaffrezo, M., 1974, Les algues calcaires du Jurassique supérieur et du Crétacé de Physique et d’Histoire naturelle de Genève, nouvelle série, v. 12, p. 69–83. inférieur des Corbières (2ème partie): Revue de Micropaléontologie, Conrad, M.A., Praturlon, A., and Radoičić, R., 1974, The Genus Actinoporella v. 17, p. 23–32. Guembel in Alth 1882, Dasycladales, green algae. A revision: Geologica Kützing, F.T., 1843, Phycologia Generalis oder Anatomie, Physiologie und Romana, v. 13, p. 1–15. Systemkunde der Tange: Leipzig, F.A. Brockhaus, 458 p. Decaisne, J., 1842, Essais sur une classification des algues et des polypiers cal- Kützing, F.T., 1863, Diagnosen und Bemerkungen zu drei und siebenzig neun cifères de Lamouroux: Annales des Sciences Naturelles, Botanique, seconde Algenspecies. Zu der öffentlichen Prüfung sämmtlicher Klassen der série, v. 17, p. 297–380. Realschule zu Nordhausen: Nordhausen, Realschule zu Nordhausen, p. 1–19. De Castro, P., 1997, Introduzione allo studio in sezione sottile delle dasicladali Lamouroux, J.V., 1816, Histoire des Polypiers Coralligènes Flexibles, Vulgaire- fossili: Quaderni dell’Accademia Pontaniana, v. 22, 261 p. ment Nommés Zoophytes: Caën, F. Poisson, 635 p. Deloffre, R., 1988, Nouvelle taxonomie des Algues Dasycladales [New tax- Linnaeus, C., 1758, Systema Naturae (tenth edition), Volume 1: Holmiae, onomy of Dasycladale Algae]: Bulletin du Centre Recherches Exploration- Impensis Direct. Laurentii Salvii, 824 p. Production Elf Aquitaine, v. 12, p. 165–217. Lorenz, T., 1902, Geologische Studien im Grenzgebiete zwischen helvetischer Dumais, J., and Harrison, L.G., 2000, Whorl morphogenesis in the dasycladalean und ostalpiner Facies: Berichte der Naturforschenden Gesellschaft zu Frei- algae: The pattern formation viewpoint: Philosophical Transactions of the burg im Breisgau, v. 12, p. 34–62. Royal Society of London, Biological Sciences, series B, v. 355, p. 281–305. Maslov, V.P., 1958, New findings of Dasycladaceae in Jurassic of Krym: Dok- Elliott, G.F., 1968, Permian to Paleocene calcareous algae (Dasycladaceae) of lady Akademii Nauk SSSR, v. 121, p. 354–357. the Middle East: Bulletin of the British Museum of Natural History (Geol- Masse, J.P., 1993a, Valanginian–Early Aptian Carbonate Platforms from Pro- ogy), Supplement 4, 111 p. vence, Southeastern France, in Toni Simo, J.A., Scott, R.W., and Masse, Elliott, G.F., 1977. A consideration of the tribe Thyrsoporelleae, dasyclad algae: J.P., Cretaceous Carbonate Platforms: American Association of Petroleum Palaeontology, v. 20, p. 705–714. Geologists, v. 56, p. 363–374. Emberger, J., 1955, Les Clypeines (Algues siphonees verticillees) des Monts Masse, J.P., 1993b, Early Cretaceous Dasycladales biostratigraphy from Pro- Oulad Nail (Atlas saharien, Algerie): Bulletin de la Société Géologique vence and adjacent regions (South of France, Switzerland, Spain). A refer- de France, v. 5, p. 543–552. ence for Mesogean correlations, in Barattolo, F., De Castro, P., and Emberger, J., 1956, Les Clypéines (Algues siphonées verticillées) des Monts Parente, M., eds., Studies on Fossil Benthic Algae: Bollettino della Società des Oulad-Naïl, Atlas saharien, Algérie : Bulletin de la Société Géologique Paleontologica Italiana, spec. vol. 1, p. 311–324. de France, v. 5, p. 543–552. Michelin, H., 1840, Iconographie zoophytologique. Description par localités Farinacci, A., and Radoičić, R., 1991, Late Jurassic–Early Cretaceous Dasycla- et terrains des Polypiers fossiles de France et pays environnants: Paris, dales (green algae) from Western Pontides, Turkey, in Farinacci, A., P. Bertrand, 348 p. Ager, D.V., and Nicosia, U., eds., Geology and Paleontology of Western Milanovic, M., 1966, Likanella, a new Permian genus of the family Dasyclada- Pontides, Turkey: Geologica Romana, v. 27, p. 135–165. ceae: Geoloski vjesnik, v. 19, p. 9–14. Favre, J., and Richard, A., 1927, Etude du Jurassique supérieur de Pierre-Châtel Pascher, A., 1931, Systematische Übersicht über die mit Flagellaten in Zusam- et de la Cluse de la Balme: Mémoires de la Société paléontologique suisse, menhang stehenden Algenreihen und Versuch einer Einreihung dieser v. 46, p. 1–38. Algenstämme in die Stämme des Pflanzenreiches: Beihefte zum Bota- Fourcade, E., Jerez, L., Rodriguez, T., and Jaffrezo, M., 1972, El Jurasico ter- nischen Centralblatt, v. 48, p. 317–332. minal y el Cretacico inferior de la Sierra de la Muela (Provincia de Murcia). Peybernès, B., 1979, Les Algues du Jurassique et du Crétacé inférieur des Consideraciones sobre las biozonas con foraminiferos del Albense-Aptense Pyrénées Franco-Espagnoles. Intérêt biostratigraphique et paléoécologique, del Sureste de Espana: Revista Española de Micropaleontologia, num. extra- in 2ème Symposium International sur les Algues Fossiles. Paris, 23–26 avril ord., XXX Aniversario E.N. Adaro, p. 215–248. 1979: Bulletin du Centre de Recherches Exploration-Production Génot, P., Saint-Martin, J.-P., and Bucur, I.I., 2008, Polyphysaceae fertile caps Elf-Aquitaine, v. 3, p. 733–741. in Hungarian Sarmatian sediments: Geologia Croatica, v. 61, p. 341–344. Pia, J., 1917, Familie: Dasycladaceae (Endl.) Cram. em., in Trauth, F., ed., Granier, B., 1987, Révision de Likanella campanensis Azema et Jaffrezo, 1972 Das Eozänvorkommen bei Radstat im Pongau und seine Beziehungen algue dasycladacée du Crétacé inférieur du sud-est de l’Espagne: Revue de zu den gleichalterigen Ablagerungen bei Kirchberg am Wechsel und Paléobiologie, v. 6, p. 207–212. Wimpassing am Leithagebirge: Denkschriften der mathematisch- Granier, B., 1988, Algues chlorophyceae du Jurassique terminal et du Crétacé naturwissenschaftlichen Klasse, v. 95, p. 209–213. inférieur en Alicante: Mediterránea: Serie de Estudios Geológicos, v. 5, Pia, J., 1920, Die Siphoneae verticillatae vom Karbon bis zur Kreide. Abhan- p. 5–96. dlungen der Zoologisch-Botanischen Gesellschaft in Wien, v. 11, p. 3–263. Granier, B., 1992, Les Algues et Foraminifères benthiques du Jurassique supér- Praturlon, A., 1964, Calcareous algae from Jurassic–Cretaceous limestones of ieur et du Crétacé inférieur du Sénégal: Journal of African Earth Sciences, Central Apennines (Southern Latium - Abruzzi): Geologica Romana, v. 3, v. 14, p. 239–253. p. 171–202.

Radoičić, R., 1969, Likanella? danilovae n. sp. and some other Lower Cret- chronostratigraphic and cyclostratigraphic units: Revue de Paléobiologie, aceous Dasycladaceae from the outer Dinarides: Vestnik Zavod za Geološka v. 13, p. 59–96. i Geofizička Istraživanja, (A) 26, p. 176–194, 237–275. Silva, P.C., 1952, A review of nomenclatural conservation in the algae from the Radoičić, R., 1974, A new Palaeocene Actinoporella (Dasycladaceae) (A pre- point of view of the type method: University of California Publications in liminary report): Bulletin Scientifique du Conseille de l’Academie des Botany, v. 25, p. 241–323. Sciences RSF de Yougoslavie, Section A Sciences Naturelles, Techniques Simmons, M.D., Emery, D., and Pickard, N.A.H., 1991, Hensonella dinarica, et Médicales v. 19, p. 230–231. an originally calcitic Early Cretaceous dasycladacean alga: Palaeontology Radoičić, R., 1975, Likanella hammudai sp. nov. from the Upper Cretaceous of (London), v. 34, p. 955–961. Tripoli area (Libya) and the age of strata containing Dissocladella undulata Sokač, B., and Velić, I., 1978a, Biostratigraphic investigations of the Lower Raineri: Annales Géologiques de la Péninsule Balkanique, v. 39, p. 147–152. Cretaceous of the Outer Dinarides I. The Neocomian of western Istria: Radoičić, R., 1978, O rasprostranjenju dazikladacea u donjoj kredi Kurilova, Geoloski vjesnik, v. 30, p. 243–250. Istočna Srbija [On distribution of dasycladacea in the Lower Cretaceous Sokač, B., and Velić, I., 1978b, Redescription of the genus Selliporella (Calcar- of Kurilovo, Eastern Serbia]: Bulletin du Museum d’Histoire Naturelle de eous algae; Dasycladaceae): Geoloski vjesnik, v. 30, p. 225–242. Belgrade, v. 33, série A, p. 183–197. Sokač, B., and Velić, I., 1981, Humiella teutae n. gen. n. sp. (Dasycladaceae) Radoičić, R., 1986, O jednoj novoj klipeini i nekim drugim dazikladaceama iz from the Neocomian of Southern Herzegovina: Geoloski Vjesnik, v. 33, Jure i Krede Crne Gore [On a new Clypeina and some other dasyclads from p. 101–105. the Jurassic and the Cretaceous of Montenegro]: Geoloski Glasnik - Bulletin Sokač, B., Velić, I., Grgasović, T., Ćosović, V., and Vlahović, I., 2012, Tax- Géologique, v. 10, p. 53–65. onomy and stratigraphy of an algal assemblage in Palaeogene deposits of Radoičić, R., 1995, On the age of limestones with Actinoporella kukoci the northern foothills of Mt. Biokovo (Southern Croatia): Geologia Croatica, Radoičić, 1975 and Praturlonella salernitana Barattolo, 1978 (Dasyclada- v. 65, p. 161–205. cean algae): Revue de Paléobiologie, v. 14, p. 95–105. Solms-Laubach, H. Graf Zu, 1895, Monograph of the Acetabularieae: Radoičić, R., Jurkovšek, B., and Jovanović, D., 2011, Clypeina teakolarae sp. Transactions of the Linnean Society of London, series 2, Botany, v. 5, nov., a dasycladalean alga from the lower Eocene of Rakitovec (Čičarija, p. 1–39. Slovenia): Geologija, v. 54, p. 13–30. Soták, J., and Mišik, M., 1993, Jurassic and Lower Cretaceous dasycladacean Raviv, V., and Lorch, J., 1970, Verticilloporella, a new Mesozoic genus algae from the Western Carpathians, in Barattolo, F., De Castro, P., and of Dasycladaceae, with discussion on Munieria and Actinoporella: Israel Parente, M., eds., Studies on Fossil Benthic Algae: Bollettino della Società Journal of Botany, v. 19, p. 225–235. Paleontologica Italiana, special volume 1, p. 383–404. Raviv, V., and Lorch, J., 1992, Mesozoic Dasycladaceae (Chlorophyceae) from Valet, G., 1968, Contribution à l’étude des Dasycladales. 1. Morphogenèse: Israel: Israel Journal of Botany, v. 41, p. 1–54. Nova Hedwigia, v. 16, p. 21–82. Sartoni, S., and Crescenti, U., 1963, Ricerche biostratigrafiche nel Mesozoico Valet, G., 1969, Contribution à l’étude des Dasycladales. 2 - Cytologie dell’Appennino meridionale: Giornale di Geologia, v. 29, p. 153–302. et reproduction, 3 - Révision systématique: Nova Hedwigia, v. 17, Schafhäutl, K.E., 1853, Beiträge zur näheren Kenntnis der Bayern’schen p. 551–644. Voralpen: Neues Jahrbuch für Mineralogie, Geologie und Paläontologie, Yamada, Y., 1934, The marine Chlorophyceae from Ryukyu, especially from v. 1853, p. 299–319. the vicinity of Nawa: Journal of the Faculty of Sciences, Hokkaido Imperial Schafhäutl, K.E., 1863, Süd-Bayerns Lethaea Geognostica. Der Kressenberg University, series 5, Botany, v. 3, p. 33–88. und die südlich von ihm gelegenen Hochalpen geognostisch betrachtet in ihren Petrefacten: Leipzig, Voss, 504 p. Schindler, U., and Conrad, M.A., 1994, The Lower Cretaceous Dasycladales from the Northwestern Friuli Platform and their distribution in Accepted: 24 December 2018