 Geologia Croatica 68/3 173–178 1 Fig. 2 Pls. Zagreb 2015 Montenegrella? gracilis n. sp., a new calcareous alga (Dasycladales) from the Upper Barremian of Mt. Biokovo (Dinarides Mts., Croatia)  Branko Sokač and Tonći Grgasović Croatian Geological Survey, Sachsova 2, Zagreb, Croatia; (tgrgasovic@hgi-cgs.hr) doi: 10.4154/gc.2015.14 Geologia CroaticaGeologia Croatica ABSTRACT Montegrella? gracilis n.sp. (Dasycladales) is characterized by a thick calcareous envelope and narrow central cavity. Branches display two orders of laterals with alternate arrangement between consecutive whorls. The primaries are vis- ibly differentiated into a thin, tendril-like stalk in the proximal part and a club-shaped or ellipsoidal inflated swelling in the distal part. The secondaries are poorly visible, of unclear shape and number, with a supposedly common start- ing point. In the type-locality, the alga occurs within a rich, typically Upper Barremian, algal assemblage. The valid- ity of the genus Montenegrella, being disputed by BARATTOLO (1983), is commented upon by showing the generic attribution of the same or different species either to Suppiluliumaella or to Montenegrella by different authors. Keywords: calcareous algae (Dasycladales), taxonomy, Barremian, Dinarides Mts., Croatia 1. INTRODUCTION While re-reviewing thin-sections made from sample KJ-18 from Barremian deposits of Mt. Biokovo we noticed sections of some dasycladalean algae that were not mentioned in pre- viously published papers. The description and illustrations of these previously identified algae, accompanied by short descriptions of lithology, lithofacies changes, and the strati- graphic position of the sample KJ-18 in the local geological column, were given in SOKAČ (2004, 2005, 2007) and SOKAČ & GRGASOVIĆ (2004, partly 2008). In addition, the outcrop in question was visited and discussed by the par- ticipants of the 9th International Congress on Fossil Algae, held in 2007 in Croatia. Data given in the afore-mentioned publications will not be repeated here. The new species has been questioningly ascribed to Montenegrella, in spite of the validity of that genus being disputed. BARATTOLO (1983), in the description of his new species Suppiluliumaella schroederi, compared morpholog- ical characteristics of the species ascribed hitherto to Sup- piluliumaella and Montenegrella and concluded, that mor- phological details mentioned in the generic description of Montenegrella SOKAČ & NIKLER (1973) were not suffi- cient for the erection of the new genus, which, should, there- fore, be considered a younger synonym of Suppiluliumaella. Consequently, BARATTOLO (1983) suggested the transfer of all species described as Montenegrella by SOKAČ & NIKLER (1973), except perhaps M. fusiformis, as well as those described by DRAGASTAN (1978; M. transdanubica and M. alimani), to Suppilulliumaella. He accepts the pos- sible identity of Montenegrella verae and M. tubifera, ac- cording to BASSOULLET et al. (1978). BERNIER (1978) thinks that M. florifera is inadequately known to be ascribed to Suppiluliumaella, whereas M. corbarica JAFFREZO (1975) could be ascribed to some other genus. Later authors also espoused different views. Some authors follow (or partly follow) BARATTOLO’s (1983) opinion, whereas other con- tinue to use Montenegrella in their identifications. Thus DRAGASTAN (1989) transfers his species M. transdanu- bica and M. alimani (DRAGASTAN, 1978) without any comment to Suppiluliumaella, adding simultaneously two new species (S.leini and S. riedeli). These, however, are rep- resented by insufficiently clear illustrations, which make their generic attribution questionable. Similarly, there is no correct explanation for their inclusion in the synonymy of Salpingoporella pygmaea, as has been done by CARRAS et al. (2006). GRANIER & DELOFFRE (1993), in their inven- tory of Jurassic and Cretaceous fossil Dasycladales, omit any critical analysis of the relationship between Suppiluliumaella ELLIOTT 1968, and Montenegrella SOKAČ & NIKLER, Geologia Croatica 68/3Geologia Croatica 174 1978, the latter being invalidly described (no holotype was designated for its type-species). In this paper lectotypes are selected by the authors, thus giving Montenegrella and the species described on this occasion a valid status. In contrast to BARATTOLO (1983), CARRAS (1995) accepted the originally described Montenegrella corbarica JAFREZZO (1975) as a valid species, but transferred it to Suppiluliu- maella, with one illustration (pl. 45, fig. 2) labelled S. cor- barica (JAFFREZO). BUCUR (2001), in the chapter on the Lower Cretaceous, (p. 68), mentions Suppiluliumaella (Mon- tenegrella) tubifera and in the same publication, on p. 142 (Stop 6), mentions and illustrates (pl. 8, figs 7-8; pl. 9) Mon- tenegrella tubifera SOKAČ & NIKLER as the dominant spe- cies in the algal association. BUCUR & SĂSĂRĂN (2005), in their description of the Upper Jurassic and Lower Creta- ceous deposits of the Trascău Mountains, mention and illus- trate (p. 28, pl. 1, fig. 11), among others, Montenegrella cf. florifera BERNIER; that is, they accept both the genus Mon- tenegrella and the species M. florifera. In MICHETIUC, CATINCUT & BUCUR (2008), the type-species of Mon- tenegrella (M. tubifera) is mentioned and illustrated (p.220, pl. VI, fig. 2) as Suppiluliumaella tubifera. SCHLAGINT- WEIT et al. (2006) give a picture (fig. 3/16) of an alga that is neither mentioned in the list of fossils nor in the list of sections pictured in fig. 3. However, an identical section is figured in GAWLICK et al. (2008, fig. 5/7) under the origi- nal name Montenegrella tubifera SOKAČ & NIKLER. Comments above show that there reigns a bewildering confusion regarding the attribution of individual species to one genus or another of Suppiluliumaella and Montenegrella. The usage of the two generic denominations by individual or even the same authors on different occasions, for the same or different species originally attributed to Montenegrella, shows the author’s dilemmas, resulting in inconsistent generic attri- butions. This question can be viewed (and explained) from different perspectives, but the opinion here is that the main reason seems to be the lack of clearly defined criteria, sum- ming up variously validated biologic parameters, in a suffi- cient and unambiguously applicable manner in order to estab- lish a new taxon (genus or species). Such criteria should be accompanied by photomicrographs of three, four or more clear and unequivocal characteristic sections. Unfortunately, the es- tablishing of new taxa is frequently based upon one or two, (seldom more), poorly preserved, often heavily recrystallized sections that do not permit the insight into real and robust mor- phological characteristics. The establishment of new taxa based on such inadequate sections, adjusted to the author’s goals and personal opinion, makes the validity of a newly de- scribed taxon questionable, opening, at the same time, possi- bilities for further new combinations (and speculation). A de- scription need not restrain the author’s freedom of expression: the authors are free to write up, line up and follow their own ideas, but the facts should be the same and equally visible for everybody, without personal, supposed conclusions, being ei- ther described or shown graphically, and aimed at supporting the author’s interpretation. The consequences of such inade- quate and variable ranking of individual morphological pa- rameters (e.g., can a bigger or smaller D and d in relation to an original description be a decisive criterion for separating species and, if so, which will be the limiting values?), may (unjustifiably) encourage various authors to establish and/or transfer taxa according to somebody’s personal opinion, with- out providing new data. Following the aforementioned lack of critical analysis, and accepting only the last published opin- ion (which is often the case), opens a number of questions re- garding the validity of individual species and their generic at- tribution. Examples abound, but we don’t want to single out any particular example (each having an author) or to exempt ourselves. We are not convinced that these questions can be objectively treated by partial corrections on a case-by-case basis, with uncoordinated criteria,; instead, they can only deepen the existing confusion. The difference between the genera Suppiluliumaella and Montenegrella is clearly visible in the growth of secondary branches. In Suppiluliumaella secondaries grow out of the en- tire surface of the distal end of the primary branch in the form of finger-like protrusions, but in Montenegrella secondaries grow out as a bunch or a bundle of branches (see comparison drawings in SOKAČ & NIKLER, 1973, tab. I). This is com- pletely clear when the original material of the type species of both genera are compared (ELLIOTT, 1968, pl. 95, fig. 1-3; SOKAČ & NIKLER, 1973, pl. 2, fig. 1-4, pl. 3, fig. 1, 3). Therefore we retain the genus Montenegrella to which we assign the new species, Montenegrella? gracilis n. sp. However, because of the unclear shape of the secondaries and their relationships with the distal ends of the primaries (a common point of departure is supposed), its generic attri- bution remains uncertain. 2. PALAEONTOLOGICAL DESCRIPTION Genus Montenegrella SOKAČ & NIKLER in GRANIER & DELOFFRE, 1993, non SOKAČ & NIKLER, 1973 Montenegrella? gracilis n. sp. (Pls. I-II) Origin of the name: The species has been named after the thin, tendril-like stalks of the primary branches, giving the alga the appearance of a tender structure. Type locality: Mt. Biokovo (South Croatia), in a road cut situated approximately in the middle between the Vošac mountain hut and the Sv. Jure peak (1762 m, Fig. 1). Coor- dinates: 43o 19’ 36’’ N, 17o 03’ 44’’ E. Type stratum: The sample KJ-18 with the new alga be- longs to the level of light brown, well-bedded limestone, with bed thickness ranging from 0.3 to 1.2 m. The micro- facies types are fenestral, skeletal- and oncoid-bearing, in- traclastic grainstone and skeletal–peloidal packstone. The allochems include irregular intraclasts, small pellets, micri- tized skeletons of benthic foraminifera and gastropods, and, less commonly, oncoids, skeletons and bioclasts of dasycla- dalean algae, and centripetally micritized shell bioclasts. The lithological succession is represented by an alternation of mudstone, algal and foraminiferal wackestone, and sporadic occurrences of skeletal–intraclastic grainstone. The stratigraphic position of sample KJ-18 is the Upper Barremian, determined by the rich dasycladalean associa- Sokač & Grgasović: Montenegrella? gracilis n. sp., a new calcareous alga (Dasycladales) from the Upper Barremian of Mt. Biokovo (Dinarides Mts., Croatia) Geologia Croatica 175 tion, and with the first occurrence of Palorbitolina lenticu- laris in the strata immediately above the investigated sample. The first occurrence of P. lenticularis, before the occurrence of the Early Aptian praeorbitolinids, indicates a Late Bar- remian age in the Karst Dinarides (VELIĆ, 2007), although the lowermost occurrence of this foraminifera has been proven to be Early Barremian (GRANIER et al., 2013). For a detailed description of the fossil assemblage, ge- ographic and stratigraphic position see SOKAČ (2004, 2005, 2007), SOKAČ & GRGA SO VIĆ (2004, 2008). Holotype: Transverse-oblique section in slide KJ- 18/126, pictured in Pl. I, fig. 4. Isotypes are re presented by variously oriented sections figured in Pls. I-II. The original material will be stored in the Collection of holotypes at the Croatian Natural History Museum in Zagreb, Demetrova 1. Diagnosis: Cylindrical, unbranched thallus with a rela- tively narrow central cavity (small value of d) with regard to the outer diameter (D). Bipartite branches alternate in densely arranged whorls. Primaries are characterized by a thin tendril-like stalk, which, in the outer half of their total length, visibly thickens, becoming club-shaped or ellipsoi- dal. The secondaries are of unclear shape, appearing to be very short, their existence being indicated by a funnel-shaped widening as a con tinuation of the primaries’ distal ends. The shape and the dimensions of the funnel-shaped widening suggest their common starting point. Dimensions in mm Maximum observed thallus length L 6,80 Outer thallus diameter D 1,76-3,90 Innet thallus diameter d 0,40-1,20 Relation inner/outer diameter d/D 0,150 – 0,330 Distance between neighbouring whorls h 0,16-0,24 Proximal diameter of primaries p 0,04-0,09 Distal diameter of primaries p’ 0,22-0,34 Length of primaries l 0,92-1,56 Number of primaries per whorl w 26-30 Inclination of primaries α 30o-40o Length of secondaries l’ ? 0,16-0,20 Number of secondaries on a primary branch w’ ? 3-4 Description: The new species is characterized by an un- branched, cylindrical thallus, consisting of fine- to medium- grained mosaic calcite which replaced the original, probably aragonite skeleton. The clearly delimited, narrow central cav- ity occupies from about 15% up to one third of the total diam- eter at the most. The outer surface of the thick calcite enve- lope, perforated by pores and small canals of the branches, depending on the position of the plane of section, appears to be rather uniformly abraded, thus being relatively regular and usually delimited by a thin, dark micritic rim. The inner sur- face is smooth and also delimited by a thin micritic rim. Bipartite branches are arranged in regular whorls; in the neighbouring whorls, the primaries occupy alternating posi- tions (Pl. II, figs. 1, 4). Phloiophorous primaries are charac- terized by a thin, tendril-like stalk with a diameter of 0.04 to 0.09 mm in their proximal half; they gradually widen into a swollen distal half (p’ 0.22-0.34 mm), thus assuming, in gen- eral, a relatively regular club-like shape (Pl. I, figs. 1, 4, 8). In oblique and oblique-tangential sections, the swollen distal parts appear as more or less rounded or ellipsoidal pores (Pl. I, figs. 1, 6; Pl. II, figs. 1, 4). The variable shape of the pores, some of which suggest the possibility of being vertically compressed, is probably only the consequence of relation- ships between their being vertically inclined and the position of the plane of section. The branches grow obliquely and are directed upwards. The direction of their growth makes an angle with the horizontal plane section of (α) 30°-40°. Whorls are comparatively densely arranged, so that the dis- tal parts of individual branches touch mutually, both within the same whorl and with the branches of the neighbouring whorls (Pl. I, fig. 2; Pl. II, fig. 4). The existence of secondaries seem without question; however, due to corrosion of the outer surface they are mostly erased, only seldom being preserved, and poorly vis- ible in only some parts of some sections (Pl. I, fig. 4). De- spite not knowing how much of the outer surface of the skel- eton has been abraded during transport, the secondaries seem to have been very short indeed (length - l’ 0.16-0.20 mm). The existence of secondaries, their dense arrangement, and probably their common starting point is also indicated by partially visible, shallow, funnel-shaped widenings occur- ring at the distal ends of the primaries (Pl. I, figs. 4, 8; Pl. II, fig. 8). However, their number and shape remain unknown. The shape and width of the funnel-shaped widening suggests a phloiophorous shape for the secondaries and 3?-4 in number per bundle. These uncertainties in the position and growth form of the secondaries make the generic attribution debatable. Similarities and differences: With regard to the origi- nally described species of Montenegrella, the new species is distinguished by the regularity of the primaries, pronounced thin, tendril-like stalks in the proximal half and regular, club- shaped or ellipsoidal swelling in the distal (outer) half of the calcareous envelope. In addition, it is distinguished by a thick calcite envelope and thin, regularly shaped central cavity. With regard to Montenegrella florifera BERNIER and M. corbarica JAFREZZO, the difference is visible in the dis- tance between neighbouring whorls (h), the shape of the pri- maries, and in clearly visible and well-developed, rather massive, secondaries in both these species. With regard to Suppiluliumaella transdanubica (DRAGASTAN) and S. al- imani (DRAGASTAN), of which the latter does not show any remarkable difference to the type-species of Suppiluliu- maella, the differences concern the value of the inner diam- eter, shape of the primaries, and independent growth of the secondaries out of the primaries’ distal ends, as distinct from these characteristics being well-developed in the mentioned species. With regard to S. leini DRAGASTAN and S. riedeli DRAGASTAN, their unclear photographic illustrations make direct comparison impossible. The differences to S. elliotti BAKALOVA and S. schroederi BARATTOLO con- cern the shape and massiveness of the primary branches, the relationship between the inner and outer diameters, and the growth form of the secondaries, regarding the same features in the new species. Geologia Croatica 68/3Geologia Croatica 176 ACKNOWLEDGEMENT The authors would like to thank reviewers Felix SCHLA- GINT WEIT (Munich) and Bruno GRANIER (Brest) for their very helpful and constructive comments which improved the manuscript. This study was supported by the Ministry of Sci- ence, Education and Sports of the Republic of Croatia through Projects 181-1811096-1093, 181-1951126-1134, and 195- 1953068-0242. REFERENCES BARATTOLO, F. (1984): Osservazioni su Suppiluliumaella schroederi n. sp. (alghe verdi, Dasicladali) del Cenomaniano del Matese (Appennino me- ridionale, Italia).– Boll. Soc. natur. Napoli, 92 (1983), 1–47. BASSOULLET, J.P., BERNIER, P., CONRAD, M.A., DELOFFRE, R. & JAFFREZO, M. (1978): Les Algues Dasycladacées du Jurassique et du Crétacé.– Géobios, Mém. spéc, 2, 330 p. BERNIER, P. (1978): Une nouvelle algue Dasycladacee du Portlandien du Jura meridional francais: Montenegrella florifera nov. sp.– Geobios, 11/1, 141–147. BUCUR I.I. 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(eds.): Proceedings XVIIIth Congress of the Carpathian- Balkan Geological Association (2006), 527–530. SOKAČ, B. (2004): On some Peri-Mediterranean Cretaceous dasyclad spe- cies (calcareous algae; Dasycladales) previously assigned to different genera.– Geol. Croatica, 57/1, 15–53. SOKAČ, B. (2005): Linoporella vesiculifera n.sp., a new calcareous alga (Dasycladales) from the Upper Barremian of Mt. Biokovo (Karst Dinar- ides, Croatia).– Geol. Croat., 58/2, 119–131. SOKAČ, B. (2007): Stop 11 – Upper Hauterivian algae – south of Mali Tro- glav.– In: GRGASOVIĆ, T. & VLAHOVIĆ, I. (eds.): 9th International Symposium on Fossil Algae – Croatia 2007, Field Trip Guidebook and Abstracts. Croatian Geological Survey, Zagreb, 79–81. SOKAČ, B. & GRGASOVIĆ, T. (2004): Megaporella nikleri n.sp. new cal- careous alga (Dasycladales) from the Upper Barremian of Mt. Biokovo, Croatia.– Riv. Ital. Paleont. Strat., 11/3, 651–658. SOKAČ, B. & GRGASOVIĆ, T. (2008): On some Salpingoporella species from the Lower Cretaceous of Dinaric karst.– Geol. Croat., 61/2–3, 251–271. SOKAČ, B. & NIKLER, L. (1973): Calcareous algae from the Lower Creta- ceous of the environs of Nikšić, Crna Gora (Montenegro).– Paleont. ju- goslav., 13, 7–57. VELIĆ, I. (2007): Stratigraphy and Palaeobiogeography of Mesozoic Ben- thic Foraminifera of the Karst Dinarides (SE Europe).– Geologia Croat- ica, 60/1, 1–113, 26 pl. Manuscript received July 23, 2015 Revised manuscript accepted September 16, 2015 Available online October 31, 2015 Figure 1: (A) General location of the sample KJ–18 locality in the Croatian territory. (B) Simplified map of the area of the central Biokovo Mt., type locality of Montenegrella? gracilis n.sp. (from SOKAČ, 2005). Sokač & Grgasović: Montenegrella? gracilis n. sp., a new calcareous alga (Dasycladales) from the Upper Barremian of Mt. Biokovo (Dinarides Mts., Croatia) Geologia Croatica 177 PLATE I Montenegrella? gracilis n. sp. 1 Longitudinal-oblique section. Slide KJ-18/4, x22. 2 Oblique section. Slide KJ-18/141, x14. 3 Tangential-longitudinal section. Slide KJ-18/142, x17. 4 Transverse-oblique section. Holotype. Slide KJ-18/146, x17. 5–6 Longitudinal-tangential sections. Slides: Fig. 5, KJ-18/245, x22; Fig. 6, KJ-18/242, x14. 7 A Montenegrella? gracilis n. sp., transverse section. B Cylindroporella lyrata MASSE & LUPERTO-SINNI, longitudinal-tangential section. Slide KJ-18/93, x14. 8 Transverse section (fragment). Slide KJ-18/216, x22. Geologia Croatica 68/3Geologia Croatica 178 PLATE II Montenegrella? gracilis n. sp. 1 Oblique-tangential section. Slide KJ-18/27, x17. 2–3 Oblique sections. Slides: Fig. 2, KJ-18/229, x14; Fig. 3, KJ-18/156, x14. 4 Tangential section. Slide KJ-18/218, x17. 5–7 Transverse sections. Slides: Fig. 5, KJ-18/64, x14; Fig. 6, KJ-18/183, x17; Fig. 7, KJ-18/215, x22. 8 Part of oblique section. Slide: Kj-18/16, x22. 9 A Cymopolia velici SOKAČ & NIKLER, transverse section. B Montenegrella? gracilis n. sp., fragment. Section KJ-18/214, x22.