Reproductive Phases of Miocene Algae from Central Paratethys and Their Bearing on Systematics
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Reproductive phases of Miocene algae from central Paratethys and their bearing on systematics JURAJ HRABOVSKÝ Hrabovský, J. 2019. Reproductive phases of Miocene algae from central Paratethys and their bearing on systematic. Acta Palaeontologica Polonica 64 (2): 417–439. Coralline algae were widespread in shallow-water environments during the middle Miocene in the central Paratethys. Their distribution and diversity patterns are useful in paleoenvironmental and paleoclimatic studies providing that spe- cies identifications are accurate and robust. Taxonomy of fossil coralline algae was traditionally based on morphology of bi/tetrasporophytes whereas taxonomy of extant species is additionally supported by characters that can be observed in the remaining life cycle phases. Therefore, investigations of such characters in fossil record are also essential to avoid misidentifications of ancient coralline algae. Here, I analyse morphology of reproductive phases in subfamilies Hydrolithoideae and Mastophoroideae from the middle Miocene of the central Paratethys. Analyses of published and newly collected hydrolithoid and mastophoroid coralline algae allow: (i) identification of three species of the ge- nus Hydrolithon and two species of the genus Lithoporella; (ii) description of complete life cycle phases for extinct Hydrolithon lemoinei, H. corculumis, and Hydrolithon sp. 1 and (iii) description of incomplete life cycles for Lithoporella minus and Lithoporella sp. 1. The results also prove that characters associated with trichocytes can be used in some fossil coralline algae, and bi/tetrasporic pore canal anatomy is reliable diagnostic character also for fossil species, as it is used in extant coralline algae of the order Corallinales. Key words: Corallinales, Hydrolithon, Lithoporella, reproductive phases, Miocene, central Paratethys, Slovakia, Romania. Juraj Hrabovský [[email protected]], Earth Science Institute of the Slovak Academy of Sciences, Dúbravská cesta 9, 84005 Bratislava, Slovakia. Received 4 December 2018, accepted 15 March 2019, available online 10 May 2019. Copyright © 2019 J. Hrabovský. This is an open-access article distributed under the terms of the Creative Commons Attribution License (for details please see http://creativecommons.org/licenses/by/4.0/), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. deposits indicate rather warmer climate conditions during Introduction the middle Miocene climatic optimum with analogues in the Recent Red Sea (Riegl and Piller 2000). Despite many anal- Coralline algae are commonly dominant rock-build- yses from the last century (Schaleková 1969, 1973 and 1978; ing organisms of the middle Miocene red-algal limestone Schaleková and Čierna 1985; Golonka 1981; Pisera 1985; (Schaleková 1962, 1978; Pisera 1985, 1996; Studencki 1988a, Pisera and Studencki 1989; Studencki 1988b, Zdražílková 1999; Doláková et al. 2008). The presence of coralline al- 1988), temporal and spatial changes in coralline algae diver- gae in analysed fossil associations is of high importance sity and distribution remain poorly known in the Miocene of for palaeoecological analyses as their distribution is largely the central Paratethys. This insufficiency in our knowledge controlled by light and temperature (Adey and McKibbin has several reasons, including: (i) taxonomic changes caused 1970; Adey et al. 1982; Athanasiadis and Ballantine 2014). by the type collections revision, new analytic techniques The composition of a rich coralline algae limestone is vari- and discrimination of new diagnostic characters were ap- able in the local and regional scales of the central Paratethys plied to few taxa only (Woelkerling 1988; Braga et al. 1993; (Baráth 1992; Baráth et al. 1994; Pisera 1996; Riegl and Basso et al. 1996); (ii) molecular taxonomy results can some- Piller 2000). Recent Adriatic Sea, Tyrrhenian Sea, or Eastern times contradict morphology-based taxonomy (Penrose and Mediterranean Sea assemblages are perceived as analogues Woelkerling 1992; Harvey et al. 2006; Bahia et al. 2014; to the late Badenian (early Serravallian) Paratethyan assem- Kato et al. 2011; Rösler et al. 2016); (iii) differences in the blages (Pisera 1985; Seneš and Ondrejčíková 1991; Basso treatment of morphological characters between biologists 1998; Basso et al. 2008). In contrast, coral carpets or patch and palaeontologists: classification of fossil coralline algae is reefs occurring mainly in the early Badenian (Langhian) typically based on bi/tetrasporophytes morphology, whereas Acta Palaeontol. Pol. 64 (2): 417–439, 2019 https://doi.org/10.4202/app.00579.2018 418 ACTA PALAEONTOLOGICA POLONICA 64 (2), 2019 extant species taxonomy is also supported by characters of Material and methods other reproductive phases (Braga et al. 2005; Athanasiadis et al. 2004; Athanasiadis and Ballantine 2014; Bahia et al. Studied material represents: (i) the collection published in 2015). The recognition and detection of gametangial and car- Schaleková (1969, 1973, 1978), Schaleková and Čierna (1985), posporangial life cycles stages in fossil coralline algae can Novák (1975), and Zdražílková (1988), (ii) new samples from solve some of these difficulties (Basso et al. 1996; Basso and selected localities in the Central Paratethys, including Lopadea Rodondi 2006; Hrabovský et al. 2015; Chelaru and Bucur Veche in Romania (Transylvania Basin), Maksymivka in 2016). Therefore, revision of published data and analyses Ukraine (Carpathian Foredeep), Modrý Majer, Sazdice, and of the newly collected material from the central Paratethys Veľký Pesek in Slovakia (Danube Basin), Devínska Kobyla, scoped on gametophyte, carposporophyte bearing plants and Vývrat, Rohožník, and Stupava, in Slovakia (Vienna Basin) bi/tetrasporophyte life phases are necessary. and (iii) personal collections of Daniel Pivko from Department Life cycle phases morphology of fossil coralline algae of Paleontology and Geology of the Comenius University were reported only sporadically in the 20th century (Conti 1943; Mastrorilli 1968). However, since the end of the Table 1. The list of the studied material. Numbers indicate number of 20th century fossil specimens of the genera Lithophyllum thin sections. (Basso et al. 1996), Hydrolithon (Woelkerling et al. 2012), Reference Neogoniolithon (Quaranta et al. 2007), Spongites (Chelaru Zdražílková Schale- Locality Pivko and Bucur 2016), Lithothamnion (Chelaru and Bucur 2016), this paper 1988, ková 1969, 2017 Phymatolithon (Basso et al. 1997; Hrabovský et al. 2015), Novák 1975 1973, 1985 Mesophyllum (Hrabovský et al. 2015; Coletti et al. 2018), Vienna Basin and in the Sporolithon (Vannucci et al. 2000; Hrabovský et Devínska Kobyla 28 28 44 al. 2015; Chelaru and Bucur 2016) have been documented. Stupava 4 The mode of the gametangial conceptacles development Vývrat 7 is comparable to bi/tetrasporangial conceptacles in some Rohožník 17 50 coralline algal genera formerly assigned to the subfamily Wolfsthal 8 Mikulov 3 Mastophoroideae Setchell, 1943. Many of these genera were Hlohovec 1 assumed to represent bi/tetrasporophytes (Quaranta et al. Müllendorf 2 2007; Basso et al. 1996; Basso and Rodondi 2006). Bi/tet- Danube Basin rasporophytes with this type of conceptacles inhabit shal- Modrý Majer 9 27 low to medium depths in the Recent tropics, subtropics and Kamenica nad 13 warm temperate seas (Aguirre et al. 2000; Adey et al. 1982). Hronom Therefore, misidentification of gametangial with bi/tetra- Pavlová 4 sporangial conceptacles could distort an information about Čierny Hill 11 coralline algae diversity and distribution. Štúrovo 4 Here, I focus on the genera from the subfamilies Hydro - Veľký Pesek 7 lithoideae Kato and Baba in Kato et al., 2011 and Masto- Sazdice 5 phoroideae Setchell, 1943 from the middle Miocene of Novohrad Basin the Carpathian Foredeep, Vienna Basin, Danube Basin, Kosihovce 8 Novohrad Basin, and Transylvania Basin. The work purpose Transylvania Basin is to: (i) describe specimens both in terms of gametangial Lopadea Veche 48 and carposporangial phases as is possible, (ii) discriminate Carpathian Foredeep species among a members belonging to the subfamilies Pratecký Vrch 23 Hydrolithoideae and Mastophoroideae in the fossil record Lomnice 7 Blučina 4 and (iii) to provide the identification key for fossil hydro- Telnice 1 lithoid and mastophoroid species. Holubice 2 Institutional abbreviations.—NHM, Natural History Mu- Vyškov 1 seum; VEGA, Vedecká Grantová Agentúra (Scientific Ochoz 1 Kralice nad Grant Agency) internal grant system of the Ministry of 1 Oslavou Education, Research and Sport of the Slovak Republic, and Rebešovice 3 3 Slovak Academy of Sciences. Hostim 1 Other abbreviations.—D, diameter; H, height; L, length; Hluchov 1 MSi, Miocene Stable isotopes (event); SD, standard devia- Židlochovice 12 tion; TS, thin section(s); W, wide. Maksymivka 7 HRABOVSKÝ—MIOCENE ALGAE FROM CENTRAL PARATETHYS 419 14° 28° 16° 26° 18° 20° 22° 24° 32 31 50° Kraków Carpathian Foredeep Lviv 29 17,18 15,16 12,13 19 Brno 49° 14 Western 20 8–11 Carpathians 7 Košice Transcarpathian Basin 1,2 Carpathian Foredeep 28 3 4 25 26 48° Wien Vienna Basin Bratislava 21,22 Novohrad Basin 5 23,24 CarpathiansEastern Eastern 6 Nyirseg Basin Alps 27 Danube Basin Budapest Graz 47° Great Hungarian Plain Styrian Basin Cluj Zala Basin Bekes Basin 30 Mura Basin Mako Basin 46° Transylvania Basin Southern North Croatia Alps Zagreb Basin Bačka Basin Banat Basin 45° Southern Carpathians Kolubara Basin