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3. Pottiodicranum, a New Moss Genus from the Late Eocene Мхи Из Ровенского Янтаря (Украина), 3

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3. Pottiodicranum, a New Moss Genus from the Late Eocene Мхи Из Ровенского Янтаря (Украина), 3 Arctoa (2016) 25: 229–235 doi: 10.15298/arctoa.25.18 MOSSES FROM ROVNO AMBER (UKRAINE), 3. POTTIODICRANUM, A NEW MOSS GENUS FROM THE LATE EOCENE МХИ ИЗ РОВЕНСКОГО ЯНТАРЯ (УКРАИНА), 3. POTTIODICRANUM, НОВЫЙ РОД МХОВ ИЗ ПОЗДНЕГО ЭОЦЕНА MICHAEL S. IGNATOV1,2, ALFONS SCHÄFER-VERWIMP3 & EVGENY E. PERKOVSKY4 & JOCHEN HEINRICHS5 МИХАИЛ С. ИГНАТОВ1,2, АЛЬФОНС ШЭФЕР-ФЕРВИМП3, ЕВГЕНИЙ Э. ПЕРКОВСКИЙ4, ЙОХЕН ХАЙНРИКС5 Abstract Pottiodicranum gen. nov. is described based on an inclusion in Late Eocene Rovno amber. The fossil is characterized by an acrocarpous habit, common for many genera of haplolepideous mosses, but differs from all known Paleogene amber fossils of acrocarpous mosses in the presence of several high solid papillae above the lumen of the lamina cells. The papillae are present on the dorsal and ventral side of the lamina and resemble modern Pottiaceae species. However, Pottiodicranum papillosum has a striate dorsal costa surface consisting of low ridges similar to those of some species of Dicranaceae s.l., especially Campylopus and Dicranum. However modern Dicranaceae lack this kind of papillae, while Pottiaceae may have lamellae on the ventral side of the costa, but not on the dorsal side. Thus the Eocene moss fossil possesses a unique combination of characters and apparently belongs to an extinct group of Dicranidae, while the family relationships remain unclear. Резюме Из познеэоценового ровненского янтаря описан новый род мхов, Pottiodicranum gen. nov. В целом по своему строению он сходен с верхоплодными мхами, особенно представляющими группу гаплолепидных мхов, или подкласс Dicranidae. В то же время, он отличается от всех прочих верхоплодных мхов, извеcтных из янтарей палеогеновых отложений, наличием нескольких высоких цельных папилл над просветом клеток листа. Папиллы развиты как на дорсальной, так и на вентральной сторонах листа и весьма напоминают по своему строению папиллы многих современных родов семейства Pottiaceae. В то же время, Pottiodicranum papillosum, единственный вид рода, имеет на дорсальной стороне жилки гребни, а также ясную продольную исчерченность, известную у современных мхов семейства Dicranaceae s.l., особенно у Campylopus и Dicranum. В то же время у представителей Dicranaceae не бывает таких папилл, а в семействе Pottiaceae неизвестны пластиночки или гребни на дорсальной стороне листа. Хотя все признаки Pottio- dicranum известны в подклассе Dicranidae, их необычная комбинация делает род уникальным. KEYWORDS: bryophytes, mosses, fossils, Dicranaceae, Pottiaceae, Dicranales INTRODUCTION four represent form genera. A few inclusions were classi- Study of moss inclusions in Paleogene amber initiat- fied to family level only, and a few taxa were not consid- ed in the 19th century and was at first limited to Baltic ered because the original material was not available. amber (Goeppert & Berendt, 1845; Goeppert, 1853; Cas- The first mosses from Late Eocene Ucrainian Rovno pary & Klebs, 1907). A second source of Paleogene moss amber were reported only recently (Ignatov & Perkovsky, fossils – Bitterfeld amber – became available in the sec- 2011, 2013) and so far include representatives of ten gen- ond half of the 20th century (Liehmann, 2013). Frahm era (extant ones are marked with an asterisk): *Hypnum (2004, 2010) extensively studied the mosses in Baltic and Hedw., Sematophyllites J.-P. Frahm, *Hypnodontopsis Z. Bitterfeld amber and assigned them to 33 genera in some Iwats. & Nog., Calymperites Ignatov & Perkovsky, *Tra- 25 families. Twenty seven of these genera are extant and chycystis Lindb., Neckerites Ignatov & Perkovsky, 1 – Moscow State University, Faculty of Biology, Moscow 119234, Russia; e-mail: [email protected] 2 – Main Botanical Garden, Russian Academy of Sciences, Botanicheskaya 4, Moscow 127276 Russia 3 – Mittlere Letten 11, 88634 Herdwangen-Schönach, Germany; e-mail: [email protected] 4 –Schmalhausen Institute of Zoology of the Ukrainian National Academy of Sciences, B. Khmelnitsky str., 15, Kiev 01-106, Ukraine; e-mail: [email protected] 5 –Systematic Botany and Mycology, and GeoBio-Center, Ludwig Maximilians University Munich, Menzinger Str. 67, 80638 Munich, Germany; e-mail: [email protected] 230 M.S. IGNATOV, A. SCHÄFER-VERWIMP, E.E. PERKOVSKY & J. HEINRICHS Ê 2 Ê Ê Ê 200 m Ê Ê 200 m 3–4 1 2 100 m 200 m 3 4 Figs. 1–4. Pottiodicranum papillosum sp. nov. (from SIZK-K-10070): 1: Holotype, total view; details of further images arrowed and numbered; 2: upper part of leaf with retained apical part; 3–4: leaves of lower portion of shoot, views at different angles, contrasting areolation (3) and ridged dorsal costa (4). Pottiodicranum, a new moss genus from Rovno amber of Late Eocene 231 50 m 100 m 5 6 Figs. 5–6. Pottiodicranum papillosum sp. nov. (from SIZK-K-10070): dorsal side of leaves showing median (5) and median to lower parts, note the increase of cell size towards the leaf base (6). Note also the ridged dorsal side of costa. *Palamocladium Müll. Hal., *Ctenidium (Schimp.) Mitt., MATERIAL AND METHODS *Tristichella Dixon, and *Isopterygium Mitt. The late Eocene amber specimen SIZK-K-10070 orig- Most of these genera have also been recognized in inates from the Pugach quarry (Klesov) in the Rovno Baltic and Bitterfield amber, and despite some differenc- Province of the Ukraine. Detailed information on the lo- es in the species composition of these ambers, the moss cality and its age is available from Ignatov & Perkovsky floras resemble each other (1) in the prevalence of pleu- (2011). Having about the same age as Baltic amber (Perk- rocarps: seven out of ten moss genera in Rovno amber, ovsky et al., 2003, 2007, 2010), Rovno amber comprises and 22 out of 33 in Baltic and Bitterfield amber; and (2) a somewhat more southern fauna (Dlussky & Rasnitsyn, the presence of well-recognizable species, e.g. represen- 2009; Naden et al., 2016; Perkovsky 2011, 2016). So far, tatives of the genera Trachycystis and Hypnodontopsis. only a few fossils of bryophytes have been recognized. So far only a single acrocarpous moss from Rovno am- Specimen SIZK-K-10070 was studied under a com- ber is unknown from Baltic and Bitterfield amber: Ca- pound light microscope Olympus CX41 and a stereomi- lymperites ucrainicus Ignatov & Perkovsky (Ignatov & croscope Olympus SZX16 equipped with an Infinity 4 Perkovsky, 2013). A second fossil genus which is so far digital camera, and also with a Carl Zeiss AxioScope A1 unique to Rovno amber is described below. compound microscope equipped with a Canon 60D dig- 232 M.S. IGNATOV, A. SCHÄFER-VERWIMP, E.E. PERKOVSKY & J. HEINRICHS 20 m 20 m 7 8 20 m 20 m 9 10 Figs. 7–10. Pottiodicranum papillosum sp. nov. (from SIZK-K-10070): dorsal sides of leaves in upper part, magnified to show areolation and papillosity patterns, better seen at margins. The presence of several papillae per cell is especially apparent from the compari- son of cell size and papillae density. ‘10’ shows irregularly isodiametric laminal cells and rectangular costal cells. Consider that ‘10’ shows an area just a little to the rigth of ‘9’ where costal cells are seen with poor focus at the right margin of the picture. Pottiodicranum, a new moss genus from Rovno amber of Late Eocene 233 100 m 11 Figs. 11–12. Pottiodicranum papillosum sp. nov. (from SIZK-K-10070): upper portions of leaves in dorsal view show- ing leaf areolation and papillosity. The presence of several pa- 50 m pillae per cell is apparent from the comparison of cell size and density of papillae. ‘12’ shows irregularly isodiametric lamina 12 cells and rectangular costa cells ital camera. Incident and transmitted light were often pillae per leaf cell do not occur among extant represen- used simultaneously. Most micrographs were obtained from tatives of Dicranaceae s.l. A few genera of Dicranaceae a series of optical sections and composed using the soft- s.l. such as Amphidium Nees (Amphidiaceae or Rhabdo- ware package HeliconFocus 4.50 (Kozub et al., 2008) for weisiaceae, if a narrow family concept is applied) have a better illustration of the three-dimensional inclusion. papillose cells yet their papillae differ from those of Pot- TAXONOMY tiodicranum in size and shape. Papillose cells are also known in some members of the large tropical moss fam- Genus Pottiodicranum gen. nov. [order Dicranales, ily Calymperaceae, which is also nested in Dicranales family insertae sedis] s.l. (i.e., including Pottiaceae). However, the dorsal sur- Description: Stem erect; densely foliated, terete. face of the costa of extant Calymperaceae is smooth or Leaves erect, oblong-lanceolate, gradually acuminate; dentate to spinose but not lamellate or striate/ridged and, costa strong, percurrent, on dorsal surface striate to in addition, most species of this family have coarsely ser- ridged; margins entire; lamina cells irregularly round- rate leaf margins and very large, hyaline and thin-walled quadrate, moderately thick-walled, with several simple, basal leaf cells (cancellinae). The papillae of some spe- high cylindric papillae per cell; basal cells slightly en- cies of Orthotrichaceae are very similar to those of Pot- larged, short rectangular. tiodicranum, however, their costae are never ridged. Etymology: The name of the genus points to a com- Ridged to almost winged costae occur in several species of bination of characters of two extant families, Dicranace- Grimmia (Grimmiaceae), but most species of this family ae (s.l) and Pottiaceae. have smooth cells. Round papillae are known, e.g., in G. Type species: Pottiodicranum papillosum sp. nov. caespiticia (Brid.) Jur. and G. elatior Bruch ex Bals.-Griv. Comparison: Pottiodicranum has the most prominent & De Not., but their cells are never pluripapillose and the papillae among the Paleogene moss fossils. Among ex- papillae are restricted to the lamina cells of the distal leaf tant bryophytes, this type of papillae commonly occurs portion. Pluripapillose cells are known from species of the in the species-rich family Pottiaceae. The vast majority Grimmiaceae genus Niphotrichum (Bedn.-Ochyra) Bedn.- of its 1500 extant species has papillose lamina cells Ochyra & Ochyra (or the Racomitrium canescens-group), (Zander, 1993).
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