Mesocyparis Rosanovii</Emphasis> Sp. Nov
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ISSN 0031-0301, Paleontological Journal, 2006, Vol. 40, No. 3, pp. 328–338. © Pleiades Publishing, Inc., 2006. Original Russian Text © T.M. Kodrul, M.V. Tekleva, V.A. Krassilov, 2006, published in Paleontologicheskii Zhurnal, 2006, No. 3, pp. 93–102. A New Conifer Species, Mesocyparis rosanovii sp. nov. (Cupressaceae, Coniferales), and Transberingian Floristic Connections T. M. Kodrula, M. V. Teklevab, and V. A. Krassilovc aGeological Institute, Russian Academy of Sciences, Pyzhevskii per. 7, Moscow, 109017 Russia e-mail: [email protected] bPaleontological Institute of Russian Academy of Sciences, ul. Profsoyuznaya 123, Moscow, 117997 Russia cInstitute of Evolution, University of Haifa, Mount Carmel 31905, Haifa, Israel Received October 25, 2005 Abstract—A new species, Mesocyparis rosanovii sp. nov. from the Lower Paleocene of the Amur Region, Rus- sia, is characterized by a combination of advanced and primitive features: mixed opposite-alternate branching of vegetative shoots, leafy microsporophylls, as well as a relatively high number of sporangia per microsporo- phyll and seeds per cone scale. The Late Cretaceous–Early Paleogene genus Mesocyparis McIver et Basinger, which obviously embraces a natural group of closely related species, was a significant component of the vege- tation in the middle and high latitudes of eastern Asia and western North America. The range of the genus tes- tifies to terrestrial Transberingian connections between the continents during the Cretaceous warming phases. DOI: 10.1134/S0031030106030142 Key words: conifers, Cupressaceae, Paleocene, Amur Region, Beringia, phytogeography. INTRODUCTION from each other, which would have virtually excluded the possibility of migration. The problem of floristic exchanges between eastern In this context, phytogeographic relations between Asia and western regions of North America has long Asia and North America should be revised using mod- attracted the attention of phytogeographers. If in the ern paleobotanical methods. The similarity between the geological past such exchanges occurred through Ber- Paleocene floras of the western United States and Can- ingia (land mass that repeatedly appeared on the Berin- ada, on the one hand, and the Chukchi Peninsula, gian shelf), they show that a land bridge for migration Sakhalin Island, and the Amur Region, on the other of terrestrial organisms existed during this geological hand, was noted by many researchers. The Paleocene epoch, and, therefore, should be taken into account in flora of Tsagayan (Amur Region) shares more than a tectonic reconstructions. The floristic connections via half of its species with the floras of North America Beringia are also important in the context of paleocli- (Krishtofovich, 1939; Krassilov, 1976a). However, a matology, since even during the period of existence of significant proportion of these species are artificial taxa the land bridge the climate at high latitudes impeded the that remain unstudied in morphological detail. There- migration of thermophilic species. The exception to fore, taxonomic revision of this major East Asiatic fos- this rule were periods of warm polar climate, during sil flora and the analysis of the resulting phytogeo- which plants of temperate and even subtropical zones graphic concepts are highly necessary. might have reached Beringia. Along with other charac- To accomplish such a revision, we are going to teristics, mass Transberingian migrations that included study, in the first place, the well-preserved plant thermophiles can serve as indicators of such periods. remains from the newly found localities of the Amur Region, the richest of which are outcropped in the One of the warm polar periods took place during the Arkhara-Boguchan open-pit coal mine. Among fossil Campanian of the Late Cretaceous (Jenkyns et al., plants collected from this locality, coniferous shoots 2004), when the floras of Alaska and northeastern Rus- and cones that belong to the family Cupressaceae and sia had many elements in common (Krassilov, 1975, were previously included in the morphological group 1976a, 1976b, 1979). After the cooling at the Creta- Cupressinocladus interruptus (Newberry) Schweitzer ceous/Paleogene boundary, the global Late Paleocene stand out because of their good preservation. The appli- warming also might have favored Transberingian cation of light and electron microscopy allows these migrations. However, some tectonic reconstructions remains to be classified in terms of the natural system claim that in the Late Cretaceous and Early Paleogene and to be compared with recently described closely Asian and American parts of Beringia were still remote related species of North America and northeastern Russia. 328 A NEW CONIFER SPECIES, MESOCYPARIS ROSANOVII SP. NOV. 329 MATERIAL AND METHODS branches at least twice as long as on ultimate branches. The material comes from the coal-bearing deposits Facial and lateral leaves approximately equal in length; of Early Paleocene age that are exposed in the open-pit facial leaves flatly adpressed, thickened along midline, coal mines of the Arkhara-Boguchan brown coal field, typically keeled, apices acute to acuminate, thickened; 15 km southeast of Arkhara in the Amur Region lateral leaves folded, decurrent, free portion narrowly (49°18′52.3″ N, 130°12′42.7″ E). Plant remains occur spreading, acute, clasping facial leaves or subtending in bluish gray clays that overlie the coal layer “Nizhnii” branches of higher order. Leaves of axes bearing seed (Akhmetiev et al., 2002; Markevich et al., 2004). The cones of one type, decurrent, distally free, falcate. cupressaceous shoots and cones are preserved as Leaves amphistomatic, stomata in irregular bands on impressions with coaly compressions. Pollen cones are abaxial side of facial leaves, mostly oblique to midline. either attached to the shoots or preserved detached, Stomata mono- or (rarely) amphicyclic, with typically associating with the vegetative shoots. Isolated five subsidiary cells, which are indistinctly differenti- microsporophylls of disintegrated pollen cones fre- ated into polar and lateral cells, often contiguous. Tri- quently occur scattered on the bedding plane. Seed chomes as irregularly developed small pits on subsid- cones are partly preserved as casts of inner cavities iary and ordinary cells. between the scales. The compressions were macerated Pollen cones up to 7 mm long, 1–4 mm in diameter, with concentrated nitric acid. We also succeeded in terminal on ultimate branchlets, consisting of three to extracting pollen grains from sporangia. five decussate pairs of peltate microsporophylls. Peltae The remains of shoots and cones were studied mor- of microsporophylls leafy, 1–2.2 mm long, 0.5–2 mm phologically using a Leica MZ6 stereomicroscope. The wide; larger (proximal) pelta with stomata. Sporangia macerated cuticles were studied in transmitted light abaxial, two to five at base of peltae, globose to obo- vate, 0.4–0.5 mm long, 0.35–0.45 mm in diameter. Pol- under an Axioplan 2 compound microscope. Leaf cuti- µ cles, microsporophylls, and inner casts of seed scales len about 20 m in diameter, spherical to slightly ellip- were mounted on SEM stubs and covered with gold for tical, irregularly gemmate, with leptomatic germinal study under a CAMSCAN SEM. Photographs were area of reduced ectexine discernible under TEM. made using Leica DFC 320 and Nikon Coolpix 8700 Seed cones erect, globose to oblong, 3.5–5 mm digital cameras. long, 3–4 mm in diameter, borne on short stalks in one Pollen grains were prepared for transmission elec- to six opposite pairs in two side rows or decussate over tron microscopy after Meyer-Melikian and Tel’nova fertile zones of penultimate axes, which is leafy above. (1990). Ultrathin sections were made with a ultramicro- Fertile cone scales four in two decussate pairs, approx- tome (LKB-3). Some of the sections obtained were imately equal in size, woody, subpeltate, umbo subapi- additionally stained with lead citrate. Ultramicrographs cal with prominent, erect point. Seeds three to five per were obtained using a TEM (Jeol 100 B). scale, inverted on short funiculus, bilaterally winged. The collection of several hundred vegetative shoots, Description (Figs. 1–3). Vegetative shoots are pollen and seed cones, and microsporophylls with spo- plagiotropic; ultimate branchlets are prostrate in one rangia is housed in the Geological Institute of the Rus- plane, leafy, mostly opposite or, more rarely, alternate. sian Academy of Sciences (GIN), no. 4867. Most shoot remains bear branches of two or three orders. The branches may be attached oppositely or alternately. Ultimate branchlets are often present only SYSTEMATIC PALEOBOTANY on the acroscopic side facing the shoot apex (Fig. 1a). Family Cupressaceae Richard ex Bartling, 1830 Leaves of vegetative shoots are scaly, decussate, and dimorphic: facial and lateral. Genus Mesocyparis McIver et Basinger, 1987 Mesocyparis rosanovii Kodrul, Tekleva et Krassilov, sp. nov. The axes of the first order 4–10 cm long bear up to 16 branches, which are borne in the axils of lateral Plate 11, figs. 1–6, Plate 12, figs. 1–8 leaves at an angle of 20°–45° and at intervals of 0.5– Etymology. In honor of Prof. A.Yu. Rozanov, 0.7 cm and are up to 2.5 cm long. Facial leaves on thick acknowledging his contribution to the development of axes are 4–6 mm long and 0.8–1 mm wide, flatly paleobotanical research at the Paleontological Institute adpressed, linear, slightly expanded in the upper por- of the Russian Academy of Sciences. tion, with an acute