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Ditaxocladus (extinct Cupressaceae, Cupressoideae) from the Upper Cretaceous and Paleocene of the Northern Hemisphere
Abstract
An emended diagnosis of the extinct cupressoid genus Ditaxocladus S. X. Guo & Z. H. Sun (syn. FokieniopsisMcIver & Basinger) and its three species is proposed based on new collections from the type locality in the Paleocene Wulonggu Formation of Altai in NW China, and other sites from China, SE Russia and western North America. Ditaxocladus planiphyllus S. X. Guo & Z. H. Sun (generitype) is now known from compound racemose fertile branches bearing several opposite pairs of globose to sub-globose seed cones composed of 6-12 decussate peltate scales, in addition to typically oppositely branched sprays. Besides D. planiphyllus, two more species are recognized based on differences in seed cone morphology. D. kivdensis Kodrul in Krassilov et al. from the Paleocene of Raichikha in Amur Province, Russia, is distinguished by similar but slender and pendulous cones, while D. catenulatus (W. A. Bell) S. X. Guo, Kvaek, Manchester & Z. K. Zhou comb. n. (syn. Fokieniopsis catenulata (W. A. Bell) McIver & Basinger and Fokienia ravenscragensisMcIver & Basinger) typified by early Paleocene records in NorthAmerica bears ovoid seed cones with a higher number of cone scales. All three species share the same type of bipinnately and oppositely ramified elongate sprays of cladode-like foliage and pinnately and oppositely arranged seed cones at successive nodes on compound racemose fertile branches. They differ from the living Fokienia, with which they were formerly compared, in oppositely (vs. alternately) branched leafy sprays and oppositely clustered (vs. solitary) seed cones consisting of much fewer cone scales (6-12 vs. 12-18 in Fokienia). Sterile foliage attributed to Ditaxocladus occurs in the Upper Cretaceous to Paleocene of North America, Russia and Paleocene of Spitsbergen. Tracing differentiation of Ditaxocladus populations (or species) over the Northern Hemisphere is hampered by incompleteness of most records based on quite uniform foliage without seed cones. Tetraclinis salicornioides (Unger) Kvaek from the Eocene to Pliocene of Europe and the Oligocene to Miocene of North America, which shares very similar gross morphology of foliage, differs from Ditaxocladus in the quadrivalvate seed cones and papillate leaf epidermis with undulate anticlinal walls. © 2012 E. Schweizerbart'sche Verlagsbuchhandlung, Stuttgart, Germany.
... Amber inclusions representing cupressoid foliage shoots are commonly short, unbranched portions, likely branchlets of (sub)ultimate order. Some genera of the Cupressaceae display distinct branching pattern, e.g., opposite in fossil Ditaxocladus and in extant Tetraclinis, but alternate in Fokienia (Guo et al. 2012). However, features of branching patterns are not discernible in the amber inclusions, since they only represent short shoot fragments. ...
... Besides, T. salicornioides has different leaf shapes, i.e., lateral leaves are long, almost linear, the conduplicate laterals do not overlap at their bases, and leaf tips are mucronate. The leaf epidermis is characterized by undulate anticlinal walls observed from cuticles Guo et al. 2012). Therefore, affinities with T. salicornioides can be rejected. ...
... Small shoot portions from Baltic amber can hardly be compared with these taxa. Recently, however, Alekseev (2018) assigned Libocedrus sub decurrens sensu Caspary & Klebs (1907a;1907b) to Ditaxocladus which occurred across North America, Asia and Spitsbergen from the Late Cretaceous Table 5. (Guo et al. 2012). The foliage of Ditaxocladus is distinct from L. subdecurrens in the following features: flattened sprays which branch very symmetrical, divided into cladode-like segments, stomata occur on both sides of the spray, leaf segments triveined (Guo et al. 2012). ...
Baltic amber represents one of the most important Cenozoic fossil Lagerstätten, providing numerous fossils including plenty of invertebrates. Although it is essential for understanding the past habitats and ecology of these fossil organisms, the Baltic amber forest and its palaeoecology is not satisfactorily known for answering questions on the identity of the amber-producing conifer(s) and the specific habitat and climate conditions under which the resins were exuded. Knowledge of the Baltic amber flora mostly dates back to the 19th century and the early decades of the 20th century and needs revision on a larger scale. Here, we continue our revision and monographic treatment of conifer inclusions based on historical and new collections. On this account, type specimens and original material of the classical monographs on Baltic amber conifers were identified from museum collections, and newly discovered material complemented the study. The studied material encompasses portions of foliage shoots, single leaves and pollen cones. The preservation mode of these conifer inclusions is briefly described and discussed. Identification of the conifer remains is based on gross-morphological (shoot architecture, leaf shape and size, phyllotaxis) and leaf epidermal characters (distribution and arrangement of stomata, cells structure of leaf margin). For taxonomic purposes, we use both the modern classification system of conifers but also artificial fossil-genera, when available characters were inadequate to accommodate the fossils into ‘natural’ genera. Based on our present and previous revisions of holotypes from Baltic amber conifers, we confirm five families, 17 genera including two fossil-genera and 26 fossil-species of conifers, including the following taxa: Cupressaceae [eight genera, 14 species; Quasisequoia couttsiae Sequoia sp. aff. S. abietina Taxodium sp., aff. Glyptostrobus europaeus Cryptomeria sp., Cupressinanthus (four spp.), Cupressinocladus (four spp.), Calocedrus sp.], Pinaceae [six genera, nine species: Pinus (four spp.), Pseudolarix sp., aff. Tsuga Cathaya sp., Nothotsuga protogaea Abies sp.], Geinitziaceae (Cupressospermum saxonicum), Sciadopityaceae (Sciadopytis sp. cf. S. tertiaria) and Cephalotaxaceae (Cephalotaxus sp. cf. C. messelensis). In particular, cupressaceous foliage shoot portions are herein reconsidered and affiliated to the fossil-genus Cupressinocladus Seward 1919, and new combinations are proposed, C. borealis (Casp. et R.Klebs 1907) comb. nov., C. breynianus (Göpp. et Berendt 1845) comb. nov. et emend., C. kleinianus (Göpp. et Berendt 1845) comb. nov., and C. lamelliformis (Casp. in Caspary & Klebs 1907) comb. nov. et emend. To sufficiently accommodate cupressaceous pollen cones, we amend the fossil-genus Cupressinanthus Casp. in Caspary et Klebs 1907 and designate Cupressinanthus magnus Casp in Caspary & Klebs 1907 emend. as type. Besides, Cupressinanthus hartmannianus (Göpp. et Berendt 1845) comb nov., C. linkianus (Göpp. et Berendt 1845) comb. nov. et emend. and C. klebsii sp. nov. can be distinguished. Inclusions of Cryptomeria (Cupressaceae), Cephalotaxus (Cephalotaxaceae) and aff. Tsuga (Pinaceae) represent first evidence of these genera from the Baltic amber flora. We discuss palaeoecological, palaeofloristic and palaeoclimatic implication for the Baltic amber forest based on data from other fossil occurrences and co-occurrences of the fossil conifers. Our results show a hyperdiversity of conifer taxa which is unique among European Eocene fossil assemblages. The conifer flora from Baltic amber is distinct from both paratropical/subtropical floras from Central Europe as well as from cool-temperate floras in the Arctic zone of the early and middle Eocene. Assemblages from early-middle Miocene coastal lowland vegetation of Central Europe are most similar in composition and their palaeoecology, implying that the Baltic amber forest might have been part of the Atlantic-Boreal phytoprovince. Our survey substantiates that the Baltic amber forest was growing under a warm-temperate and humid palaeoclimate. Based on the autecology of individual fossil-species, the Baltic amber forest represented heterogeneous vegetation with a mosaic of forests and open habitats in alluvial plains, coastal swamps and mesic hinterland. In light of these new results, we discuss the age of Baltic amber and its Lagerstätte and conclude that the revealed conifer diversity supports the palynostratigraphic late Eocene age estimations.
... Guo & Z.H. Sun des Paläozäns und der Oberkreide, mit extrem dünnen, geradwandigen Epidermen, gut unterscheinbar (Guo et al. 2012). Mai & Walther (1985: 30-31) Beschreibung: Siehe die Beschreibungen bei Mai (1975: 567-575 (Mai 1975: 567-568, 572-573 (Schneider 2007). ...
... Guo & Z.H. Sun occurring in the Palaeocene and Upper Cretaceous throughout the Northern Hemisphere, which has extremely thinly cutinized, straight-walled epidermis, less distinct Florin rings and the seed cones with multiple cone scales (Guo et al. 2012). Tetraclinis brachyodon (Brongn.) ...
A complete reinvestigation of historical fossil plant collections made in the 19th century at Orsberg (Rhineland-Palatinate, late Oligocene) and published by C.O. WEBER and P. WESSEL (1851-1855) resulted in new interpretations on the fossil flora. Previous morphological and anatomical investigations of leaves and carpological are revised and now complemented leading to the following new combinations of taxa: Laurophyllum hemisphaerium (ROSELT & SCHNEIDER 1969) comb. nov., Corylopsiphyllum celtifoliurn (C.O. WEBER in P. WESSEL & C.O. WEBER 1855) comb. nov.,Ampelopsis dubia (C.O. WEBER 1852) comb. nov., Tetrastigmophyllum betulaelbrme (C.O. WEBER 1852) comb. nov., and Alnus rhenana (P. WESSEL & C.O. WEBER 1855) comb. nov. Furthermore, the flora is supplemented by new records of Magnolia, cf. Lithocarpus, Laurus, Hydrangea, ? Cleyera, Ternstroemites, Symplocos and Vaccinioides. Preliminary evaluation of the fossil material suggests two types of taphocoenoses bound to different sedimentary settings: pear forming Glyptostrobus swamp forests within the lignite and a Mixed Mesophytic Forest of the East Asian type buried in the oil shale.
... Genus DITAXOCLADUS Guo and Sun (in Guo et al., 1984) Ditaxocladus catenulatus (Bell) Guo et al., 2012 Morphotype. MT001; Fig. 5A. ...
The Cretaceous-Paleogene (K/Pg) boundary marks a mass extinction resulting in global biotic turnover. Exposures of the Hell Creek Formation in northeastern Montana contain some of the most well-studied vertebrate localities recording this mass extinction; however, very little is known of the floral record in this area. As part of an effort to reconstruct floral changes across the K/Pg in northeastern Montana, this study presents a taxonomically diverse flora from the Seafood Salad locality, located ∼ 65 m below the K/Pg boundary in the Hell Creek Formation, Garfield County, Montana. Leaves, stems, and reproductive structures (e.g., cones and seeds) are preserved as compression and impression fossils in massive, bedded siltstones and very fine sandstones. Seafood Salad is significant in that it is represents a “pre-disaster” community approximately 1.3 m.y. before the K/Pg mass extinction. We interpret the plants in these deposits as reflecting a local riparian community. The vegetation was taxonomically diverse and dominated by angiosperm trees; it also included abundant conifer specimens of a few taxa and relatively few ginkgoes and ferns. We describe 34 morphotypes and propose taxonomic affinities to modern groups and to fossil taxa from contemporaneous-age deposits in Montana and North Dakota. The Seafood Salad flora shares several taxa with other Late Cretaceous floras of the Western Interior, but substantial differences in taxonomic composition and relative abundances among these assemblages indicate that regional plant communities in the latest Cretaceous were spatially heterogeneous, rapidly changing, or both.
... Of note for age assignment is the single find of two ovulate cones of Fokienia in the Kanaka Creek flora (Fig. 6F), previously described by McIver and Basinger (1990) from the Ravenscrag flora. More recently, Guo et al. (2012) designated Fokienia a junior synonym of the extinct cupressaceous genus Ditaxocladus (2012). The foliage of Fokienia is not known from Kanaka Creek, but occurs elsewhere in the Paleocene of the Rocky Mountains and Great Plains (Brown 1962), the Ravenscrag Formation of Saskatchewan Basinger 1990, 1993), and Paleocene floras of western Alberta as Androvettia (Bell 1949) ( Table 2). ...
The late Paleocene to early Eocene sediments of Ellesmere and Axel Heiberg islands, Nunavut, of the Canadian High Arctic contain a rich fossil flora and fauna. Although the megafloral fossils have been known for more than a century, limited descriptions of the fossil flora have been presented. Here, we provide a comprehensive morphotype catalogue of fossil plants from multiple localities from Ellesmere and Axel Heiberg islands that form a systematic framework for establishing an early Paleogene polar flora from High Arctic latitudes in Canada. Described are 62 ‘dicot’ angiosperm morphotypes, three monocotyledonous angiosperms, 13 gymnosperms, and five pteridophyte morphotypes. This work presents a significant contribution to the understanding of north-polar diversity and environments during the warm greenhouse climate of the early Paleogene.
... Of note for age assignment is the single find of two ovulate cones of Fokienia in the Kanaka Creek flora (Fig. 6F), previously described by McIver and Basinger (1990) from the Ravenscrag flora. More recently, Guo et al. (2012) designated Fokienia a junior synonym of the extinct cupressaceous genus Ditaxocladus (2012). The foliage of Fokienia is not known from Kanaka Creek, but occurs elsewhere in the Paleocene of the Rocky Mountains and Great Plains (Brown 1962), the Ravenscrag Formation of Saskatchewan Basinger 1990, 1993), and Paleocene floras of western Alberta as Androvettia (Bell 1949) (Table 2). ...
Paleogene sediments of the Huntingdon Formation, a correlative to the Chuckanut Formation of neighboring Washington State, USA, are exposed in the greater Vancouver area, British Columbia, Canada. Palynology and plant macrofossils suggest the Kanaka Creek section is Paleocene rather than Eocene in age. Detrital zircon dating is less decisive, yet indicates the Kanaka rocks are no older than Maastrichtian. Analyses of plant macro- and microfossils suggest an early to middle Paleocene age for the Kanaka fossil flora. Paleocene indicators include macrofossils such as Platanus bella, Archeampelos, Hamamelites inequalis, and Ditaxocladus, and pollen taxa such as Paraalnipollenites, Triporopollenites mullensis, and Duplopollis. Paleogene taxa such as Woodwardia maxonii, Macclintockia, and Glyptostrobus dominate the flora. Fungal spores including the Late Cretaceous Pesavis parva and the Paleogene Pesavis tagluensis are notable age indicators. Physiognomy of 41 angiosperm leaf morphotypes from Kanaka Creek yields mean annual temperatures in the microthermal to lower mesothermal range (11.2 ± 4.3°C to 14.6 ± 2.7°C from LMA; 14.8 ± 2.1°C from CLAMP), with mild winters (cold month mean temperature 3.9 ± 3.4°C). Paleoclimate was cooler than the upper Paleocene and Eocene members of the Chuckanut Formation. Mean annual precipitation is estimated at ~140 cm with large uncertainties. The Kanaka paleoflora is reconstructed as a mixed conifer-broadleaf forest, sharing common taxa with other western North American Paleocene floras and growing in a temperate moist climate. Kanaka Creek is a rare coastal Paleocene plant locality that provides new insights into coastal vegetation and climate prior to the Paleocene-Eocene Thermal Maximum.
... The other specimen shows only the wing without the seed part. (Guo et al. 2012). Similar records are known from the European Palaeogene and Neogene, more rarely from the Oligocene and Miocene of North America (Kvaček et al. 2000). ...
Geology and palaeontology of the Roudniky area, situated between the Most Basin and the Česke středohoři Mountains in North Bohemia, the Czech Republic, is treated in detail. The volcano-sedimentary structure underlying the basin with the included fossiliferous layers is newly interpreted as a maar-diatreme. The radiometric age of the basalts was determined to 35.4 ± 0.9 Ma and 37.1 ± 0.9 Ma - late Eocene, Priabonian. The fish fauna consists of amiids characteristic of other adjacent late Eocene sites while the co-occurring macroflora includes many elements that are typical of the Oligocene. Closest relations are to the fauna and flora of the Větruše Hill at Usti nad Labem. Based on palaeoclimatic estimates the major turnover from subtropical to warm-temperate climatic regimes in North Bohemia appears to have initiated in the late Eocene by an increase of seasonality and the resulting vegetation change from the evergreen to mixed mesophytic forest types well before the maximum temperature drop of mean annual temperature at ca 33.5 Ma.
... The cupressaceous foliage that Brown (1962) referred to as Fokienia catenulata (BELL) R. W. BROWN (MCIVER and BASINGER 1990) was subsequently found with attached cones and placed in the extinct genus Ditaxocladus GUO et SUN. The species was transferred to Ditaxocladus catenulatus by Guo et al. (2012). Fokeniopsis MCIVER et BASINGER (1993) is a junior synonym of Ditaxocladus GUO et SUN (in Guo et al. 1984). ...
The Paleocene megafossil flora of the Rocky Mountains and Great Plains region in the United States of America, including leaves, cones, fruits, and seeds, monographed by Roland W. Brown in 1962, has been reevaluated and updated to include subsequent taxonomic revisions. The scope of this investigation included thousands of specimens from more than 450 localities of the Fort Union, Evanston, Ferris, Raton, Bear, Lebo, Melville, Ludlow, Tongue River and Sentinel Butte strata of New Mexico, Colorado, Wyoming, Montana and North and South Dakota. A large number of floristic elements remain uncertain as to their modern familial affinities due to limited diagnostic characters, or insufficient comparative investigations. Nevertheless, many of Brown’s determinations have been upheld and several newly recognized genera and families have strong support. The flora includes greater diversity of Platanaceae and Cornales than Brown had recognized. These, together with Fagales (particularly Betulaceae and Juglandaceae), Saxifragales (Trochodendroides, Archeampelos and Nyssidium), are widespread and prominent members of the flora. New combinations introduced here include Ensete goldianum (LESQUEREUX) comb. nov., Macginitiea nobilis (NEWBERRY) comb. nov., Platanites raynoldsii (NEWBERRY) comb. nov., Trochodendroides genetrix (NEWBERRY) comb nov., Cucurbitaciphyllum lobatum (KNOWLTON) comb. nov., and Mciveraephyllum nebrascense (SCHIMPER) comb. nov. Georeference data are provided for all of the localities cited by Brown. © 2014, Acta Musei Nationalis Pragae, Series B - Historia Naturalis. All right reserved.
We present the oldest known occurrences of crown-group Trochodendraceae based on new material from the Palaeocene of Wyoming, USA. Two genera are recognized, Trochodendron and Eotrochion gen. nov. The fossil fruit of Trochodendron infernense sp. nov. is represented by a pedicellate, apically dehiscent capsular fruit composed of nine follicle-like units, each bearing a persistent convex style. The basal part is ornamented with numerous raised stamen scars. From the same deposits, Eotrochion is represented by infructescences, fruits and associated leaves. The infructescences are racemes of numerous apically dehiscent capsules, each with c. 14-16 styles, each with an underlying nectary and receptacles lacking stamen scars, but possessing a prominent perianth scar. A phylogenetic assessment of the modern species, plus representatives of four extinct genera of fossil Trochodendraceae based on available morphological characters, yields a favoured topology of Trochodendron(Eotrochion(Concavistylon kvacekii(C. wehrii (Pentacentron, Tetracentron)))). A parsimony analysis of currently available characters indicates that C. wehrii renders Concavistylon non-monophyletic. Accordingly, we transfer it to Paraconcavistylon gen. nov., characterized by pendent, rather than erect infructescences. We also reconsider the extinct Nordenskioeldia (Late Cretaceous to Miocene), the prior placement of which in Trochodendraceae has been challenged, and we consider it to fall outside the crown group of the family.
The botanical composition of the ‘Baltic amber forest’ was similar to recent subtropical forests of South Eastern Asia and North America. Abundant remains of gymnosperm plants are typical inclusions in succinite but their taxonomical diversity is studied insufficiently. We confirm the presence of 30 gymnosperm species in Baltic amber. Family Pinaceae includes 14 species: Pinus succinifera (Goepp. et Berendt.) Conw., Pinus subrigida Goepp. et Menge, Pinus dolichophylla Casp., Pinus silvatica Goepp. et Menge, Pinus cembrifolia Casp., Pinus wredeana (Goepp. et Berendt) Casp., Pinus multicellularis Casp. et R. Klebs, Pinus reichiana (Goepp. et Berendt) Conw., Pinus kleinii Conw., Picea engleri Conw., Pseudotsuga sukerii (Casp. et R. Klebs) P.Alekseev comb. nov., Pseudotsuga linearis (Casp. et R. Klebs) P.Alekseev comb. nov., Tsuga kowalewskii (Caspary et R.Klebs) P.Alekseev comb. nov., Carpinites dubius Goepp. et Berendt; family Podocarpaceae includes one species Podocarpus nathorstii (Conw.) P.Alekseev comb. nov.; family Sciadopityaceae includes one species Sciadopitys tertiaria Menzel; family Cupressaceae includes 14 species: Glyptostrobus europaeus (Brongniart) Unger, Sequoia abietina (Brongniart in Cuvier) Knobloch, Cryptomeria sp., Ditaxocladus subdecurrens (Casp.) P.Alekseev comb. nov., Thujopsis breyniana (Goepp. et Berendt) P.Alekseev comb. nov., Chamaecyparis carinata (Casp. et R. Klebs) P.Alekseev comb. nov., “Chamaecyparis casparyi” R. Klebs, Chamaecyparis liukianus (Goepp. et Berendt) P.Alekseev comb. nov., Thuja succinea (Casp. et R. Klebs) P.Alekseev comb. nov., Thuja sambiensis (Casp. et R.Klebs) P.Alekseev comb. nov., Cupressus mengeana (Goepp.) P.Alekseev comb. nov., Platycladus conwentzii (R.Klebs) P.Alekseev comb. nov., “Thuites” lamelliformis Casp., “Cupressinanthus” polysuccus Casp.
The presence of Ditaxocladus subdecurrens in succinite extends both the geographic and stratigraphic ranges of this genus. This is the first finding of Ditaxocladus in Europe and the youngest (late Eocene) record for this genus. The presence of Agatis, Cedrus, Pseudolarix, Abies, Taxodium, Libocedrus, Widdringtonia and Juniperus in Baltic “amber” forest is not confirmed.
A detailed survey is presented of the early Miocene flora of the abandoned Kristina Mine at Hrádek nad Nisou in the Hrádek part of the Zittau Basin, North Bohemia. It is based on leaf morphological and anatomical study of macrofossils recovered since 1963 with additional information on carpological records. The flora belongs to the Younger Mastixioid Floras sensu Mai (1964) and is correlated with the early Miocene floristic assemblage ("Florenkomplex") of Eichelskopf-Wiesa (Mai 1995), i.e., the floristic zone VI in Saxony sensu Mai (1967). It includes in the foliage record representatives of ferns (Thelypteridaceae, Polypodiales fam. inc.), conifers (Pinaceae, Cupressaceae, Geinitziaceae) and angiosperms (prevailingly Lauraceae, Fagaceae, rarely Hamamelidaceae, Myricaceae, Juglandaceae, Leguminosae (including Leguminosites hradekensis (E. KNOBLOCH et KVAČEK) KVAČEK et TEODORIDIS comb. n.), Oleaceae and uncertain families). The carpological record adds, in addition to the above families, representatives of Magnoliaceae, Nymphaeaceae, Schisandraceae (incl. Illiciaceae), Menispermaceae, Cornaceae, Lythraceae (incl. Microdiptera donata (HOLÝ) KVAČEK et TEODORIDIS comb. n.), Halorhagaceae, Altingiaceae, Ulmaceae, Rosaceae, Ericaceae, Symplocaceae, Styracaceae, Rutaceae, Aquifoliaceae, Staphyleaceae, Vitaceae, Sabiaceae, Caprifoliaceae, Potamogetonaceae, Arecaceae, Cyperaceae, Stemonaceae and Sparganiaceae. An informal term "plexus" is attached to names of fossil species whose parts have not yet been found in organic connection. Vegetation is classified as broad-leaved evergreen forest type according to the new statistical method developed for zonal forest formations (IPR vegetation analysis). Palaeoenvironmental and climatic proxy data based on angiosperm leaf record using a combination of the physiognomic (CLAMP, LMA) and Nearest Living Relatives (CA) techniques are, according to CLAMP estimates - MAT 14.2 °C, WMMT 24.0 °C, CMMT 6.2 °C, 3-WET 131.7 cm, 3-DRY 20.0 cm, GROWSEAS 10.3 month, GSP 203.1 cm, MMGSP 9.9 cm, RH 54.5 %, SH 5.5 g/kg, and ENTHAL 29.9 kJ/kg; LMA estimates - MAT1 is 21.1 °C (sensu Wolfe 1979) and MAT2 is 18.7 °C (sensu Su et al. 2010), and value of the sampling error sensu Miller et al. (2006) is 3.1 °C; CA proxy data intervals: MAT 17-18 °C, WMT 26.5-26.9 °C, CMT 9.6-12.6 °C, and MAP 1146-1146 mm. The obtained climate proxy datasets correspond more or less with those of stratigraphically analogous sites from Wackersdorf, Wiesa and Berzdorf in Germany as well as the summarised proxy datasets of the Mydlovary Fm. in South Bohemia.
Fossil fruits and a vegetative axis assignable to the extant genus Ceratophyllum are described from four North American Tertiary localities. Fossil fruits assignable to the extant species C. muricatum and C. echinatum are reported from the Eocene Green River and Claiborne formations, and the Miocene Esmerelda Formation, respectively. An extinct species, C. furcatispinum, is described from the Paleocene Fort Union Formation and represents the oldest published report of Ceratophyllum in the fossil record. The existence of extant angiosperm species in the Eocene is very unusual and may be attributable in this case to slow evolutionary rates and unusual evolutionary properties associated with hydrophily in the genus Ceratophyllum.
Today Tetraclinis Mast. (Cupressaceae) and Metasequoia Miki (Taxodiaceae) are monotypic relict conifers with a successively narrowed geographical range and a reduced number of species since the Tertiary. The former genus occurred frequently in many palaeofloras from the Palaeocene to the Pliocene in Europe and still remains an endemic in S.W. Europe and N. Africa. The latter genus covered much of the northern hemisphere from the Late Cretaceous to the Pliocene but extended in to the edges of Europe only for a short time. Changes in ecological tolerances obviously brought about the range restriction to the west Mediterranean arid area (Tetraclinis) and the central Chinese humid floral province (Metasequoia). Recent relicts should be considered inaccurate indicators in the Tertiary because the ecological tolerances shown by the relict species may be very different from those of their Tertiary representatives.