Late Miocene Palaeocarya (Engelhardieae: Juglandaceae) from Southwest China and its biogeographic implications

Fossil fruits of Palaeocarya (Juglandaceae) are described from late Miocene sediments of southeastern Yunnan, China. The fruits present a tri-lobed wing consisting of an intact oblong-ovate middle lobe and two lateral lobes. The lobes are apically obovate, and have pinnate venation. The middle lobe is thicker at the base and gradually tapers to the apex. The nutlet, located at the base of the winged fruit, is round and hispid, and is subdivided by a septum into two compartments. Based on extensive morphological comparisons to previously documented fossil fruits, we found that the fossil fruits align most closely with members of the genusPalaeocarya, but have a unique combination of characters. Thus, we describe the fossils as a new species, Palaeocarya hispida sp. nov. This species represents an important range expansion for low-latitude occurrences of Palaeocarya in the late Miocene and therefore substantially improves our understanding of the biogeographic history of the genus. We propose that the wide distribution of Palaeocarya and relatively narrow distributions of close relatives, Engelhardia, Alfaropsis, and Oreomunnea, might be associated with a stepwise cooling and a major ice sheet expansion in the Antarctic and Arctic from the late middle Miocene to early Pliocene. In particular, the climatic oscillations during the Quaternary, such as the last glacial maximum, might have led to a decrease in the geographic distribution of Engelhardieae.     

Paliurus Fruits from the Oligocene of South China and Their Phytogeographic Implications

Paliurus favonii Unger is recognized and described based on fruits from the Oligocene Ningming flora of Guangxi, South China. Characteristics of the present specimens include circular winged fruits that are 10.0–11.5 mm in diameter with a central endocarp at 3.0 to 4.0 mm in diameter. The specimens fall into the morphological range of the fossil species P. favonii, which has been observed in other Cenozoic sites in the Northern Hemisphere. The present discovery represents the lowest latitude distribution of P. favonii in the world, and we are presenting the first P. favonii fossil described with detailed cuticular characteristics from China. Further, this finding demonstrates that the genus existed in the Oligocene Ningming region, South China, and provides new information for understanding the fossil history. The dispersal mode for winged fossils demonstrates that wind dispersal is well-represented in the Oligocene Ningming flora.     

A fossil fruit wing of Dipterocarpus from the middle Miocene of Fujian, China and its palaeoclimatic significance

Dipterocarpus zhengae sp. nov. is described from the middle Miocene Fotan Group of Zhangpu county, Fujian Province, Southeast China on the basis of a fruit wing. Three kinds of venation of the calyx longer lobes (enlarged sepals) occur in the winged fruits of extant Dipterocarpus. The fossil species is referred to the kind having three primary veins and within that category is most similar to extant Dipterocarpus gracilis Blume in the size of the longer lobe as well as its venation. The occurrence of the fruit wing of Dipterocarpus, together with palynological evidence, indicates unequivocally that a tropical climate and tropical rain forest occurred in Zhangpu during the middle Miocene, warmer and more humid than at the present day. The palaeobiogeography of Dipterocarpus is reviewed.     

First fossil evidence of Palaeocarya (Engelhardioideae: Juglandaceae) from India and its biogeographical implications

Even though presently indigenous to eastern Himalaya in India, no Engelhardioideae have been reported from the Cenozoic sediments of India till date. Here, we report the first Indian occurrence of a characteristic engelhardioid winged samaroid fruit having a tri-lobed wing (oblong-ovate median lobe and two lateral lobes) and a globose nut from the latest Neogene (Pliocene: Rajdanda Formation) sediments of Chotanagpur Plateau, eastern India. This is the first fossil evidence of relict family Juglandaceae from the Indian Cenozoic. We determine its taxonomic position on the basis of detailed macromorphological comparison with similar extant and fossil specimens and discuss its palaeoclimatic significance in terms of the present-day distribution of modern analogous species. We assign this Pliocene winged fruit specimen to the morphogenus Palaeocarya sect. Monocosta Manchester and describe it as a new species, namely Palaeocarya indica Hazra, Hazra M & Khan sp. nov. Palaeocarya sect. Monocosta has rich fossil records from the Cenozoic sediments of Europe, North America, and eastern Asia (China, Korea), but the modern analog, Engelhardia, is presently native only to India and neighboring Southeast Asia. We discuss the possible causes of disappearance of Engelhardia from the present-day vegetation of Chotanagpur Plateau. Its disappearance may be related to the gradual intensification of monsoonal rainfall seasonality since the Pliocene. Here, we also review in detail the biogeographic history of Palaeocarya sect. Monocosta and suggest its possible migration routes.     

INVESTIGATIONS OF ANGIOSPERMS FROM THE EOCENE OF NORTH AMERICA: JUGLANDACEOUS WINGED FRUITS

The winged fruits of Engelhardia have been reported in many early Tertiary floras in the northern Hemisphere. Recent systematic revisions of the extant genera, Engelhardia, Oreomunnea, and Alfaroa provide a helpful guide to the study of the fossil fruits. Fruit form, venation, nut septations and flower parts are features that separate the extant Asian Engelhardia from the American Oreomunnea. Four types of fossil winged fruits can be recognized in Middle Eocene sediments of southeastern North America: Engelhardia, Paraoreomunnea gen. n., Paleooreomunnea gen. n., and Pararengelhardia. Both Paleooreomunnea and Paraengelhardia represent extinct lines; no living forms have similar winged fruits. The fossil fruits of Engelhardia and Paraoreomunnea are similar to the fruits of extant Engelhardia and Oreomunnea, respectively. The fossil taxa reported in this paper were sympatric. The present allopatric distribution of Engelhardia and Oreomunnea is the result of Neogene regional extinctions. Engelhardia and Oreomunnea have probably been distinct genera since early Paleogene time.     

Two Eocene fossil fruits from the Changchang Basin of Hainan Island,China

Two types of fossil fruit, one belonging to Palaeocarya sp. (Juglandaceae) and the other to Acer cf. A. miofranchetii Hu et Chaney (Aceraceae), are found in the Eocene coal-bearing series from the Changchang Basin of Hainan Island, China. This is the first fossil record of Palaeocarya and Acer in a tropical area of China. These fossils provide evidence for an investigation of the phytogeographic history of these two genera. Since their extant relative genera are distributed mostly in northern temperate or tropical–subtropical mountainous regions, I propose that the Changchang Basin of Hainan Island was close to a mountainous region in the Eocene; the plants bearing these fruits were growing at a mid-high altitude with a relatively cool climate, and the fruits were not preserved in situ but transported to the fossil site. The characters of other associated fossil plants and palynological data also support this hypothesis.     

Two new species of Palaeoarthroteles Kovalev and Mostovski (Diptera, Rhagionidae) from the Callovian-Oxfordian (Jurassic) Daohugou biota of China

Two new species of fossil snipe flies (Rhagionidae) from the Daohugou Formation of Chifeng City in Inner Mongolia Autonomous Region (northeastern China) are described as Palaeoarthroteles jurassicus nov. sp. and P. pallidius nov. sp. They are the first record of the genus Palaeoarthroteles Kovalev and Mostovski beyond Siberia, which have implications for stratigraphic correlation of the formations in which they are found.     

Hemiptelea mikii sp. nov. (Ulmaceae), fossil fruits and woods from the pleistocene of central Japan

A new fossil species,Hemiptelea mikii, is described on the basis of fruits from the Pleistocene of central Japan. It is distinguished fromH. davidii, the only extant species of the genus restricted to Korea and China, in having shorter fruits. Fossil woods discovered from the same horizon differ from the extant one in some anatomical characteristics, and are assigned to the new fossil species. Taken together with earlier records of fossil fruit occurrence in the early and the middle Pleistocene of central Japan, the latest finding of fossil fruits and woods from the last glacial sediments (ca. 50,000 years ago) at Kita-egota site of Tokyo suggests thatH. mikii was widespread in central Japan throughout the Pleistocene and survived until as late as the Last Glacial age.     

Pliocene Albizia (Fabaceae) from Jharkhand,eastern India: reappraisal of its biogeography during the Cenozoic in Southeast Asia

Albizia, a diverse tree genus, occupies monsoonal warm humid rain forests in tropical and subtropical regions. We recovered a well-preserved compound fossil leaf and two fossil fruits of Albizia (Fabaceae) from the latest Neogene (Rajdanda Formation: Pliocene) sediments of Jharkhand of the Chotanagpur Plateau, eastern India. On the basis of the architectural features of the fossil leaf, a new species is established as A. mahuadanrensis Hazra, Hazra and Khan, n. sp., characterised by a bipinnate, compound leaf having a rachis bearing opposite, asymmetrically ovate to sub-rhomboid leaflets, pulvinus on leaflet petiolule and brochiodromous secondary veins. Based on both morphological and anatomical characters of the fossil fruits, A. palaeoprocera Hazra, Hazra and Khan, n. sp. is erected, characterised by flattened to broadly linear shaped, wingless fruits; ovate-elliptic shaped seed chambers having ellipsoidal seeds in one series; irregularly polygonal to rectangular epidermal cells with oblique end walls and randomly oriented, scattered, paracytic stomata. Analysis of Albizia fossil occurrences indicates that the legume taxon was common in Neogene forests of India and elsewhere. The present-day distribution of the closely affiliated modern species of the fossil taxa indicates a warm and humid tropical environment during the time of deposition. We also review the biogeographic history of Albizia in India and other Asian countries.     

Diversity of Menispermaceae from the Paleocene and Eocene of South China

We present here the earliest known Asian fossil records of the Menispermaceae based on fossil fruits from Paleocene and Eocene localities in South China. A new genus and species, Paleoorbicarpum parvum sp. nov., and two new species of Stephania Loureiro, S. ornamenta sp. nov. and S. geniculata sp. nov., are recognized from Paleocene deposits of the Sanshui Basin, Guangdong, and a new occurrence of the widespread Eocene species Stephania auriformis (Hollick) Han & Manchester is recognized from the Maoming Basin, Guangdong. The Paleocene Stephania specimens described here represent the earliest fossil endocarp record of the Menispermaceae in eastern Asia. This discovery shows that the moonseed family had arrived in tropical and humid South China by at least the middle Paleocene, which provides important evidence for the origin and phytogeographic history of the family.     

ComprehendingCornus: Puzzles and progress in the systematics of the dogwoods

Dogwoods evolved in two main lineages, a red-fruited line in which the inflorescences have basal bracts, and a blue-(or white-)fruited line in which the bracts are rudimentary or lacking. The 15 “red-line” species are mostly well marked, whereas “blue-line” species—numbering roughly 50 if some newer treatments be accepted—are mostly hard to tell apart. Red-line ovules are tenuinucellate, blue-line ovules crassinucellate. Dividing the red line separates cornelian cherries from the showy-bracted species; dividing the latter separates dwarf cornels from big-bracted dogwoods. Adding in the blue line there are thus four subgroups, which differ with regard to inflorescences— and with regard to iridoid glucosides, among other things. I downplay further subdivision (into opposite- and alternateleaved blue-line groups, for instance) and focus on few traits because I aim to trace the subgroups backward in the fossil record, to decide the direction of evolutionary changes, and to infer the causes for divergence. Unlike pollen, leaves, and wood, fossil dogwood fruits can often be assigned to one of the four subgroups and sometimes to a species group within a subgroup.Dunstania (based on fruits from England’s early Tertiary and here transferred toCornus’s cornelian cherry subgroup) or something close to the dunstanias gave rise to modernC. volkensii of Africa and four related species found today in California and Eurasia. Fossil fruit-stones of the blue line, like their modern counterparts, can be subglobose, compressed and asymmetric, fluted, or topped by a deep depression. Such a deep depression serves to link a fossil to the extant speciesC. alternifolia andC. controversa, but other features of the blue-line fruitstones, overlapping and less constant, cannot be used to prove a fossil’s tie to groups within the blue line. Evidence of several kinds makes the blue-line dogwoods oldest and connectsCornus to the nyssoids:Nyssa, Camptotheca, andDavidia. Commonly called Nyssaceae after 1910, when Wangerin got their traits wrong, the nyssoids are surer relatives ofCornus than any of Wangerin’s (“Das Pflanzenreich”) cornaceous genera.Mastixia, however, may be the actual sister group because it shares withCornus 1-celled 2-armed hairs that nyssoids lack.Cornus, nyssoids, and mastixioids (modernMastixia and its closest fossil allies) are here regarded as the true Cornaceae. Other genera once thought to be cornaceous are doubtfully to not at all related. As is now widely known, discordant features makeCorokia andKaliphora (for example) out of place in the Cornaceae, butAucuba andGarrya (for example) are neither easily excluded by a listing of their traits nor easily included by construction of branched diagrams. The novelty that brought about the radiation of true cornads was, I think, epigyny, followed fairly quickly by the hard endocarp and singleseeded locules. Mastixioid fruits of the late Cretaceous were dry or leathery and smaller than their successors. This suggests that early members of the family spread abiotically or by means of animals that ate whole plants. Bigger, fleshier fruits came later through interaction with evolving frugivores. The cause of early radiation withinCornus probably was interaction with insect predators and pollinators. One species of the blue line escaped competition for spring pollinators by switching to fall flowering and thereby kept some traits that other dogwoods lost. Though birds are now the main dispersers of the dogwoods, rodents likely play a minor role, and monkeys likely played a major role when compound fruits evolved. Though gynoecial reduction was the rule in dogwoods,C. (Dunstania) multilocularis’s multiple seed chambers reflect an evolutionary increase. I postulate that modernCornus’s 4-merous flower with 2-merous ovary has a strong developmental tie to pairing of the leaves and branches.     

Two new species of Electra (Bryozoa, Cheilostomata) from the Miocene of the Aquitaine Basin, France

Post-Cretaceous examples of Electridae, a primitive family of cheilostome bryozoans, are poorly represented in the fossil record, probably because of their thinly calcified zooids and preference for nearshore environments. Two new electrid species are here described from the Lower Miocene (Burdigalian) of Pontpourquey, Aquitaine, France: Electra triaurata nov. sp. and Electra aquitanica nov. sp. Both species belong to extant species groups, the E. indica and E. biscuta groups, respectively, that presently occur in the Indo-Pacific; both are the only fossil examples of these species groups. Whereas E. triaurata nov. sp. has uniserial colonies, zooids with porous gymnocysts, three flattened spines and basal windows allowing etching of the substrate to produce the trace fossil Leptichnus, E. aquitanica nov. sp. has multiserial colonies and zooids with a proximal gymnocyst bearing 2 to 5 spines.     

Exploring the Formation of a Disjunctive Pattern between Eastern Asia and North America Based on Fossil Evidence from Thuja (Cupressaceae)

Thuja, a genus of Cupressaceae comprising five extant species, presently occurs in both East Asia (3 species) and North America (2 species) and has a long fossil record from Paleocene to Pleistocene in the Northern Hemisphere. Two distinct hypotheses have been proposed to account for the origin and present distribution of this genus. Here we recognize and describe T. sutchuenensis Franch., a new fossil Thuja from the late Pliocene sediments of Zhangcun, Shanxi, North China, based on detailed comparisons with all living species and other fossil ones, integrate the global fossil records of this genus plotted in a set of paleomaps from different time intervals, which show that Thuja probably first appeared at high latitudes of North America in or before the Paleocene. This genus reached Greenland in the Paleocene, then arrived in eastern Asia in the Miocene via the land connection between East Asia and western North America. In the late Pliocene, it migrated into the interior of China. With the Quaternary cooling and drying, Thuja gradually retreated southwards to form today’s disjunctive distribution between East Asia and North America.     

An extinct juglandaceous fruit from the Upper Palaeocene of southern England

A new genus of juglandaceous winged fruit, is described from the Reading Beds (Upper Palaeocene) of southern England. It comprises one of the earlier macrofossil records of the Juglandaceae, and is the earliest from the European Tertiary. The fruit represents an extinct genus related to the extant tribe Engelhardieae, but excluded from it by its simple unlobed bract. Cladistic analysis shows Casholdia to display generalized engelhardioid fruit morphology. It lacks the tri-lobed bract diagnostic of the Engelhardieae, and predates the first occurrence of such bracts in the fossil record. Casholdia adds to the mounting evidence indicating an early Palaeogene radiation of the Juglandaceae.     

A new species of Nelumbo from South China and its palaeoecological implications

A new species of Nelumbo Adanson is described from the Eocene of the Changchang Basin, Hainan Island, South China. This is the first comprehensive report on a fossil Nelumbo species from the Eocene of the Changchang Basin, Hainan Island. The co-occurrence of fruits, tubers, rhizomes with numerous adventitious roots, leaves, and receptacles indicates that the plants grew where they are preserved. These specimens, combined with palynological assemblage and other macrofossils, indicate a tropical or subtropical humid climate in the Changchang Basin from the late early Eocene to early late Eocene.     

Ozakia,a new genus of winged fruit shared between the Miocene of Japan and western North America

A new genus is recognized based on winged fruits with a single species shared between the Miocene of southwestern Honshu, Japan, and the Miocene of Oregon and Idaho, USA. Calyces of Ozakia emryi gen. et sp. n. were formerly attributed to Heptacodium (Caprifoliaceae) and Amelanchier (Rosaceae); however, newly recovered specimens reveal additional characters that contradict these assignments. The pedicellate fruits are obovate, tapering basally and truncate apically, with about 10 longitudinal ribs, a prominent epigynous synsepalous calyx of five lobes, each with a midvein and a pair of weaker, ascending intramarginal primary veins. The single style has a capitate stigma. Ozakia is considered to represent an extinct eudicot genus, the familial affinities of which remain uncertain. The eastern Asian–western North American disjunction of Ozakia occurrences suggests that this plant traversed the Beringia land bridge during or prior to the Middle Miocene. Relatively few extinct angiosperm genera are known as late as the Miocene.     

The earliest fossil record of Panorpidae (Mecoptera) from the Middle Jurassic of China

The early history of Panorpidae (Mecoptera) is poorly known due to sparse fossil records. Up to date, only nine fossil species have been described, all from the Paleogene, except the Early Cretaceous Solusipanorpa gibbidorsa Lin, 1980. However, we suggest S. gibbidorsa is too incompletely preserved to permit even family classification. A new genus with two new species, Jurassipanorpa impunctata gen. et sp. n. and Jurassipanorpa sticta sp. n., are described based on four well-preserved specimens from the late Middle Jurassic Jiulongshan Formation of Daohugou, Inner Mongolia, China. These two new species are the earliest fossil records of Panorpidae. The new genus is erected based on a combination of forewing characters: both R₁ and Rs₁ with two branches, 1A reaching posterior margin of wing distad of the forking of Rs from R_1, and no crossveins or only one crossvein between veins of 1A and 2A. In all four specimens, long and robust setae ranging from 0.09 to 0.38 mm in length and pointing anteriorly, are present on anal veins of forewings. The function of these setae is enigmatic.     

First record of fossil angiosperm wood (Ulmoxylon,Ulmaceae) from the famous locality of Bı́lina (Czech Republic,Early Miocene)

A fossil angiosperm wood is described for the first time from the famous Early Miocene locality of Bı́lina. It represents a fossil elm wood, attributed to Ulmoxylon marchesonii Biondi. The fossil wood can be compared to extant North American soft elms, also to Ulmus macrocarpa Hance and U. parvifolia Jacq. from China or to the European common elm U. carpinifolia Gled. The wood together with fossil leaves/fruits of Ulmus pyramidalis Goeppert forms a single natural fossil species that lived in the Bı́lina area during the Early Miocene. The influence of two types of preservation, permineralised and xylitic, on the same wood species is also discussed. To cite this article: J. Sakala, C. R. Palevol 1 (2002) 161–166.     

Fruits of Tetrapterys (Malpighiaceae) from the Oligocene of Hungary and Slovenia

Propeller-like winged fruits of Tetrapterys harpyiarum Unger from the Oligocene of Sotzka, Budapest, Eger-Vécsey valley, and a new occurrence at Eger-Kiseged, were reinvestigated and compared in detail with extant species of Tetrapterys (Malpighiaceae) and with other dicotyledonous genera with four winged fruits. T. harpyiarum fruits are bilaterally symmetrical, consisting of a globose nut surrounded by four elongate wings with parallel venation. Tetrapterys is now distributed only in tropical America and this implies that there was an opportunity for Tetrapterys to spread between the Partethys region and the New World during the Tertiary.