A systematic overview of fossil osmundalean ferns in China: Diversity variation,distribution pattern,and evolutionary implications

The order Osmundales is a unique fern taxon with extensive fossil records in geological past. Diverse osmundalean fossils have been reported from China, ranging in age from the Late Palaeozoic to the Cenozoic. Most of them are based on leaf impressions/compressions, but permineralized rhizomes are also well documented. In this study, we provide a systematic overview on fossil osmundalean ferns in China with special references on diversity variations, distribution patterns, and evolutionary implications. Fossil evidence indicates that this fern lineage first appeared in the Late Palaeozoic in China. The Late Triassic to Middle Jurassic interval was the radiation stage. From the Late Jurassic onward, fossil diversity declined rapidly. Cenozoic osmundalean taxa are represented by the relict species of Osmunda. Geographically, osmundalean fossils are found from both the Northern and Southern phytoprovinces of China, though variations are documented for geographical ranges. The Chinese fossil records cover almost all important stages for the macroevolution of the Osmundales, and contribute to further understanding of evolutionary processes of this peculiar fern lineage.     

Carnivora from the Plio-Pleistocene hominin site of Drimolen, Gauteng, South Africa

Drimolen is one of the newest and most productive hominin sites in South Africa, and is dated on faunal grounds between 2.0 Ma to 1.5 Ma. This paper provides the first overview of the Carnivora from Drimolen, updating the previously published preliminary faunal list, and describing all currently prepared craniodental and postcranial material. The Drimolen specimens are described in comparison with other modern and fossil South African carnivore material. The carnivores cover a range of taxa including hyaenids, felids, canids and herpestids. Most notable amongst these are the sabretooth Dinofelis aff. piveteaui craniodental and postcranial remains, which are described in detail, and a Chasmaporthetes nitidula cranium. The genus Chasmaporthetes is found at three other sites in the area - Sterkfontein, Swartkrans and Coopers D. There are two models for the geographic origin of Dinofelis piveteaui, in that it may have arisen in either eastern or southern Africa. These possibilities are discussed in the light of the new South African Dinofelis material, as the Drimolen material appears to represent a more primitive form with affinities with D. piveteaui. Fossil leopard material from Kromdraai B and Drimolen is also discussed, as the metapodia assigned to P. pardus from these two sites are very small, but lie within the variation of modern leopards. Such size differences in fossil postcrania may have implications for the niches that these animals may have occupied in the past.     

Fertile organs and in situ spores of a new dipteridaceous fern Hausmannia sinensis from the Jurassic of northern China

As a representative fossil member of the dipteridaceous fern, genus Hausmannia was reported worldwide from the Mesozoic strata; however, little is known about the fertile structures, including sporangia and in situ spores, of this genus. In this study, a new species Hausmannia sinensis was identified from the Middle Jurassic of Nei Mongol (Inner Mongolia), northern China. The specimens are compressions and are well preserved with details of sporangia and in situ spores. The leaf laminae are broadly fan-shaped, with an almost entire margin. Primary and lateral veins dichotomously branch to form square or polygonal meshes. Each ultimate mesh bears one to two circular sori of 0.4 mm in diameter. Sori are exindusiate; each sorus contains three to six round to ovoid sporangia. The annulus is developed and oblique, with stomial region present in proximal position. Spores are trilete, circular to oval in shape. Both proximal and distal surfaces are covered with baculate to subverrucate sculptures. Spores range from 20 to 30 μm in diameter (average 28 μm), and are comparable to the dispersed genera Baculatisporites Thomas and Pflug and Apiculatisporis Potonié and Kremp. Hausmannia sinensis represents the first compression species of genus Hausmannia form Eurasia, which shows the combination of well-preserved sori, sporangia, annuli and in situ spore characters, and is therefore helpful for further understanding the diversity and evolution of the Dipteridaceae fern lineage through time.     

A new fossil record of Thyrsopteridaceae (Cyatheales) from the mid-Cretaceous amber of Myanmar

Paleopalynology evidence and megafossil records indicated that tree fern order Cyatheales, played an important role in Mesozoic terrestrial ecosystem. Few fossil records of the family Thyrsopteridaceae have been reported so far. In the present study, we describe a distinctive fertile pinnule segment of a fern plant preserved in a mid-Cretaceous amber from Myanmar. The well-preserved fertile pinnule is identified as a new species of the extant tree fern genus Thyrsopteris (Thyrsopteridaceae): Thyrsopteris cyathindusia n. sp. It represents the second fossil species of Thyrsopteridaceae found in the mid-Cretaceous amber from Myanmar. The new species shows distinctive characteristics of the sporophyll that are considered paleoecologically/paleobiogeographically significant for evolution of the Thyrsopteridaceae.     

Early Cretaceous Gleicheniaceae and Matoniaceae (Gleicheniales) from Alexander Island,Antarctica

The first Cretaceous representatives of Matoniaceae and Gleicheniaceae in the Antarctic are described from the Albian flora of Alexander Island. Two new species are recognized, one is assigned to Matonia, and another to the emended gleicheniaceous form-genus Gleicheniaceaephyllum. The taxonomy of fossil genera assigned to Gleicheniaceae is problematic, and as an alternative to the invalid genus Gleichenites, we propose that Microphyllopteris be utilised as a form-genus for species that have either an unknown branching pattern or pseudo-dichotomous branching without resting buds; Microphyllopteris is reserved for ferns that cannot be ascribed conclusively to Gleicheniaceae. Alternatively, Gleicheniaceaephyllum should be used for gleicheniaceous material that has a resting bud amongst two to four primary branches. Gleicheniaceaephyllum acutum sp. nov. is preserved as sterile and fertile fronds, pinnae and pinnules. The other new species, Matonia jeffersonii sp. nov., occurs as sterile and fertile pinnae and pinnules, and is aligned to Matoniaceae based on the presence of circular-oval sori with peltate indusia. The two taxa are inferred to have either a pedate or scrambling habit. These morphologies are relatively common at Alexander Island, compared to other high latitude fossil floras, and are interpreted as an adaptation to the high disturbance, polar conditions and unusual photoperiod experienced at Alexander Island during the Early Cretaceous. The theory that angiosperms caused the demise of ferns is discussed in relation to the Matoniaceae and Gleicheniaceae, and there does not appear to be association between angiosperm expansion and the decline of these two fern families.     

Remarks on some Eurasian pliocervines: Characteristics, evolution, and relationships with the tribe Cervini

In order to define the ancestors of the Eurasian genera Axis and Rusa, the representatives of the tribe Pliocervini have been reviewed, with the exclusion of the genus Pliocervus. The fossil remains referred to Pliocervini stored in the Palaeontological Museum of Mechnikov Odessa National University and studied by Khomenko [Khomenko, I., 1913. Meotian fauna of v. Tarakliya of Bendery district. Annuaire géologique et minéralogique de la Russie. 1. The ancestors of modern and fossil Cervinae. 2. Giraffidae and Cavicornia 15, 107-132 (in Russian)] are reappraised: these remains are attributed to Cervavitus novorossiae (Khomenko, 1913). The fossil deer collected from the locality of Novoelizavetovka (Odessa region, Ukraine) and observed in the same Museum of the Odessa National University, is referred to Cervavitus variabilis Alexeev (1915).The fossil material from Shanxi and Honan provinces (China), described by Zdansky [Zdansky, O., 1925. Fossil Hirsche Chinas. Palaeontologia Sinica C 2, 1-94] and Teilhard de Chardin and Trassaert [Teilhard de Chardin, P., Trassaert, M., 1937. The Pliocene Camelidae, Giraffidae and Cervidae of southeastern Shansi.] is referred to Cervavitus shanxius Dong and Hu, 1994. The phylogenetic relationships between Pliocervini from China and the European ones are investigated on the basis of the morphological, biochronological and paleogeographical data. These elements and the suggestions of the more recent genetical and biometrical data may indicate that the origin of the two genera Axis and Rusa could be recognized into the Chinese pliocervines.     

Davallia (Polypodiales: Davalliaceae) macrofossils from Early Miocene Otago (New Zealand) with in situ spores

Fossil fern fronds referable to the extant fern genus Davallia (Polypodiales: Davalliaceae) bearing sporangia with in situ spores are described from the Early Miocene Foulden Maar diatomite deposit, Otago, New Zealand. The fronds are the first published Southern Hemisphere macrofossil record for the family and provide valuable palaeoclimate data supporting warm conditions in Early Miocene New Zealand. The matching of Davallia fronds to the form spore taxon Polypodiisporites radiatus shows that the genus has had a long, apparently continuous history throughout late Cenozoic New Zealand.     

Palaeobiogeographic and metric analysis of the Mesozoic fern Weichselia

Eighty-three publications have been examined to recover information on the palaeogeography, the chronostratigraphy and the presence of the fossil fern genus Weichselia Stiehler in different depositional environments. This fossil fern foliage was reported in 72 localities worldwide. Its fossil record ranges from the Bathonian to the Cenomanian, predominating from the Berriasian to the Barremian, but still common during the Aptian to Cenomanian. Weichselia mostly occurred in continental depositional environments in Europe, whereas it has been mostly related to marginal marine depositional environments in Africa, North America, and India. Three species have been described: W. reticulata, W. peruviana, and W. negevensis. We analyse the measurements on specimens collected from sixteen localities as well as published data from twelve others. Our results suggest that: (i) the climate changes during the Aptian might have affected the pinnule size of Weichselia, as the Aptian-Cenomanian pinnules are larger than those from the Berriasian-Barremian; (ii) some differences might be explained by the metric variation of the frond parts; (iii) there are no clear differences in size between remains collected from different depositional environments; (iv) there is no sufficient evidence to determine how many species of Weichselia have existed.     

An analysis of two Myosorex species (Soricidae) from the Early Pliocene site of Langebaanweg (West coast, South Africa) using geometric morphometrics, linear measurements, and non-metric characters

This represents the first study of the soricid community from the Early Pliocene site of Langebaanweg (LBW) which is represented by four species, two of which belong to the soricid genus Myosorex (Forest shrew). Using geometric morphometrics the two Myosorex fossil species are compared with each other, and the extant western and eastern Cape species: Myosorex varius, M. cafer and M. longicaudatus. In addition, the results from an analysis of a number of non-metric characters and linear measurements on the fossil species are presented. The mandibles of the fossil species are characterized by extremely broad teeth, ascending rami which are relatively gracile and differ in shape and size from modern species, and the existence of a number of primitive features of the mandibles and teeth. This evidence is interpreted as indicating that the two LBW Myosorex species represent an archaic lineage now extinct in southern Africa.     

Filicales y Bennettitales del Cretácico Inferior de Carazuelo, provincia de Soria, España

Silicified woods of the fern Tempskya riojana Barale and Viera and the Bennettitales Cycadeoidea barbarae Crisafulli nov. sp., found together with undetermined conifer woods, from the NW of Carazuelo, province Soria, are described. This site is located south of the so-called San Saturio fault which runs parallel to Road N-234. The fossil trunks are included in the “Conglomerados poligénicos, areniscas y arcillas” fluviatile sediments, dated as Paleogene. The original source of these fossils could be Tera and/or Oncala Groups, both Early Cretaceous in age and outcropping close by at Fuentetecha, Fuensaúco and North of Soria. Similar plant fossil findings from this region are reported and some mistakes in the geological mapping are analyzed. The Tempskya specimens were histologically poorly preserved, but had enough anatomical characters to be undoubtedly assigned to Tempskya riojana. The new species Cycadeoidea barbarae is described, with fairly well preserved cellular structure, viz., pith, vascular cylinder, cortex, foliar/branch traces, as well as strobili surrounded by foliar scales and apical seminal cones. An important and new aspect of the new species is that it includes morphologically and anatomically preserved both fructifications and vegetative parts, whereas these are usually find separated by other species of Bennettitales.     

The identity of Tetracyclus constrictus (Bacillariophyceae) and a description of Tetracyclusbradburyii, a new miocene species

The siliceous valves from species of Tetracyclus (Bacillariophyceae) have variable but characteristic shapes. In this paper, species characterised by having valves with a central constriction are discussed. This distinctive shape occurs during only a period of its lifecycle. Tetracyclus constrictus and Tetracyclus celatom have been applied to fossil specimens occurring in deposits from the west coast of the USA and the east coast of Russia. Do these names apply to two closely related species or are they one and the same taxon? Two further species, Tetracyclus javanicus and Tetracyclus peragalli, have been involved in the identity of T. constrictus–T. celatom. T. constrictus M. Peragallo is the correct name and author for specimens of the extinct Miocene, circum-Pacific (or northern) fossil species with valves panduriform with rounded poles; T. javanicus refers to a species found living on Java only; fossil specimens usually referred to T. javanicus (Java) belong to a new taxon, Tetracyclus bradburyii (USA); T. peragalli is an extinct Miocene fossil, with a northern hemisphere distribution; small (≤10 μm) circular/elliptical fossil specimens, found alongside other specimens, are difficult (if not impossible) to determine to species level.     

Dammaradiene synthase, a squalene cyclase, from Dryopteris crassirhizoma Nakai

Ferns produce a variety of cyclic triterpene hydrocarbons in large amount. Squalene cyclases (SCs) are responsible enzymes for formation of cyclic triterpene hydrocarbon skeletons. Although more than ten bacterial SCs have been cloned and four of them characterized for their enzymatic products, the only example of a fern SC is ACH, from Adiantum capillus-veneris, which produces hydroxyhopane. To obtain a deeper understanding of the molecular evolution of SCs and the origin of the structural diversity of fern triterpenes, further cloning and characterization of SCs have been pursued. In this study, a SC cDNA, DCD, was cloned from Dryopteris crassirhizoma by homology-based RT-PCR. DCD contains a 2058-bp open reading frame that encodes a 685 amino acid polypeptide exhibiting 66% identity to the previously identified fern SC, ACH, and 35-40% identity to bacterial SCs. Heterologous expression of DCD in yeast established it to be a dammaradiene synthase affording dammara-18(28),21-diene, a tetracyclic triterpene hydrocarbon. Although neither this compound nor any derived metabolites have been previously reported from D. crassirhizoma, re-investigation of the leaflets demonstrated the presence of dammara-18(28),21-diene. DCD represents the first SC that produces a tetracyclic triterpene hydrocarbon.     

A new large barn owl (Aves, Strigiformes, Tytonidae) from the Middle Pleistocene of Sicily, Italy, and its taphonomical significance

A new species of Tytonidae, Tyto mourerchauvireae, is described from the Sicilian cave deposits of Spinagallo, Luparello and Marasà, which have yielded a common vertebrate fossil assemblages referred to the early Middle Pleistocene. T. mourerchauvireae nov. sp. shows a pronounced increase in body size compared to other congeneric taxa. It is larger than the extant Tyto alba and the extinct Tyto balearica and Tyto sanctialbani and comparable in size among the Mediterranean taxa only with the extinct Tyto robusta, which differs in some morphological characteristics of the long bones. The insular adaptations of T. mourerchauvireae nov. sp. and the differences from the other forms of the genus Tyto which spread through the Mediterranean area during Neogene and Pleistocene are discussed. The fossil assemblages of the early Middle Pleistocene of Sicily are dominated by extinct giant Gliridae of the genus Leithia and Maltamys; these taxa are regarded as the primary prey items of T. mourerchauvireae nov. sp. The Gliridae remains were analysed microscopically to detect possible modifications of bones and teeth caused by ingestion and digestion processes. This type of analysis, together with the qualitative study of the fossil assemblages, allows to show the taphonomical importance of T. mourerchauvireae nov. sp. as agent of accumulation in such Sicilian fossil assemblages. Incidentally the taxonomic validity of T. robusta is discussed and confirmed.     

Phylogeny of early Australopithecus: new fossil evidence from the Woranso-Mille (central Afar,Ethiopia)

The earliest evidence of Australopithecus goes back to ca 4.2 Ma with the first recorded appearance of Australopithecus ‘anamensis’ at Kanapoi, Kenya. Australopithecus afarensis is well documented between 3.6 and 3.0 Ma mainly from deposits at Laetoli (Tanzania) and Hadar (Ethiopia). The phylogenetic relationship of these two ‘species’ is hypothesized as ancestor–descendant. However, the lack of fossil evidence from the time between 3.6 and 3.9 Ma has been one of its weakest points. Recent fieldwork in the Woranso-Mille study area in the Afar region of Ethiopia has yielded fossil hominids dated between 3.6 and 3.8 Ma. These new fossils play a significant role in testing the proposed relationship between Au. anamensis and Au. afarensis. The Woranso-Mille hominids (3.6–3.8 Ma) show a mosaic of primitive, predominantly Au. anamensis-like, and some derived (Au. afarensis-like) dentognathic features. Furthermore, they show that, as currently known, there are no discrete and functionally significant anatomical differences between Au. anamensis and Au. afarensis. Based on the currently available evidence, it appears that there is no compelling evidence to falsify the hypothesis of ‘chronospecies pair’ or ancestor–descendant relationship between Au. anamensis and Au. afarensis. Most importantly, however, the temporally and morphologically intermediate Woranso-Mille hominids indicate that the species names Au. afarensis and Au. anamensis do not refer to two real species, but rather to earlier and later representatives of a single phyletically evolving lineage. However, if retaining these two names is necessary for communication purposes, the Woranso-Mille hominids are best referred to as Au. anamensis based on new dentognathic evidence.     

Starch grain morphology of the seagrasses Halodule wrightii, Ruppia maritima, Syringodium filiforme, and Thalassia testudinum

Starch grains are a ubiquitous component of plants that have been used in tandem with phytoliths, pollen, and macrofossils to reconstruct past floral diversity. This tool has yet to be fully explored for aquatic plants, specifically seagrasses, which lack phytoliths and are rarely preserved as macrofossils or pollen. If starch grains in seagrasses are morphologically distinct, this method has the potential to improve seagrass identification in the fossil record in such cases where its starch is preserved (e.g. scratches and occlusal surfaces of tooth enamel from seagrass consumers). The goals of this study were twofold: (1) to determine if starch is present in seagrass material and (2) to assess how starch grain morphology differs between different seagrasses.This study focused on four abundant and ecologically distinct seagrasses from the Caribbean: Halodule wrightii, Ruppia maritima, Syringodium filiforme, and Thalassia testudinum. Starch grains were observed in all species except S. filiforme. Grains from H. wrightii are typically observed in side-on orientation, are sub-round to angular, and are fairly small (3-19 μm, end-on). Grains of R. maritima are small spherical grains (4-8 μm) that have a centric hilum and a straight extinction cross with a median angle between the arms of 90°. Grains from T. testudinum are large (9-31 μm, end-on), conical in side-on and round/sub-round in end-on orientation, have a slightly eccentric hilum with an obvious particle, and prominent lamellae.Visual assessment and comparative statistics demonstrate that the morphology of starch grains from T. testudinum, R. maritima, and H. wrightii are significantly different. With more extensive research, there is potential for the positive identification of starch grains from an unknown seagrass. The ability to identify seagrass from starch grains could facilitate the identification of seagrasses in the fossil record and supply information on seagrass evolution and distribution, climate effects on seagrass distribution, and the diets of seagrass consumers.     

First discovery of colobine fossils from the early to middle Pleistocene of southern Taiwan

Here we report on two kinds of cercopithecid fossil monkeys (Cercopithecinae and Colobinae) from the early to middle Pleistocene sediments of the Chochen (=Tsochen) area (Tsailiao-chi or Shinhua Hill), southern Taiwan. The fossil specimens include the first fossil record of colobine monkeys from Taiwan, where only macaque monkeys now occur. All cercopithecine fossils were identified as Macaca cf. Macaca cyclopis, the extant Taiwan macaque, except for one extremely large isolated upper molar, which may belong to another macaque species. On the other hand, all colobine specimens fall within the size variation of extant and extinct Rhinopithecus, but its specific status cannot be determined because of the scantiness of the fossil material. In Taiwan, Rhinopithecus presumably became extinct in the late Pleistocene, probably owing to global cooling and vegetation change, whereas macaques, which are of almost the same body size as Rhinopithecus, survived as M. cyclopis to the present. The contrasting history of survival between the two kinds of monkeys may be due to ecological/behavioral differences between them or as a result of accidental events that occurred in the Pleistocene of Taiwan.     

A closer look at the “Protopithecus” fossil assemblages: new genus and species from Bahia,Brazil

The recently extinct large-bodied New World monkey Protopithecus brasiliensis Lund 1836 was named based on a distal humerus and proximal femur found in the Lagoa Santa cave system in the southeastern Brazilian state of Minas Gerais. These bones are from an animal about twice the size of the largest extant platyrrhines. One hundred and seventy-five years later, a nearly complete skeleton was discovered in the Toca da Boa Vista caves in the neighboring state of Bahia and was allocated to the same taxon as it was the first platyrrhine fossil of comparable size found since the originals. Our detailed study of the equivalent elements, however, reveals important morphological differences that do not correspond to intraspecific variation as we know it in related platyrrhine taxa. The presence of both an expanded brachioradialis flange on the humerus and gluteal tuberosity on the femur of the Bahian skeleton distinguishes it from the Lagoa Santa fossil as well as from all other platyrrhines. Further cranial and postcranial evidence suggests a closer relationship of the former with the alouattine Alouatta, while the limited Lund material fits more comfortably with the ateline clade. Therefore, we propose to limit P. brasiliensis Lund to the distal humerus and proximal femur from Lagoa Santa and erect a new genus and species for the skeleton from Toca da Boa Vista. Cartelles coimbrafilhoi was a large-bodied frugivore with a relatively small brain and diverse locomotor repertoire including both suspension and climbing that expands the range of platyrrhine biodiversity beyond the dimensions of the living neotropical primates.     

Comparative pollen morphology of Brachylena, Tarchananthus and two species of Tubulifloridites (Asteraceae) from the Eocene, Knysna Lignite of South Africa

Two fossil taxa Tubulifloridites antipodica and T. viteauensis recovered from the Eocene Knysna Lignite of South Africa were examined with scanning (SEM) and transmission electron microscopy (TEM). The details of their sculpturing and wall structure are similar to the same species of fossil dispersed pollen taxa recovered from southwestern Africa and South America. Fifteen species of the woody South African taxa, Brachylaena (9 species) and Tarchonanthus (6 species) were investigated with SEM and TEM. All of the taxa are tricolporate, spherical to slightly prolate, microechinate to echinate and have a bilayered columellate infrastructure, except B.ilicifolia, which has a single columellate infrastructural level with the “granularization” of the outer portion of the infrastructural layer or the inner layer of the tectum. There is a similar distribution of plesiomorphic and derived pollen characters in a number of aster subfamilies and tribes suggesting a similar evolutionary progression of pollen, and pollen wall character evolution was occurring synchronously in a variety of aster subfamilies during the middle Tertiary and that these unique pollen features may be important to the evolution and diversification of the Asteraceae.