Fossil Araceae from a Paleocene neotropical rainforest in Colombia

Both the fossil record and molecular data support a long evolutionary history for the Araceae. Although the family is diverse in tropical America today, most araceous fossils, however, have been recorded from middle and high latitudes. Here, we report fossil leaves of Araceae from the middle-late Paleocene of northern Colombia, and review fossil araceous pollen grains from the same interval. Two of the fossil leaf species are placed in the new fossil morphogenus Petrocardium Herrera, Jaramillo, Dilcher, Wing et Gomez-N gen. nov.; these fossils are very similar in leaf morphology to extant Anthurium; however, their relationship to the genus is still unresolved. A third fossil leaf type from Cerrejón is recognized as a species of the extant genus Montrichardia, the first fossil record for this genus. These fossils inhabited a coastal rainforest ～60-58 million years ago with broadly similar habitat preferences to modern Araceae.     

INVESTIGATIONS OF ANGIOSPERMS FROM THE EOCENE OF SOUTHEASTERN NORTH AMERICA: PHILODENDRON LEAF REMAINS

Large leaves, new to the fossil record, from the Claiborne Formation in western Tennessee have been collected and analyzed. Careful analyses of venation and cuticular anatomy indicate that these fossil leaves contain specific characters found in modern species of the genus Philodendron subgenus Meconostigma. Features of venation were taxonomically more useful than other features. Reports of fossils of the Araceae are scarce and those few fossils assigned to the family frequently have been inadequately studied. This report establishes a reliable and carefully documented occurrence of the Araceae in the fossil record, and provides information concerning the differentiation of the family in time and the distributions of the genus Philodendron.     

Sequences,stratigraphy and scenarios: what can we say about the fossil record of the earliest tetrapods?

Past research on the emergence of digit-bearing tetrapods has led to the widely accepted premise that this important evolutionary event occurred during the Late Devonian. The discovery of convincing digit-bearing tetrapod trackways of early Middle Devonian age in Poland has upset this orthodoxy, indicating that current scenarios which link the timing of the origin of digited tetrapods to specific events in Earth history are likely to be in error. Inspired by this find, we examine the fossil record of early digit-bearing tetrapods and their closest fish-like relatives from a statistical standpoint. We find that the Polish trackways force a substantial reconsideration of the nature of the early tetrapod record when only body fossils are considered. However, the effect is less drastic (and often not statistically significant) when other reliably dated trackways that were previously considered anachronistic are taken into account. Using two approaches, we find that 95 per cent credible and confidence intervals for the origin of digit-bearing tetrapods extend into the Early Devonian and beyond, spanning late Emsian to mid Ludlow. For biologically realistic diversity models, estimated genus-level preservation rates for Devonian digited tetrapods and their relatives range from 0.025 to 0.073 per lineage-million years, an order of magnitude lower than species-level rates for groups typically considered to have dense records. Available fossils of early digited tetrapods and their immediate relatives are adequate for documenting large-scale patterns of character acquisition associated with the origin of terrestriality, but low preservation rates coupled with clear geographical and stratigraphic sampling biases caution against building scenarios for the origin of digits and terrestrialization tied to the provenance of particular specimens or faunas.     

The fossil pollen record of Araceae

The fossil record of Araceae pollen beginning in the late Early Cretaceous and peaking in the Paleocene/Eocene is very sparse up to now, consisting of three highly distinctive types: zona-aperturate pollen of the Monstera or Gonatopus type (very similar to Proxapertites operculatus), an ulcerate-spiny type typical for Limnobiophyllum, and a polyplicate, omniaperturate pollen type (an ephedroid pollen with non-gnetalean affinities) which was recently reported from the late Early Cretaceous (Mayoa portugallica). An extensive literature search has shown that some distinctive Ephedripites forms (the Paleogene Ephedripites vanegensis, and the Late Cretaceous Ephedripites elsikii) are very similar to pollen of Spathiphyllum and both species are here transferred from Ephedripites to Spathiphyllum (as comb. nov.). We also add new fossil findings to the Araceae record. The new findings include a zona-aperturate, microperforate to microreticulate pollen type from the Palaeocene of Colombia, highly similar to extant Gonatopus or Zamioculcas or Monstera pollen (Araceae) and to fossil Proxapertites operculatus, which is currently seen as a fossil equivalent; and, an ulcerate, spiny pollen from the Eocene of Stolzenbach, Germany, extending the range of Limnobiophyllum (Pandaniidites), which is thought to be an extinct member of extant Araceae. The three pollen types add considerably to the reliable fossil record of the family that now contains more than 20 records of these three pollen types: with the zona-aperturate type recorded from the tropical or subtropical regions of Northern and Southern America, Central Africa, Southern and Central Europe, from the Indian subcontinent and the Malayan Archipelago; the ulcerate type occurring in North America and Europe; and the polyplicate type mainly occurring in South America and South-West Europe. Now we have good evidence that some of the aroid subfamilies were already in existence in the Cretaceous, increasing in diversity and worldwide distribution in the Paleogene. Dedicated to Prof. Dr. Stefan Vogel on the occasion of his 80th birthday.     

辽西义县下白垩统沙海组木化石新材料

辽西地区下白垩统沙海组是我国北方地区重要的白垩纪木化石产出层位,其产出的木化石数量丰富、类型多样.尽管前人开展了部分研究,但目前对沙海组木化石植物群组成特征的认识仍相对薄弱.本文报道了辽西义县上石洞沟地区沙海组产出的木化石新材料,共鉴定出2属3种,分别为:Phyllocladoxylon cf.eboracense(H...     

On the systematic position of the Early Cretaceous fern genus “Athyrium”

The oldest fossils assigned to Athyrium (mostly based on the sorus morphology) comprise fronds and spores from the Lower Cretaceous of Northeast Asia. However, most molecular dating suggests that extant Athyrium diverged from its sister genus during the Eocene or later, implying that the Cretaceous fossils probably belong to another polypodiaceous taxon. By examining the sorus morphology of extant genera related to the family Athyriaceae, we found that the primary diagnostic feature for assigning the Cretaceous fossils to Athyrium, i.e., the sorus shape, is common to the entire extant family, or plesiomorphic for the genus. As the fronds are more commonly preserved than the reproductive parts, we compared the fossil frond morphology with those of living taxa of the family that is divided into two types. The Cretaceous fossil we examined here bears the frond’s costal groove characters on adaxial side, which is more closely related to that of the Deparia-clade instead of the clade including Athyrium and other genera of the family. The observation is further confirmed by the cladistic analysis using morphological characters. The systematic position of the Early Cretaceous “Athyrium” was resolved as a stem member of the total Athyriaceae using a tip-dating approach with the Fossilized Birth-Death model in a Bayesian framework. Our study suggests that Early Cretaceous fossils previously assigned to Athyrium require taxonomic revision.     

吉林乾安大布苏晚更新世野牛化石

我国东北地区更新世野牛化石的发现已有近一个世纪历史, 尽管先后有不少零星报道, 但至今无一篇专门文献, 并且前人报道的头骨化石材料无一是完整的。本文记述了保存基本完好的2件野牛头骨和3件下颌骨; 化石来自吉林省乾安县大布苏地区, 地质时代为距今约2万年。基于其形态特征和测量数据, 本文将其归入草原野牛Bison priscus(Bojanus, 1827)。前人将东北平原晚更新世野牛归入Bison exiguus Matsumoto, 1915的做法存在诸多问题, 因为后者在模式产地、地质时代及形态特征等方面都与东北晚更新世的野牛差异甚大。我国东北平原晚更新世的野牛应归入草原野牛种, 该种是中-晚更新世广泛分布于全北区的种类, 也是猛犸象-披毛犀动物群的最主要成员之一。     

Biogeography of cedrela (meliaceae, sapindales) in central and South america

Dated phylogenies have helped clarify the complex history of many plant families that today are restricted to the world's tropical forests, but that have Eocene, Oligocene, and Miocene fossils from the northern hemisphere. One such family is the Meliaceae. Here we infer the history of the neotropical Meliaceae genus Cedrela (17 species), the sister clade of which today is restricted to tropical Asia. Sequences from the nuclear ribosomal spacer region and five plastid loci obtained for all ingroup species and relevant outgroups were used to infer species relationships and for molecular-clock dating under two Bayesian relaxed clock models. The clock models differed in their handling of rate autocorrelation and sets of fossil constraints. Results suggest that (1) crown group diversification in Cedrela started in the Oligocene/Early Miocene and intensified in the Late Miocene and Early Pliocene, and (2) Central American Cedrela species do not form a clade, implying reentry into Central America after the closure of the Panamanian Isthmus. At present, Cedrela is distributed in both dry and humid habitats, but morphological features suggest an origin in dry forest under seasonal climates, fitting with Miocene and Pliocene Cedrela fossils from deciduous forests.     

贵州盘县中三叠统混鱼龙类化石的初步研究

记述贵州省盘县新民乡羊圈村中三叠统关岭组上段的混鱼龙3种,其中有一新种:Mixosaurus mao-taiensis Young,1965,emend.nov.,Mixosaurus cornalianus(Bassani),1886和Mixosaurus yangjuanensis sp.nov.。与混鱼龙类化石同层产出的有真颌鱼类和龙鱼类化石,真颌鱼类的特征比意大利和瑞士边界地区安尼-拉丁期和拉丁期早期普罗山头组(Prosanto Formation)所产的始真颌鱼Eoeugnathus进步,混鱼龙新种也比意-瑞边界地区安尼-拉丁期过渡期的混鱼龙进步。因此,我们推测当前混鱼龙类化石的层位可能包括从下安尼阶上部到拉丁阶下部的一段地层,不是简单的一个化石层,此问题尚待进一步深入研究。     

REVIEWS

Avian remains from the Early Miocene (～17 Ma) Moghra Formation of Egypt include new records of ‘waterbirds’ (storks, herons, pelicans and allies) and a ratite. Only a single avian fossil has been previously reported from Wadi Moghra and, thus, additional knowledge of the avifauna complements previously documented faunal and floral assemblages and provides a more complete picture of the Early Miocene ecosystem of that locality. The presence of ciconiiforms and pelecaniforms is consistent with the relatively abundant records of ‘waterbirds’ from other African freshwater fossil localities. Moreover, avian fossils from Wadi Moghra provide new insight into the avifaunal composition of the Early Miocene in Africa, a period that is relatively underrepresented in the avian fossil record of that continent.     

Maximum likelihood inference implies a high, not a low, ancestral haploid chromosome number in Araceae, with a critique of the bias introduced by 'x'

Background and Aims

For 84 years, botanists have relied on calculating the highest common factor for series of haploid chromosome numbers to arrive at a so-called basic number, x. This was done without consistent (reproducible) reference to species relationships and frequencies of different numbers in a clade. Likelihood models that treat polyploidy, chromosome fusion and fission as events with particular probabilities now allow reconstruction of ancestral chromosome numbers in an explicit framework. We have used a modelling approach to reconstruct chromosome number change in the large monocot family Araceae and to test earlier hypotheses about basic numbers in the family.

Methods

Using a maximum likelihood approach and chromosome counts for 26 % of the 3300 species of Araceae and representative numbers for each of the other 13 families of Alismatales, polyploidization events and single chromosome changes were inferred on a genus-level phylogenetic tree for 113 of the 117 genera of Araceae.

Key Results

The previously inferred basic numbers x = 14 and x = 7 are rejected. Instead, maximum likelihood optimization revealed an ancestral haploid chromosome number of n = 16, Bayesian inference of n = 18. Chromosome fusion (loss) is the predominant inferred event, whereas polyploidization events occurred less frequently and mainly towards the tips of the tree.

Conclusions

The bias towards low basic numbers (x) introduced by the algebraic approach to inferring chromosome number changes, prevalent among botanists, may have contributed to an unrealistic picture of ancestral chromosome numbers in many plant clades. The availability of robust quantitative methods for reconstructing ancestral chromosome numbers on molecular phylogenetic trees (with or without branch length information), with confidence statistics, makes the calculation of x an obsolete approach, at least when applied to large clades.     

The quality of the fossil record of Mesozoic birds

The Mesozoic fossil record has proved critical for understanding the early evolution and subsequent radiation of birds. Little is known, however, about its relative completeness: just how 'good' is the fossil record of birds from the Mesozoic? This question has come to prominence recently in the debate over differences in estimated dates of origin of major clades of birds from molecular and palaeontological data. Using a dataset comprising all known fossil taxa, we present analyses that go some way towards answering this question. Whereas avian diversity remains poorly represented in the Mesozoic, many relatively complete bird specimens have been discovered. New taxa have been added to the phylogenetic tree of basal birds, but its overall shape remains constant, suggesting that the broad outlines of early avian evolution are consistently represented: no stage in the Mesozoic is characterized by an overabundance of scrappy fossils compared with more complete specimens. Examples of Neornithes (modern orders) are known from later stages in the Cretaceous, but their fossils are rarer and scrappier than those of basal bird groups, which we suggest is a biological, rather than a geological, signal.     

PhyloSort: a user-friendly phylogenetic sorting tool and its application to estimating the cyanobacterial contribution to the nuclear genome of Chlamydomonas

Background

Molecular phylogenetic analyses have identified Trimeniaceae, a monotypic family distributed only in Oceania, as among the earliest diverging families of extant angiosperms. Therefore, the fossils of this family are helpful to understand the earliest flowering plants. Paleobotanical information is also important to track the historical and geographical pathways to endemism of the Trimeniaceae. However, fossils of the family were previously unknown from the Early Cretaceous, the time when the angiosperm radiated. In this study, we report a seed from the late Albian (ca. 100 million years ago) of Japan representing the oldest known occurrence of Trimeniaceae and discuss the character evolution and biogeography of this family.

Results

A structurally preserved seed was collected from the early Late Albian Hikagenosawa Formation of the Yezo Group, which was deposited in palaeolatitudes of 35 to 40°N. The seed has a multilayered stony exotesta with alveolate surface, parenchymatous mesotesta, and operculate inner integument, which are characteristic to extant trimeniaceous seeds. However, the seed differs from extant seeds, i.e., in its well-developed endosperm and absence of antiraphal vascular bundle. Thus, the seed would be a new genus and species of Trimeniaceae.

Conclusion

The fossil seed indicates that seed coat characters were conserved for 100 million years or more in Trimeniaceae. It also suggests that the antiraphal vascular bundle and perispermy originated secondarily in Trimeniaceae as previously inferred from the phylogeny and character distribution in the extant basalmost angiosperms. The fossil seed provides the first evidence that Trimeniaceae was distributed in a midlatitude location of the Northern Hemisphere during the Early Cretaceous, when angiosperms radiated extensively, supporting a hypothesis that the extant austral distribution is relict.     

木兰科的化石记录

通过整理和分析木兰科植物的化石记录发现：不论是植物大化石还是花粉,迄今为止在白垩纪以前地层中尚无可靠的记录,自白垩纪以来,木兰科的许多种广泛发生于北半球,如亚洲,欧洲及北美等地,但非洲和大洋洲至今尚未发现木兰科的化石记录。该科最早的化石记录为中国东北延吉地区早白垩世大拉子组的喙柱始木兰Archimagnolia rostrato-stylose Tao et Zhang. 根据现有化石记录,并结合木兰科现代植物的地理分布,推测：1）木兰科的起源时间不迟于早白垩世Aptian-Albian期;2）木兰科起源地点可能是东亚,后来经过欧洲进入北美,再从北美迁移到达南美洲;3）在地质历史时期,木兰属的出现比鹅掌楸属早,从而支持根据形态学与分子系统学研究得出的木兰属较鹅掌楸属原始的结论。     

Amber microfossils: On the validity of species concept

Do terrestrial micro-organisms evolve morphologically? A recent concept suggests that morphological stasis over dozens of millions of years has persisted in microbial lineages. However, it is based on a weak fossil record. Indeed, it is already difficult to define a species with extant microbes, and this task is even harder when dealing with fossil micro-organisms. Based on research on fossils in amber, we highlighted the different problems that are raised when describing a new fossil species of micro-organisms and we discuss the concept of morphological stasis.     

New fossil leaves and fruits of Lauraceae from the Middle Miocene of Fujian,southeastern China differentiated using a cluster analysis

The fossil record of Lauraceae can be traced back to the Early Cretaceous of eastern Asia based on fossil flowers. Here, we refer a number of new occurrences of leaf and fruit fossils of Lauraceae from the Middle Miocene of Zhangpu, Fujian, China, to seven species. These data provide evidence supporting the fact that a diverse subtropical, or tropical, Lauraceae-dominated evergreen forest surrounded this region 15 million years ago (Mya). The Lauraceae fossils presented in this paper provide evidence for the evolution of this group as well as new materials that enable the study of the Fujian Province Neogene flora. The fossils described in this paper fill in the gaps in studies about Lauraceae pollen in the Middle Miocene from Fotan, Fujian, China. In addition, these fossils also enrich the Middle Miocene fossil records of Lauraceae in eastern Asia, especially improving the study of the macrostructures and reproductive organs of fossil Lauraceae from southern China. The similarity between fossil and modern fruits shows that during the Middle Miocene the fruit morphological of Lauraceae have changed very little. We also identify families where the fossils we report belong to their closest relatives and can be used to reconstruct the paleoenvironment of Fujian in the Middle Miocene.     

Phylogenetic relationships of aroids and duckweeds (Araceae) inferred from coding and noncoding plastid DNA

Familial, subfamilial, and tribal monophyly and relationships of aroids and duckweeds were assessed by parsimony and Bayesian phylogenetic analyses of five regions of coding (rbcL, matK) and noncoding plastid DNA (partial trnK intron, trnL intron, trnL-trnF spacer) for exemplars of nearly all aroid and duckweed genera. Our analyses confirm the position of Lemna and its allies (formerly Lemnaceae) within Araceae as the well-supported sister group of all aroids except Gymnostachydoideae and Orontioideae. The last two subfamilies form the sister clade of the rest of the family. Monophyly of subfamilies Orontioideae, Pothoideae, Monsteroideae, and Lasioideae is supported, but Aroideae are paraphyletic if Calla is maintained in its own subfamily (Calloideae). Our results suggest expansion of the recently proposed subfamily Zamioculcadoideae (Zamioculcas, Gonatopus) to include Stylochaeton and identify problems in the current delimitation of tribes Anadendreae, Heteropsideae, and Monstereae (Monsteroideae), Caladieae/Zomicarpeae, and Colocasieae (Aroideae). Canalization of traits of the spathe and spadix considered typical of Araceae evolved after the split of Gymnostachydoideae, Orontioideae, and Lemnoideae. An association with aquatic habitats is a plesiomorphic attribute in Araceae, occurring in the helophytic Orontioideae and free-floating Lemnoideae, but evolving independently in various derived aroid lineages including free-floating Pistia (Aroideae).     

Upper Cretaceous sulcate pollen from the Timerdyakh Formation,Vilui Basin (Siberia)

One inaperturate and 16 monosulcate pollen types are described from the latest Campanian to earliest Maastrichtian sediments of the Vilui basin, Siberia, using both light and scanning electron microscopy, and assigned systematically when possible to modern families or subfamilies. Despite their scant occurrence, the pollen grains show considerable diversity. Two new genera have been erected. Lasioideaecidites with two species, Lasioideaecidites hessei and Lasioideaecidites bogneri, represents the earliest record of the subfamily Lasioideae (Araceae). Aristolochiacidites with one species, Aristolochiacidites viluiensis, is assigned to the subfamily Aristolochioideae (Aristolochiaceae) and represents the first fossil pollen record of the family. A new species of Liliacidites, Liliacidites goldblattii, is closely similar to pollen of Isophysioideae (Iridaceae) and/or Doryanthaceae and is the first fossil evidence of the clade that includes these closely related families. A further nine new fossil species are described (Clavatipollenites timerdyakhensis, Liliacidites palaeofritillaria, Retimonocolpites longosucatus, R. microreticulatus, R. microrugulatus, R. lysichitonoides, R. chapmaniae, Monosulcites parvus and Arecipites tyungensis) and assigned to Chloranthaceae, Araceae (Orontioideae, ?Pothoideae), Liliaceae, ?Hypoxidaceae and Arecaceae. The Hydatellaceae (Nymphaeales) may also be present in the palynoflora of the Vilui Basin, represented by pollen similar to M. rivularis Braman from the Santonian to Campanian (Upper Cretaceous) Milk River Formation, Canada.