Rainforest conifers of Eocene Patagonia: Attached cones and foliage of the extant Southeast Asian and Australasian genus Dacrycarpus (Podocarpaceae)

? Premise of the study: Eocene caldera-lake beds at Laguna del Hunco (LH, ca. 52.2 Ma) and Río Pichileufú (RP, ca. 47.7 Ma) in Argentine Patagonia provide copious information about the biological history of Gondwana. Several plant genera from these sites are known as fossils from southern Australia and New Zealand and survive only in Australasian rainforests. The potential presence of Dacrycarpus (Podocarpaceae) holds considerable interest due to its extensive foliage-fossil record in Gondwana, its remarkably broad modern distribution in Southeast Asian and Australasian rainforests, its high physiological moisture requirements, and its bird-dispersed seeds. However, the unique seed cones that firmly diagnose Dacrycarpus were not previously known from the fossil record. ? Methods: I describe and interpret fertile (LH) and vegetative (LH and RP) material of Dacrycarpus and present a nomenclatural revision for fossil Dacrycarpus from South America. ? Key results: Dacrycarpus puertae sp. nov. is the first fossil occurrence of the unusual seed cones that typify living Dacrycarpus, attached to characteristic foliage, and of attached Dacrycarpus pollen cones and foliage. Dacrycarpus puertae is indistinguishable from living D. imbricatus (montane, Burma to Fiji). Dacrycarpus chilensis (Engelhardt) comb. nov. is proposed for Eocene vegetative material from Chile. ? Conclusions: Modern-aspect Dacrycarpus was present in Eocene Patagonia, demonstrating an astonishingly wide-ranging paleogeographic history and implying a long evolutionary association with bird dispersers. Dacrycarpus puertae provides the first significant Asian link for Eocene Patagonian floras, strengthens the biogeographic connections from Patagonia to Australasia across Antarctica during the warm Eocene, and indicates high-rainfall paleoenvironments.     

Cross-species amplification of Eucalyptus microsatellite loci

This study examined the interspecific amplification of nuclear microsatellite loci developed mainly for eucalypts in the subgenus Symphyomyrtus across five species within the second most speciose subgenus, subgenus Eucalyptus. A set of eight to 10 loci, depending on taxon, have been identified that are highly variable and easily scored. The successful transfer of microsatellite loci to these eucalypt species sidesteps the expensive and time-consuming development of species-specific microsatellite libraries. This primer set will enable the examination and cross-species comparison of the genetic resources of commercially and ecologically important members of the subgenus Eucalyptus.     

CAESALPINIA SUBGENUS MEZONEURON (LEGUMINOSAE,CAESALPINIOIDEAE) FROM THE TERTIARY OF NORTH AMERICA

Legume fruits from the Eocene of Tennessee and Wyoming and the Miocene of Idaho are described and assigned to Caesalpinia subgenus Mezoneuron (Caesalpinioideae), an extant Paleotropical taxon that does not occur in North or South America today. Morphological and anatomical details of the fruits are used in evaluating their systematic relationships. The features of the fossil fruits are accommodated only within this extant subgenus. These fossils represent the only reliable known occurrence of C. subgenus Mezoneuron in the paleobotanical record. These fossils suggest that subgenus Mezoneuron was distinct from subgenus Caesalpinia by the Middle Eocene. Further, they document the widespread occurrence of this currently Paleotropical group for at least 30 million years in North America.     

The first Gondwanan Euphorbiaceae fossils reset the biogeographic history of the Macaranga-Mallotus clade

Premise

The spurge family Euphorbiaceae is prominent in tropical rainforests worldwide, particularly in Asia. There is little consensus on the biogeographic origins of the family or its principal lineages. No confirmed spurge macrofossils have come from Gondwana.

Methods

We describe the first Gondwanan macrofossils of Euphorbiaceae, represented by two infructescences and associated peltate leaves from the early Eocene (52 Myr ago [Ma]) Laguna del Hunco site in Chubut, Argentina.

Results

The infructescences are panicles bearing tiny, pedicellate, spineless capsular fruits with two locules, two axile lenticular seeds, and two unbranched, plumose stigmas. The fossils' character combination only occurs today in some species of the Macaranga-Mallotus clade (MMC; Euphorbiaceae), a widespread Old-World understory group often thought to have tropical Asian origins. The associated leaves are consistent with extant Macaranga.

Conclusions

The new fossils are the oldest known for the MMC, demonstrating its Gondwanan history and marking its divergence by at least 52 Ma. This discovery makes an Asian origin of the MMC unlikely because immense oceanic distances separated Asia and South America 52 Ma. The only other MMC reproductive fossils so far known are also from the southern hemisphere (early Miocene, southern New Zealand), far from the Asian tropics. The MMC, along with many other Gondwanan survivors, most likely entered Asia during the Neogene Sahul-Sunda collision. Our discovery adds to a substantial series of well-dated, well-preserved fossils from one undersampled region, Patagonia, that have changed our understanding of plant biogeographic history.     

石松类卷柏科地史时期和现代地理分布格局的形成

卷柏科仅含一属,全球分布,具有宽幅生态适应性,属于维管植物基部类群石松类植物,现生约700种。卷柏科起源于泥盆纪,化石记录遍布自石炭纪至今的各地质时期。目前,现代卷柏属亚属的分类方案基本稳定,但对于化石卷柏的亚属分类、以及地质历史时期的时空分布问题还缺乏相关研究。本文全面总结了卷柏科化石记录的时代和地理分布,并结合分子系统学探讨其地理分布格局的演化,为该科植物的分类与进化研究提供参考。根据现有研究基础,结合最新的卷柏科系统分类研究成果和化石记录,本文对该类群化石的系统分类进行归纳和厘定,并对化石类群的卷柏亚属Selaginella、Hexaphyllum亚属及rhizophoric clade物种的分化和地理分布格局的形成进行了讨论。本文认为,卷柏科起源于古生代的欧美植物区,不晚于二叠纪,其后伴随着泛大陆解体扩散至华夏植物区及冈瓦纳植物区。化石亚属Hexaphyllum是rhizophoric clade的基部类群,在石炭纪末灭绝。化石证据表明,从古生代末期起,rhizophoric clade已成为卷柏科的优势类群,并在中生代形成了全球分布的格局。     

Papuacedrus (Cupressaceae) in Eocene Patagonia: A new fossil link to Australasian rainforests

The 51.9 Ma Laguna del Hunco (LH) and 47.5 Ma Río Pichileufú (RP) floras from Patagonia, Argentina are unusually rich, angiosperm-dominated assemblages with living relatives in the low-latitude West Pacific, neotropics, and temperate southern latitudes. The diverse gymnosperms in these floras are important for Gondwanan biogeographic history and paleoclimatic interpretations. "Libocedrus" prechilensis Berry 1938 (Cupressaceae), previously known only from the holotype (RP), a vegetative branch, is revised here based on new material from both localities, including a seed cone attached to a shoot with cuticle (LH). Characters of these fossils are diagnostic of monotypic Papuacedrus (highlands of New Guinea and Moluccas). Living P. papuana is most abundant in cloud forests receiving up to 4 m rainfall annually, whereas Austrocedrus (Libocedrus) chilensis, the basis of comparison when the fossil species was named, inhabits dry, cold steppe margins to mediterranean climates in southern South America. We establish Papuacedrus prechilensis comb. nov., which simultaneously invalidates a southern South American connection for the fossil floras and reveals a link to West Pacific montane rainforests. Combined evidence indicates a biome similar to extant subtropical, or tropical montane, rainforests that persisted for at least 4.4 Myr, linking elevated floral richness to abundant rainfall.     

Eocene plant diversity at Laguna del Hunco and Río Pichileufú, Patagonia, Argentina

The origins of South America's exceptional plant diversity are poorly known from the fossil record. We report on unbiased quantitative collections of fossil floras from Laguna del Hunco (LH) and Río Pichileufú (RP) in Patagonia, Argentina. These sites represent a frost-free humid biome in South American middle latitudes of the globally warm Eocene. At LH, from 4,303 identified specimens, we recognize 186 species of plant organs and 152 species of leaves. Adjusted for sample size, the LH flora is more diverse than comparable Eocene floras known from other continents. The RP flora shares several taxa with LH and appears to be as rich, although sampling is preliminary. The two floras were previously considered coeval. However, (40)Ar/(39)Ar dating of three ash-fall tuff beds in close stratigraphic association with the RP flora indicates an age of 47.46+/-0.05 Ma, 4.5 million years younger than LH, for which one tuff is reanalyzed here as 51.91+/-0.22 Ma. Thus, diverse floral associations in Patagonia evolved by the Eocene, possibly in response to global warming, and were persistent and areally extensive. This suggests extraordinary richness at low latitudes via the latitudinal diversity gradient, corroborated by published palynological data from the Eocene of Colombia.     

Revision of the Proteaceae macrofossil record from Patagonia, Argentina

Proteaceae are restricted to the Southern Hemisphere, and of the seven tribes of the subfamily Grevilleoideae, only three (Macadamieae, Oriteae, and Embothrieae) have living members in Argentina. Megafossil genera of Proteaceae recorded from Patagonia includeLomatia, Embothrium, Orites, andRoupala. In this report, we evaluate and revise fossil Argentine Proteaceae on the basis of type material and new specimens. The new collections come from the Tufolitas Laguna del Hunco (early Eocene, Chubut Province), the Ventana (middle Eocene, Río Negro Province), and the Río Ñirihuau (late Oligocene-early Miocene, Río Negro Province) formations, Patagonia, Argentina. We confirm the presence ofLomatia preferruginea Berry,L. occidentalis (Berry) Frenguelli,L. patagonica Frenguelli,Roupala patagonica Durango de Cabrera et Romero, andOrites bivascularis Romero, Dibbern et Gandolfo. Fossils assigned toEmbothrium precoccineum Berry andE. pregrandiflorum Berry are doubtful, and new material is necessary to confirm the presence of this genus in the fossil record of Patagonia. A putative new fossil species of Proteaceae is presented as Proteaceae gen. et sp. indet. Fossil Proteaceae are compared with modern genera, and an identification key for the fossil leaf species is presented. Doubtful historical records of Proteaceae fossils for the Antarctic Peninsula region and Patagonia are also discussed. Based on this revision, the three tribes of Proteaceae found today in Argentina were already present in Patagonia by the early Eocene, where they probably arrived via the Australia-Antarctica-South America connection.     

Contrasting adaptation of two Eucalyptus subgenera is related to differences in respiratory metabolism

1. The largest eucalypt subgenus Symphyomyrtus survives and grows better in diverse native and exotic environments than the second largest subgenus Monocalyptus . Previously postulated reasons for this difference, including differential resistance to native insect pests and dependence on specific ectomycorrhizal symbiotic associations with soil fungi, do not fully explain the marked adaptive differentiation.
2. This work shows that differences between the subgenera in survival and growth performance are related to respiratory parameters. We propose that effects of climatic temperature on respiratory metabolism result in Symphyomyrtus being more successful than Monocalyptus in adapting to diverse environments. This suggests a new paradigm for adaptation and evolution of eucalypts.     

Reproductive structures of Rhamnaceae from the Cerro del Pueblo (Late Cretaceous, Coahuila) and Coatzingo (Oligocene, Puebla) Formations, Mexico

Recently discovered fossil flowers from the Cretaceous Cerro del Pueblo and flowers and fruits from the Oligocene Coatzingo Formations are assigned to the Rhamnaceae. The Cretaceous flower, Coahuilanthus belindae Calvillo-Canadell and Cevallos-Ferriz, gen. et sp. nov., is actinomorphic with fused perianth parts forming a slightly campanulate to cupulate floral cup, with sepals slightly keeled and spatulate clawed petals. The Oligocene fossils include Nahinda axamilpensis Calvillo-Canadell and Cevallos-Ferriz, gen. et sp. nov. (characterized by its campanulate bisexual flower with stamens opposite, adnate to and enfolded by petals; and with the ovary ripening into a drupe), and a winged fruit assigned to Ventilago engoto Calvillo-Canadell and Cevallos-Ferriz, sp. nov. The flowers and drupe features indicate closer affinity to Zizipheae and/or Rhamneae, while the single samaroid fruit suggests the presence of Ventilagineae. However, the unique character combination in the fossil flowers precludes placing them in extant genera. Nevertheless, the history of the family is long and can be traced back to the Campanian. A detailed phylogenetic revision of the group that uses morphological characters from both extant and fossil plants is needed to better understand the significance of these records as well as other important fossils of the family.     

Chloroplast sharing in the Tasmanian eucalypts

The biogeographic pattern of chloroplast DNA (cpDNA) haplotypes in Eucalyptus on the island of Tasmania is consistent with reticulate evolution, involving at least 12 Tasmanian species from the subgenus Symphyomyrtus. Intraspecific cpDNA polymorphism in 14 of 17 species is coupled with extensive sharing of identical haplotypes across populations of different species in the same geographic area. Haplotype diversity is lowest in central regions of Tasmania formerly occupied by alpine vegetation during glacial intervals and in northern regions that were periodically linked to continental Australia by land bridges. The observed distribution of several cpDNA haplotypes unique to Tasmania coincides with modeled locations of glacial refugia in coastal areas of Tasmania and shows the power of cpDNA in unraveling the complex history of past distributions of Eucalyptus. The results suggest that the model of evolution of the eucalypts should be reassessed to allow for the anastomosing effects of interspecific hybridization and introgression.     

Phytogeographic implications of fossil endocarps of Menispermaceae from the Paleocene of Colombia

? Premise of the study: Fossil leaves of Menispermaceae were previously described from the Paleocene of Colombia. Because of strong homoplasy of leaf characters, the fossils could not be placed more specifically within recognized clades, and additional data were needed to specify intrafamilial and paleogeographic relationships during the Paleocene. ? Methods: Fossil endocarps of Menispermaceae were collected from the Cerrejón Formation, the recently discovered Bogotá flora, and Wyoming (～60 Ma). We surveyed the endocarp morphology of almost all extant genera, conducted character optimization, a molecular scaffold analysis, and critically reviewed the related fossil genera. ? Key results: Parallel syndromes of fruit characters have appeared in unrelated clades of the family according to current phylogenetic reconstructions. However, mapping selected endocarp characters across those clades that contain horseshoe-shaped endocarps facilitates identification and phylogenetic assessment of the fossils. Three fossil species are recognized. One of them belongs to the extant genus Stephania, which today grows only in Africa and Australasia. Palaeoluna gen. nov. is placed within the pantropical clade composed of extant Stephania, Cissampelos, and Cyclea; this morphogenus is also recognized from the Paleocene of Wyoming. Menispina gen. nov. shows similarity with several unrelated clades. ? Conclusions: The new fossils from Colombia reveal a complex paleobiogeographic history of the recognized clades within Menispermaceae, suggesting a more active exchange among neotropical, paleotropical, North American, and European paleoforests than previously recognized. In addition, the new fossils indicate that neotropical forests were an important biome for the radiation and dispersal of derived lineages in Menispermaceae after the Cretaceous-Paleogene boundary.     

INVESTIGATIONS OF TERTIARY ANGIOSPERMS: A NEW FLORA INCLUDING EOMIMOSOIDEA PLUMOSA FROM THE OLIGOCENE OF EASTERN TEXAS

A new fossil flower and inflorescence-bearing locality has been discovered in the Oligocene of the Texas Gulf Coast. The new flora is similar to the Middle Eocene Claiborne Flora of the southeastern USA, but the quality of preservation is sometimes better in the Oligocene fossils. One component of the new flora, a mimosoid legume inflorescence, appears identical with Eomimosoidea plumosa, first reported from the Claiborne Formation of western Tennessee. Investigations of these younger specimens indicate that the taxon had changed little during the Middle Eocene-Oligocene interval, and the better quality of preservation of the Texas specimens has provided further insights into the structure of the fossils. Comparisons of the fine structural details of the pollen of Eomimosoidea with similar pollen of extant mimosoids has confirmed that the fossil genus is indeed extinct and suggests that tetrahedral tetrads of columellate, tricolporate pollen grains are ancient, possibly primitive, in the Mimosoideae.     

Seed morphology of modern and fossil Ampelocissus (Vitaceae) and implications for phytogeography

Seeds are useful in distinguishing among extant genera of Vitaceae and provide a good basis for interpretation of fossil remains in reconstructing the evolutionary and phytogeographic history of this putatively basal Rosid family. Seeds of Ampelocissus s.l. including Pterisanthes and Nothocissus are distinguished from those of all other vitaceous genera by long, parallel ventral infolds and a centrally positioned oval chalazal scar. Principal component analysis facilitates recognition of four Ampelocissus s.l. seed morphotypes differentiated by dorsiventral thickness, width of ventral infolds, chalazal depth, and degree of dorsal surface rugosity. While these intergrade, their end-member morphologies are distinctive and coincide well with inflorescence morphology, extant geographic distribution, and ecology. Seven fossil morphospecies are recognized. Ampelocissus parvisemina sp. n. (Paleocene of North Dakota; Eocene of Oregon) and A. auriforma Manchester (Eocene of Oregon) resemble extant Central American species; A. bravoi Berry (Eocene of Peru) is similar to one group of Old World extant species; and A. parachandleri sp. n. (Eocene of Oregon) and the three European fossil species A. chandleri (Kirchheimer) comb. n., A. lobatum (Chandler) comb. n., and A. wildei sp. n. (Eocene to Miocene) resemble another group of extant Old World Ampelocissus. All these fossils occur outside the present geographic range of the genus, reflecting warmer climates and former intercontinental links.     

A new genus and species of darner dragonfl y (Aeshnidae: Odonata) from the lower Eocene of Laguna del Hunco,Patagonia, Argentina

A new genus of Aeshnidae, Huncoaeshna n. gen., based on Huncoaeshna corrugata n. gen., n. sp., is erected from Laguna del Hunco (Ypresian) in Patagonia Argentina. The specimen presents a special kind of preservation with the middle part of the wing wrinkled. The presence of only two fossil specimens of Aeshnidae in South America is surely due to the lack of paleontomologists and collections of fossil insects in the subcontinent.     

A cladistic study of Arillastrum, Angophora and Eucalyptus (Myrtaceae)

A cladistic study of Anllastrum, Angophora and Eucalyptus (Myrtaceae). Transformed cladistic; character compatibility; branch and bound, and Farris-Wagner methods gave similar solutions in a cladistic study of Arillastrum, Angophora and Eucalyptus. These analyses, based on morphological characters, indicate that Eucalyptus is a monophyletic group and that its sister taxon is Angophora.
Within Eucalyptus , subgenera Blakella and Corymbia are sister taxa to all other groups; subgenera Monocalyptus, Idiogenes and Gaubaea form a monophyletic group with subgenus Monocalyptus sister to subgenera Idiogenes and Gaubaea ; subgenera Symphyomyrtus and Telocalyptus together also form a monophyletic group and, with Eucalyptus similis (subgenus Eudesmia group 4), are sister to the Monocalyptus group. Eucalyptus subgenus Telocalyptus (4 species), Eucalyptus subgenus Idiogenes (1 species) and Eucalyptus subgenus Gaubaea (2 species) should not be recognized as subgenera and some individual species need further examination. Eucalyptus subgenus Eudesmia is a paraphyletic group.
Some characters are identified as parallelisms, e.g. axillary inflorescences, sepaline operculum, bristle glands, and clustered anthers. A more congruent interpretation of the single operculum of Eucalyptus subgenus Monocalyptus as at least partly petaline rather than solely sepaline in origin is suggested.
The area relationships for the taxa are concordant with those derived from geological and climatological information. New Caledonia is sister area to Australia, and within Australia southwestern Australia is sister area to south-eastern and north-eastern Australia.     

Fossil Ericaceae from New Zealand: Deconstructing the use of fossil evidence in historical biogeography

The Australasian Ericaceae epitomize many problems in understanding the biogeography of the southern hemisphere, especially the relative contributions of Gondwanan vicariance and dispersal. Late Cretaceous fossil pollen of the family suggests extreme antiquity of the group in Australasia, but recent phylogenetic evidence suggests much younger histories for most of the groups in that region. This paper documents two new species of latest Oligocene-Early Miocene macrofossils of Ericaceae from New Zealand. Cyathodophyllum novae-zelandiae G.J.Jord. & Bannister gen. and sp. nov. is the oldest record of the tribe Styphelieae, but is of a clade now extinct in New Zealand, possibly related to the Tasmanian genus Cyathodes. Richeaphyllum waimumuensis G.J.Jord. & Bannister sp. nov. is a member of Richeeae, but it is ambiguous as to whether it is a member of the impressive modern New Zealand radiation in Dracophyllum. These fossils emphasize the fact that at least some of the fossil pollen of Ericaceae may have been derived from extinct lineages and therefore should not be used as evidence for the antiquity of any modern New Zealand clade of Ericaceae. New fossils and/or detailed analysis of fossil and extant pollen may help resolve such uncertainty.     

Phylogeny of extant and fossil Juglandaceae inferred from the integration of molecular and morphological data sets

It is widely acknowledged that integrating fossils into data sets of extant taxa is imperative for proper placement of fossils, resolution of relationships, and a better understanding of character evolution. The importance of this process has been further magnified because of the crucial role of fossils in dating divergence times. Outstanding issues remain, including appropriate methods to place fossils in phylogenetic trees, the importance of molecules versus morphology in these analyses, as well as the impact of potentially large amounts of missing data for fossil taxa. In this study we used the angiosperm clade Juglandaceae as a model for investigating methods of integrating fossils into a phylogenetic framework of extant taxa. The clade has a rich fossil record relative to low extant diversity, as well as a robust molecular phylogeny and morphological database for extant taxa. After combining fossil organ genera into composite and terminal taxa, our objectives were to (1) compare multiple methods for the integration of the fossils and extant taxa (including total evidence, molecular scaffolds, and molecular matrix representation with parsimony [MRP]); (2) explore the impact of missing data (incomplete taxa and characters) and the evidence for placing fossils on the topology; (3) simulate the phylogenetic effect of missing data by creating "artificial fossils"; and (4) place fossils and compare the impact of single and multiple fossil constraints in estimating the age of clades. Despite large and variable amounts of missing data, each of the methods provided reasonable placement of both fossils and simulated "artificial fossils" in the phylogeny previously inferred only from extant taxa. Our results clearly show that the amount of missing data in any given taxon is not by itself an operational guideline for excluding fossils from analysis. Three fossil taxa (Cruciptera simsonii, Paleoplatycarya wingii, and Platycarya americana) were placed within crown clades containing living taxa for which relationships previously had been suggested based on morphology, whereas Polyptera manningii, a mosaic taxon with equivocal affinities, was placed firmly as sister to two modern crown clades. The position of Paleooreomunnea stoneana was ambiguous with total evidence but conclusive with DNA scaffolds and MRP. There was less disturbance of relationships among extant taxa using a total evidence approach, and the DNA scaffold approach did not provide improved resolution or internal support for clades compared to total evidence, whereas weighted MRP retained comparable levels of support but lost crown clade resolution. Multiple internal minimum age constraints generally provided reasonable age estimates, but the use of single constraints provided by extinct genera tended to underestimate clade ages.