Late paleozoic and Late Triassic limestones from north Palawan Block (Philippines): Microfacies and paleogeographical implications

Summary Selected Late Paleozoic and Triassic limestone exposures were studied on northern Palawan Island, Philippines, with regard to microfacies, stratigraphy and facies interpretation. Although some of the outcrops were already reported in literature, we present the first detailed microfacies study. Late Paleozoic carbonates in the El Nido area are represented by widley distributed Permian and locally very restricted Carbonifenous limestones. Of particular interest is the first report of Carboniferous limestones in the Philippines dated by fossils. Fusulinids indicate a ‘Middle’ Carboniferous (Moscovian-Kasimovian) age of the Paglugaban Formation only known from Paglugaban Island. The Permian Minilog Formation consists mostly of fusulinid wackestones and dasycladacean wacke-/packstones. Fusulinid datings (neoschwagerinids and verbeekinids) provide a Guadalupian (Wordian-Capitanian) age. The depositional setting of the Middle Permian carbonates corresponds to a distally steepened ramp with biostromes built by alatoconchid bivalves locally associated with richthofeniid brachiopods. Late Triassic limestones occur in isolated exposures on and around Busuanga Island (Calamian Islands). The age of the investigated carbonates is Rhaetian based on the occurrence ofTriasina hantkent Maizon. Microfacies data indicate the existence of reefs (Malajon Island) and carbonate platforms (Kalampisanan Islands, Busuanga Island, Coron Island). Reef boundstones are characterized by abundant solenoporacean red algae, coralline sponges and corals. Platform carbonates yield a broad spectrum of microfacies types, predominantly wacke- and packstones with abundant involutinid foraminifera and some calcareous algae. These facies types correspond to platform carbonates known from other parts of Southeast Asia (Eastern Sulawesi and Banda Basin; Malay Peninsula and Malay Basin). The Philippine platform carbonates were deposited on and around seamounts surrounded by deeper water radiolarian cherts. The new data on facies and age of the Philippine Permian and Triassic carbonates contradict a close paleogeographical connection between the North Palawan Block and South China and arise problems for the currently proposed origin of the North Palawan Block at the paleomargin of South China. We hypothesize that North Palawan was part of the Indochina Block during the Carboniferous and Permian, separated from the Indochina Block during the Middle Permian and collided with the South China Block in the Late Cretaceous.     

Out of Africa and the evolution of human behavior

Twenty‐one years ago, a landmark exploration of mitochondrial DNA diversity popularized the idea of a recent African origin for all living humans. 1 The ancestral African population was estimated to have existed 200 ka (thousands of years ago) plus or minus a few tens of thousands of years. A corollary was that at some later date the fully modern African descendants of that population expanded to swamp or replace the Neanderthals and other nonmodern Eurasians. The basic concept soon became known as “Out of Africa,” after the Academy Award winning film (1985) that took its title, in turn, from Isak Dinesen's classic autobiography (1937). Many subsequent genetic analyses, including those of Ingman and coworkers 2 and Underhill and coworkers, 3 have reaffirmed the fundamental Out of Africa model. The fossil and archeological records also support it strongly. The fossil record implies that anatomically modern or near‐modern humans were present in Africa by 150 ka; the fossil and archeological records together indicate that modern Africans expanded to Eurasia beginning about 50 ka.     

Island life in the Cretaceous - faunal composition,biogeography, evolution,and extinction of land-living vertebrates on the Late Cretaceous European archipelago

The Late Cretaceous was a time of tremendous global change, as the final stages of the Age of Dinosaurs were shaped by climate and sea level fluctuations and witness to marked paleogeographic and faunal changes, before the end-Cretaceous bolide impact. The terrestrial fossil record of Late Cretaceous Europe is becoming increasingly better understood, based largely on intensive fieldwork over the past two decades, promising new insights into latest Cretaceous faunal evolution. We review the terrestrial Late Cretaceous record from Europe and discuss its importance for understanding the paleogeography, ecology, evolution, and extinction of land-dwelling vertebrates. We review the major Late Cretaceous faunas from Austria, Hungary, France, Spain, Portugal, and Romania, as well as more fragmentary records from elsewhere in Europe. We discuss the paleogeographic background and history of assembly of these faunas, and argue that they are comprised of an endemic ‘core’ supplemented with various immigration waves. These faunas lived on an island archipelago, and we describe how this insular setting led to ecological peculiarities such as low diversity, a preponderance of primitive taxa, and marked changes in morphology (particularly body size dwarfing). We conclude by discussing the importance of the European record in understanding the end-Cretaceous extinction and show that there is no clear evidence that dinosaurs or other groups were undergoing long-term declines in Europe prior to the bolide impact.     

Speculations on carbon dioxide starvation, Late Tertiary evolution of stomatal regulation and floristic modernization

Ambient atmospheric CO₂ concentration ([CO₂]_a) has apparently declined from values above 200μmol mol⁻¹ to values below 200μmol mol⁻¹ within the last several million years. The lower end of this range is marginal for C₃ plants. I hypothesize that: (1) declining [CO₂]_a imposed a physiological strain on plants, and plant taxa evolving under declining [CO₂]_a tended to develop compensating mechanisms, including increased stomatal efficiency; (2) angiosperms were better able to adjust to declining [CO₂]_a than were gymnosperms and pteridophytes; and (3) angiosperm adjustment has been uneven. Fast-evolving taxa (e.g. grasses and herbs) have been better able to adapt to CO₂ starvation. If these propositions are true, stomatal adjustment mechanisms should show patterned variation, and a single pattern of stomatal regulation cannot be assumed.     

The evolution of the phenylpropanoid pathway entailed pronounced radiations and divergences of enzyme families

Land plants constantly respond to fluctuations in their environment. Part of their response is the production of a diverse repertoire of specialized metabolites. One of the foremost sources for metabolites relevant to environmental responses is the phenylpropanoid pathway, which was long thought to be a land-plant-specific adaptation shaped by selective forces in the terrestrial habitat. Recent data have, however, revealed that streptophyte algae, the algal relatives of land plants, have candidates for the genetic toolkit for phenylpropanoid biosynthesis and produce phenylpropanoid-derived metabolites. Using phylogenetic and sequence analyses, we here show that the enzyme families that orchestrate pivotal steps in phenylpropanoid biosynthesis have independently undergone pronounced radiations and divergence in multiple lineages of major groups of land plants; sister to many of these radiated gene families are streptophyte algal candidates for these enzymes. These radiations suggest a high evolutionary versatility in the enzyme families involved in the phenylpropanoid-derived metabolism across embryophytes. We suggest that this versatility likely translates into functional divergence, and may explain the key to one of the defining traits of embryophytes: a rich specialized metabolism.     

Estuarine fish and tetrapod evolution: insights from a Late Devonian (Famennian) Gondwanan estuarine lake and a southern African Holocene equivalent

The Waterloo Farm lagerstätte in South Africa provides a uniquely well‐preserved record of a Latest Devonian estuarine ecosystem. Ecological evidence from it is reviewed, contextualised, and compared with that available from the analogous Swartvlei estuarine lake, with a particular emphasis on their piscean inhabitants. Although the taxonomic affinities of the estuarine species are temporally very different, the overall patterns of utilisation prove to be remarkably congruent, with similar trophic structures. Significantly, both systems show evidence of widespread use of estuaries as fish nurseries by both resident and marine migrant taxa. Holocene estuaries are almost exclusively utilised by actinopterygians which are overwhelmingly dominated by oviparous species. Complex strategies are utilised by estuarine resident species to avoid exposure of eggs to environmental stresses that characterize these systems. By contrast, many of the groups utilising Devonian estuaries were likely live bearers, potentially allowing them to avoid the challenges faced by oviparous taxa. This may have contributed to dominance of these systems by non‐actinoptergians prior to the End Devonian Mass Extinction. The association of early aquatic tetrapods at Waterloo Farm with a fish nursery environment is consistent with findings from North America, Belgium and Russia, and may be implied by the estuarine settings of a number of other Devonian tetrapods. Tetrapods apparently replace their sister group, the elpistostegids, in estuaries with both groups having been postulated to be adaptated to shallow water habitats where they could access small piscean prey. Correlation of tetrapods (and elpistostegids) with fish nursery areas in the Late Devonian lends strong support to this hypothesis, suggesting that adaptations permitting improved access to the abundant juvenile fish within the littoral zone of estuarine lakes and continental water bodies may have been pivotal in the evolution of tetrapods.     

Divergence of the phytochrome gene family predates angiosperm evolution and suggests thatSelaginella andEquisetum arose prior toPsilotum

Thirty-two partial phytochrome sequences from algae, mosses, ferns, gymnosperms, and angiosperms (11 of them newly released ones from our laboratory) were analyzed by distance and characterstate approaches (PHYLIP, TREECON, PAUP). In addition, 12 full-length sequences were analyzed. Despite low bootstrap values at individual internal nodes, the inferred trees (neighbor joining, Fitch, maximum parsimony) generally showed similar branching orders consistent with other molecular data. Lower plants formed two distinct groups. One basal group consisted of Selaginella, Equisetum, and mosses; the other consisted of a monophyletic cluster of frond-bearing pteridophytes. Psilotum was a member of the latter group and hence perhaps was not, as sometimes suggested, a close relative of the first vascular plants. The results further suggest that phytochrome gene duplication giving rise to a- and b- and later to c-types may have taken place within seedfern genomes. Distance matrices dated the separation of mono- and dicotyledons back to about 260 million years before the present (Myr b.p.) and the separation of Metasequoia and Picea to a fossil record-compatible value of 230 Myr B.P. The Ephedra sequence clustered with the c- or a-type and Metasequoia and Picea sequences clustered with the b-type lineage. The paleoherb Nymphaea branched off from the c-type lineage prior to the divergence of mono- and dicotyledons on the a- and b-type branches. Sequences of Piper (another paleoherb) created problems in that they branched off from different phytochrome lineages at nodes contradicting distance from the inferred trees' origin. Correspondence to: H.A.W. Schneider-Poetsch     

Tangled up in two: a burst of genome duplications at the end of the Cretaceous and the consequences for plant evolution

Genome sequencing has demonstrated that besides frequent small-scale duplications, large-scale duplication events such as whole genome duplications (WGDs) are found on many branches of the evolutionary tree of life. Especially in the plant lineage, there is evidence for recurrent WGDs, and the ancestor of all angiosperms was in fact most likely a polyploid species. The number of WGDs found in sequenced plant genomes allows us to investigate questions about the roles of WGDs that were hitherto impossible to address. An intriguing observation is that many plant WGDs seem associated with periods of increased environmental stress and/or fluctuations, a trend that is evident for both present-day polyploids and palaeopolyploids formed around the Cretaceous–Palaeogene (K–Pg) extinction at 66 Ma. Here, we revisit the WGDs in plants that mark the K–Pg boundary, and discuss some specific examples of biological innovations and/or diversifications that may be linked to these WGDs. We review evidence for the processes that could have contributed to increased polyploid establishment at the K–Pg boundary, and discuss the implications on subsequent plant evolution in the Cenozoic.     

Ecology and evolution join forces to good effect

The 3rd regular meeting of the Canadian Society of Ecology and Evolution was held at the University of British Columbia, Vancouver, Canada from 11 to 14 May 2008.     

Cooperation and the evolution of anisogamy

In the lights of the concept of cooperation wholes, I discuss why the differentiation of sperm and ova can occur with a mathematical model. Most of Parker's explanations for anisogamy are not completely proper, because it is proved that sperm competition is neither sufficient nor necessary for anisogamy and cooperation to deal with fertilization risks is the real key to understand the evolution of anisogamy. According to the computer simulation results, the transport of gametes between different individuals, risks of the transport, the consequent inequality of sperm and eggs and competition among different individuals were the main causes of gamete differentiation. But these factors have different roles and effects. The transport risk is the main reason for individuals of different mating types to cooperate and differentiate into sperm and egg producers. The transported gametes have an advantage to evolve into sperm to seek for a larger gamete number over the fixed gametes, because they suffer more risks as they can encounter the same fixed gamete and less sibling competition as they can be dispersed better. Gamete competition among different individuals just causes the transported gametes to become as small as possible if they have already become smaller beyond a critical state. In the final discussion, I further put the evolution of anisogamy into a broader background of levels of selection and of the evolution of cooperation, the most important existential mode of matters that makes life as life.     

Simulation of evolution implemented in the mutualistic symbioses towards enhancing their ecological efficiency, functional integrity and genotypic specificity

We created the mathematical model for the evolution of the Efficiency of Mutualistic Symbioses (EMS) which was estimated as the microsymbiont impacts on the host’s reproductive potential. Using the example of rhizobia–legume interaction, the relationships were studied between EMS and Functional Integrity of Symbiosis (FIS) which is represented as a measure for concordance of changes in the partners’ genotypic frequencies under the environmental fluctuations represented by the minor deviations of the systemic model parameters. The FIS indices correlate positively with EMS values suggesting an enhancement of FIS via the natural selection operating in the partners’ populations in favor of high EMS. Due to this selection, nodular habitats may be closed for colonization by the non-beneficial bacterial strains and the Genotypic Specificity of Mutualism (GSM) in partners’ interactions is enhanced: the selective advantage of host-specific vs non-host-specific mutualists is increasing. The novelty of our model is to suggest a selective background for macroevolutionary events reorganizing the structure and functions of symbiotic systems and to present its evolution as a result of shifting the equilibrium between different types of mutualists under the impacts of the symbiosis-stipulated modes of natural selection.     

Ecology,evolution and genetics join together on Canada's east coast

The 4th regular meeting of the Canadian Society of Ecology and Evolution was held in conjunction with the 52nd Annual Conference of the Genetics Society of Canada at Dalhousie University, Halifax, from 14 to 17 May 2009.     

中国现代人起源与演化的考古学思考

越来越多古人类学和旧石器时代考古学的发现与研究成果极大地改变了我们对现代人类起源、演化和适应理论问题的看法。中国作为东亚一个幅员辽阔的地理区域,已经成为这项研究的热点地带。来自该地区新的人类化石和石器组合对基于非洲和欧亚大陆西部记录构建的“现代人出自非洲说”（Recent Out-of-Africa）提出了挑战。新的古人类学研究结果表明,早期现代人出现于约100 kaBP的中国南方,可能（至少部分）是由那里的土著居民演化而来。一些古老型智人表现出了与早期古人类镶嵌或过渡性的体质特征,并可能与尼安德特人和丹尼索瓦人杂交混合。同时一些遗址出土的打制石器表现出了早期现代人类技术和行为的复杂性。中国北方的小石片石器主工业和南方的砾石石器主工业贯穿于整个更新世,然而从约40 kaBP开始,石叶技术开始出现在中国北部,紧随其后的是这些地区骨制工具和个人装饰品的出现,这表明,更新世晚期西伯利亚和中亚地区与我国北部可能存在着紧密的文化关系,东北亚地区可能存在着由西北向东南的迁移路线。人类化石和考古学证据表明,中国现代人类起源和适应的过程与机制可能与欧亚大陆西部有所不同。本文对有关中国现代人类起源与演化的考古学研究所取得的新发现和进展进行了总体回顾,从考古学角度阐述了对相关重要学术问题的看法,并为未来的研究提出了方向性建议。     

On the early stages of the evolution of the geosphere and biosphere

The conditions necessary for the existence of nucleic-protein life are as follows: the presence of liquid water, an atmosphere, and a magnetic field (all of which protect from meteorites, abrupt changes in temperature, and a flow of charged particles from space) and the availability of nutrients (macro-and microelements in the form of dissolved compounds). In the evolution of the geosphere, complex interference of irreversible processes (general cooling, gravitational differentiation of the Earth’s interior, dissipation of hydrogen, etc.) with cyclic processes of varying natures and periodicities (from the endogenic cycles “from Pangea to Pangea” to Milankovitch cycles), these conditions have repeatedly changed; hence, in the coevolution of the geosphere and biosphere, the vector of irreversible evolution was determined by the geosphere. Only with the appearance of the ocean as a global system of homeostasis, which provided the maintenance and leveling of nutrient concentrations in the hydrosphere, and the conveyor of nutrients from the mantle, “the film of life” could begin its expansion from the source of the nutrients. Life itself is a system of homeostasis, but not due to the global size and a vast buffer capacity, but because of the high rate of reactions and presence of a program (genome) that allowed its development (ontogeny) independent from the outside environment. The early stages of the origin and evolution of the biosphere (from the RNA-world to the development of the prokaryotic ecosystems) were characterized by the domination of chemotrophic ecosystems. The geographical ranges of these ecosystems were directly or indirectly (through the atmosphere and hydrosphere) tied to the sources of nutrients in the geosphere, which were in turn connected to various sources of volcanic and geotectonic activity (geothermal waters, “black smokers” along the rift zones, etc.). This gave the biosphere consisting of chemotrophic ecosystems a mosaic appearance composed of separate local oases of life. The decrease of methane and accumulation of O₂ in the atmosphere in the geological evolution of the Earth caused the extinction of chemotrophic ecosystems and directed evolution of the biosphere toward autotrophy. Autotrophic photosynthesis gave the biosphere an energy source that was not connected to the geosphere, and for the first time allowed its liberation from the geosphere by developing its own vector of evolution. This vector resulted in the biosphere forming a continuous film of life on the planet by capturing the continents and occupying pelagic and abyssal zones, and the appearance of eukaryotes. The geosphere formed biogeochemical cycles in parallel to the geochemical ones, and comparable in the annual balances of participating matter.     

台湾海峡两岸生物地理学现状与展望

对生物地理学的内涵、研究领域以及进展进行了简要介绍,认为近期海峡两岸生物地理学研究的目标应当集中在:末次冰期以来的海陆变迁对海峡两岸生物分布格局的影响、海峡两岸现代动植物区系的渊源、台湾最后与大陆保持陆地连接的时间与范围、东亚岛弧对台湾生物分布格局的影响以及海峡两岸生物的物种分化与生态适应性五个方面。同时认为海峡两岸现代生物分布格局关系的一种历史生物地理学解释,应当是一个在末次冰期期间存在于长江以南的大区系片断化后对海峡两岸各自不同环境适应的结果。     

Progressive sequence alignment and molecular evolution of the Zn-containing alcohol dehydrogenase family

Summary Sequences of 47 members of the Zn-containing alcohol dehydrogenase (ADH) family were aligned progressively, and an evolutionary tree with detailed branch order and branch lengths was produced. The alignment shows that only 9 amino acid residues (of 374 in the horse liver ADH sequence) are conserved in this family; these include eight Gly and one Val with structural roles. Three residues that bind the catalytic Zn and modulate its electrostatic environment are conserved in 45 members. Asp 223, which determines specificity for NAD, is found in all but the two NADP-dependent enzymes, which have Gly or Ala. Ser or Thr 48, which makes a hydrogen bond to the substrate, is present in 46 members. The four Cys ligands for the structural zinc are conserved except in -crystallin, the sorbitol dehydrogenases, and two bacterial enzymes. Analysis of the evolutionary tree gives estimates of the times of divergence for different animal ADHs. The human class II () and class III () ADHs probably diverged about 630 million years ago, and the newly identified human ADH6 appeared about 520 million years ago, implying that these classes of enzymes may exist or have existed in all vertebrates. The human class I ADH isoenzymes (, , and ) diverged about 80 million years ago, suggesting that these isoenzymes may exist or have existed in all primates. Analysis of branch lengths shows that these plant ADHs are more conserved than the animal ones and that class III ADHs are more conserved than class I ADHs. The rate of acceptance of point mutations (PAM units) shows that selection pressure has existed for ADHs, implying that these enzymes play definite metabolic roles.Offprint requests to: B.V. Plapp     

Non‐angiosperm phytochromes and the evolution of vascular plants

The phytochromes, a class of plant light‐sensing pigments, are a gene family with a long, complex evolutionary history. Angiosperms each have five or more phytochromes (designated A to E in Arabidopsis ) with distinct functions as light receptors and only moderate sequence identities for different types within a species. The long‐term challenge taken up here is to trace the origin and function of the various motifs within the angiosperm phytochromes through gymnosperm phytochromes (types N, O and P) and lower plant phytochromes, sometimes reaching even to bacterial progenitor molecules. Particularly intriguing are the findings of homology of a C‐terminal region of phytochromes with bacterial transmitter modules and of a large N‐terminal region with a protein encoded by a gene from the cyanobacterum Synechocystis . Phylogenetic analysis helps to answer general questions such as the times of divergence of mono‐ and dicotyledons, of groups of gymnosperms or of ferns. Phytochrome sequences suggest (1) that mono‐ and dicotyledons became separated 150‐200 million years earlier than indicated by the fossil record and (2) that Ginkgo and Cycas have been separated unexpectedly late from the lineage giving rise to the Pinidae. (3) The status of Psilotum as a close relative of the primeval vascular plants is not supported. Phytochrome gene sequences additionally reveal that (4) moss and fern phytochromes have erratically acquired C‐termini which, though kinase‐like, are different from the common ones and that (5) introns have been lost, gained or shifted in position from algae to angiosperms. Phytochromes promise to be a rich source of phylogenetic information into the future as more sequences and functional data emerge, not least from studies of lower plants.     

京津冀地区生态系统服务供需测度及时空演变

生态系统服务供需状况深刻影响地区生态安全与经济社会的可持续发展。以京津冀为研究区,选取产水、碳固持以及粮食生产3项生态系统服务,应用InVEST模型、ArcGIS空间分析模块,定量测度2005年与2015年京津冀地区生态系统服务供需水平,探索京津冀地区生态系统服务供需时空格局及类型演变特征。结果表明：（1）在2005年与2015年,京津冀地区产水服务处于供需赤字状态,碳固持服务和粮食生产服务处于供需盈余状态,且各项生态系统服务供需状态呈显著的空间异质性;（2）2005年与2015年京津冀地区生态系统服务综合供需状况呈赤字状态,主要赤字范围以京津为中心呈环状分布,并在此期间范围有所扩大。（3）基于生态系统服务供需状况,将京津冀各县区划分为生态涵养区、生态保育区、生态改良区、生态恢复区以及生态控制区,并针对各类型区生态系统供需状况及其时空演变规律,提出了相应的政策建议。