ANCIENT TEPUI SUMMITS HARBOR YOUNG RATHER THAN OLD LINEAGES OF ENDEMIC FROGS

The flattop mountains (tepuis) of South America are ancient remnants of the Precambrian Guiana Shield plateau. The tepui summits, isolated by their surrounding cliffs that can be up to 1000 m tall, are thought of as “islands in the sky,” harboring relict flora and fauna that underwent vicariant speciation due to plateau fragmentation. High endemicity atop tepui summits support the idea of an ancient “Lost World” biota. However, recent work suggests that dispersal between lowlands and summits has occurred long after tepui formation indicating that tepui summits may not be as isolated from the lowlands as researchers have long suggested. Neither view of the origin of the tepui biota (i.e., ancient vicariance vs. recent dispersal) has strong empirical support owing to a lack of studies. We test diversification hypotheses of the Guiana Shield highlands by estimating divergence times of an endemic group of treefrogs, Tepuihyla. We find that diversification of this group does not support an ancient origin for this taxon; instead, divergence times among the highland species are 2–5 Ma. Our data indicate that most highland speciation occurred during the Pliocene. Thus, this unparalleled landscape known as “The Lost World” is inhabited, in part, not by Early Tertiary relicts but neoendemics.     

Potential migration routes and barriers for vascular plants of the Neotropical Guyana Highlands during the Quaternary

Aim To reconstruct in detail the potential migration routes and barriers for vascular plants from the summits of the Guyana mountains during the Quaternary, in order to test the possibility of migration among them during the glaciations. These changes in connectivity are predicted based on the altitudinal migration of plant communities associated with glacial cooling. To examine the effects of these cycles, the extent of the potential biotic interchange and its influence on patterns of endemism was modelled. Location The summits of the tepuis or table mountains of the Neotropical Guyana Highlands, which constitute the peculiar and discontinuous Pantepui phytogeographical province (total surface 5000 km², altitudinal range 1500–3014 m a.s.l.), and is characterized by a unique and diverse flora with a high degree of endemism. Methods GIS‐based palaeotopographical reconstruction using a high‐precision digital elevation model, combined with phytogeographical analysis by means of a data base built up from the Flora of the Venezuelan Guyana, which includes the geographical and altitudinal ranges for each Pantepui species. Results During the Last Glacial Maximum, which serves as a representative of a standard Quaternary glaciation, most migration pathways among tepuis were open for species with lower altitudinal levels (LAL) ≤ 1500 m (1678 species or c. 69% of the total Pantepui species), and closed for species with LAL ≥ 2300 m altitude (c. 3%). The species in between these altitudes have intermediate migratory possibilities, depending on the district and the tepui considered. If these local factors are considered, the number of species with no possibility of glacial interchange increases to 202 or c. 8% of the total. The strongest topographical barrier separated eastern locations above 1600–1700 m elevation from all others. The highest possibility of interchange was among the tepuis of the eastern sector, in which internal topographical barriers were only effective for species with LALs at or above 1900 m. Main conclusions The Quaternary evolution of the vascular flora from the Guyana Highlands took place in a predominantly migration‐prone, glacial‐era landscape, in which more than 70% of the flora (maximum estimate) was able to move from one tepuian district to another, thanks to the downward bioclimatic shift caused by cooling. Interglacials were too short to drive significant evolutionary diversification. A number of present high‐altitude local endemics are species that were unable to migrate, even during glaciations. However, some endemic species do appear to have been able to migrate among regions, suggesting that topographical isolation alone is not enough to explain patterns of endemism. Other factors such as tepui summit area, habitat heterogeneity or pre‐Quaternary evolution are considered. These studies should be complemented with palaeoecological and phylogeographical surveys.     

Molecular evolution, adaptive radiation, and geographic diversification in the amphiatlantic family Rapateaceae: evidence from ndhF sequences and morphology

Rapateaceae (16 genera, approximately 100 species) is largely restricted to the tepuis and sandplains of the Guayana Shield in northern South America, with Maschalocephalus endemic to West Africa. The family has undergone extensive radiation in flower form, leaf shape, habit, and habitat. To analyze the evolution of these distributions and traits, we derived a molecular phylogeny for representatives of 14 genera, based on sequence variation in the chloroplast-encoded ndhF gene. The lowland subfamily Rapateoideae is paraphyletic and includes the largely montane subfamily Saxofridericioideae as a monophyletic subset. Overall, the morphological/anatomical data differ significantly from ndhF sequences in phylogenetic structure, but show a high degree of concordance with the molecular tree in three of four tribes. Branch lengths are consistent with the operation of a molecular clock. Maschalocephalus diverges only slightly from other Monotremae: it is the product of relatively recent, long-distance dispersal, not continental drift--only its habitat atop rifted, nutrient-poor sandstones is vicariant. The family appears to have originated approximately 65 Mya in inundated lowlands of the Guayana Shield, followed by: (1) wide geographic spread of lowland taxa along riverine corridors; (2) colonization of Amazonian white-sand savannas in the western Shield; (3) invasion of tepui habitats with frequent speciation, evolution of narrow endemism, and origin of hummingbird pollination in the western Shield; and (4) reinvasion of lowland white-sand savannas. The apparent timing of speciation in the Stegolepis alliance about 6-12 Mya occurred long after the tepuis began to be dissected from each other as the Atlantic rifted approximately 90 Mya. Given the narrow distributions of most montane taxa, this suggests that infrequent long-distance dispersal combined with vicariance accounts for speciation atop tepuis in the Stegolepis alliance.     

Establishing a baseline of plant diversity and endemism on a neotropical mountain summit for future comparative studies assessing upward migration: an approach from biogeography and nature conservation

Climate change is forcing many plant species to shift their range in search of adequate environmental conditions, being localized endemic species particularly at risk on mountain summits. The Pantepui biogeographic province, a set of flat-topped mountain summits (called tepuis) of northern South America, contains both high plant diversity and a high degree of endemism. Previous studies based on warming projections for the area suggested that half of the Pantepui endemic flora would disappear due to habitat loss by 2100. In this study, we selected one of the best-explored tepuis, Roraima-tepui, to establish the baseline of diversity and endemism for comparisons with historical data and future monitoring surveys, aimed at testing the hypothesis of upward migration of plants in response to global warming. We also analysed floristic and physiognomic features of the Eastern Tepui Chain (ETC, the mountain range where Roraima is located), and the phytogeographic patterns of both the ETC and Pantepui. The Roraima summit contains 227 species, including 44 new records, 13 exotic species (some of them with high invasive potential), and at least one species new to science. At the ETC level, Roraima is the tepui with highest species richness and degree of endemism, and shows a relatively high floristic similarity with Kukenán and Ilú. Herbaceous species dominate over shrubs on these tepuis, Tramen and Maringma, whereas on Yuruaní, Karaurín and Uei, they reach similar abundances. At the Pantepui level, endemic species have highly localized distribution patterns (17% local endemics). Conservation opportunities are evaluated in light of these results.     

Sexual dimorphism in the hoverfly motion vision pathway

Many insects perform high-speed aerial maneuvers in which they navigate through visually complex surrounds. Among insects, hoverflies stand out, with males switching from stationary hovering to high-speed pursuit at extreme angular velocities [1]. In dipterans, 50-60 large interneurons -- the lobula-plate tangential cells (LPTCs) -- detect changes in optic flow experienced during flight [2-5]. It has been predicted that large LPTC receptive fields are a requirement of accurate "matched filters" of optic flow [6]. Whereas many fly taxa have three horizontal system (HS) LPTC neurons in each hemisphere, hoverflies have four [7], possibly reflecting the more sophisticated flight behavior. We here show that the most dorsal hoverfly neuron (HS north [HSN]) is sexually dimorphic, with the male receptive field substantially smaller than in females or in either sex of blowflies. The (hoverfly-specific) HSN equatorial (HSNE) is, however, sexually isomorphic. Using complex optic flow, we show that HSN, despite its smaller receptive field, codes yaw velocity as well as HSNE. Responses to a target moving against a plain or textured background suggest that the male HSN could potentially play a role in target pursuit under some conditions.     

Isolation and characterization of halophilic bacteria from Urmia Lake in Iran

Urmia Lake is one of the most permanent hypersaline lakes in the world which is threatened by hypersalinity and serious dryness. In spite of its importance no paper has been published regarding bacterial community of this lake. Accordingly, the present study aimed to investigate the halophilic bacteria in the aforementioned lake. In so doing, thirty seven strains were isolated on six different culture media. The isolated strains were characterized using phenotypic and genotypic methods. Growth of the strains occurred at 2535 degrees C, pH 6-9 and 7 to 20% (w/v) NaCl indicating that most of the isolates were moderately halophiles. Catalase, oxidase and urease activities were found to be positive for the majority of the isolates. Phylogenetic analysis based on 16S rRNA gene sequences indicated that the isolated bacteria belonged to two major taxa: Gammaproteobacteria (92%, including Salicola [46%], Pseudomonas [13.5%], Marinobacter [ 11%], Idiomarina [11%], and Halomonas [8%]) and Firmicutes (8%, including Bacillus [5%] and Halobacillus [3%]). In addition, a novel bacterium whose 16S rRNA gene sequence showed almost 98% sequence identity with the taxonomically troubled DSM 3050T, Halovibrio denitrificans HGD 3T and Halospina denitrificans HGD 1-3T, each, was isolated. 16S rRNA gene similarity levels along with phenotypic characteristics suggest that some of the isolated strains could be regarded as potential type strain for novel species, on which further studies are recommended.     

Notes on Guayana mosses with new information of Sphagnum ornatum

Mosses restricted to tepui summits in the Guayana Highland are relatively few.Sphagnum ornatum Crum, though, only recently described from Brazil, has been found subsequently in a number of localities in nearby Venezuela and Guyana. These additional collections make for a better understanding of the diagnostic features of a species characteristically found on the tops of the sandstone mesas of southern Venezuela and adjacent areas.     

Male chimpanzees exchange political support for mating opportunities

Male chimpanzees, Pan troglodytes, differ from males in most other mammalian taxa because they remain in their natal communities throughout their lives, form close bonds with one another, and cooperate in a range of activities [1]. However, males also compete fiercely for status within their groups [2,3], and high rank enhances male reproductive success [4,5]. Males rely partly on coalitions to achieve and maintain status [2,3,6-9], and shifts in male alliances can have dramatic political effects [2,3,6]. It is not known what benefits are obtained by low-ranking coalition partners. Here we report that the highest-ranking (alpha) male in one well-studied community of chimpanzees rewarded his allies by allowing them preferential access to mates.     

Six New Species of Dobsonflies from Venezuela (Megaloptera: Corydalidae: Corydalinae)

Six new species of dobsonflies (Megaloptera: Corydalidae: Corydalinae) from Venezuela are described and illustrated, two in the Neotropical genus Chloronia and four within the New World genus Corydalus. Chloronia contains 15 previously described species. The new species Chloronia gaianii and C. marthae, each from the southern tepui and Andean regions, appear to be most closely related to C. hieroglyphica (Rambur) and C. mirifica Navás, respectively. Corydalus contains 30 previously described species. The new species Corydalus clavijoi, of Andean affinity, appears most closely related to C. tesselatus Stitz; C. crossi, from Auyán-tepuí, might be closely related to C. nubilus Erichson; and both C. hayashii and C. mayri, from westernmost Guayana Shield area and Aracamuni tepuis, respectively, clearly belong in the distinct and monophyletic C. arpi Navás species group. Keys for identification of the Venezuelan species of both genera are provided.     

Evidence for degeneration of the Y chromosome in the dioecious plant Silene latifolia

The human Y--probably because of its nonrecombining nature--has lost 97% of its genes since X and Y chromosomes started to diverge [1, 2]. There are clear signs of degeneration in the Drosophila miranda neoY chromosome (an autosome fused to the Y chromosome), with neoY genes showing faster protein evolution [3-6], accumulation of unpreferred codons [6], more insertions of transposable elements [5, 7], and lower levels of expression [8] than neoX genes. In the many other taxa with sex chromosomes, Y degeneration has hardly been studied. In plants, many genes are expressed in pollen [9], and strong pollen selection may oppose the degeneration of plant Y chromosomes [10]. Silene latifolia is a dioecious plant with young heteromorphic sex chromosomes [11, 12]. Here we test whether the S. latifolia Y chromosome is undergoing genetic degeneration by analyzing seven sex-linked genes. S. latifolia Y-linked genes tend to evolve faster at the protein level than their X-linked homologs, and they have lower expression levels. Several Y gene introns have increased in length, with evidence for transposable-element accumulation. We detect signs of degeneration in most of the Y-linked gene sequences analyzed, similar to those of animal Y-linked and neo-Y chromosome genes.     

Collapse of a new living species of giant clam in the Red Sea

Giant clams are among the most spectacular but also the most endangered marine invertebrates. Their large size and easy accessibility has caused overfishing and collapse of the natural stocks in many places and local extinction in some of the species [1, 2]. The diversity of giant clams is extremely low because of reliction in this Tethyan group [3, 4]. The latest additions of living species date back almost two decades [5-7], fixing the number of extant Tridacna at seven species [3]. Here, we report the discovery of a new species of giant clam: Tridacna costata sp. nov. features characteristic shells with pronounced vertical folds, is genetically distinct, and shows an earlier and abbreviated reproduction than its Red Sea congeners. This species represents less than 1% of the present stocks but up to >80% of the fossil shells. The decline in proportion and shell size (20x) indicates overharvesting [8] dating back to the early human occupation of the Red Sea >125,000 years ago [9]. This earliest depletion reported so far of a shallow-water megafaunal invertebrate has important ramifications for human dispersal out of Africa [10]. Its oversight in one of the best-investigated reef provinces [11-13] illustrates the dearth of knowledge on marine biodiversity.     

Plant communities and environmental factors in the Guayana Highlands: monitoring for conservation under future climate change

The Guayana Highlands (GH) constitute a highly diverse, but relatively poorly studied Neotropical biome, comprised of ～50 flat-topped mountain summits (called tepuis). Previous studies based on warming forecasts for the region suggested that an upward displacement of environmental conditions of 500–700 m could occur by 2100, potentially resulting in the extinction of c. 50% of its endemic flora due to total habitat loss. To assess the ecological responses of the species to climate change, and select the appropriate conservation measures, long-term monitoring of the GH plant communities will be necessary. In this study, the baseline state for future comparisons was established for the best explored tepui in terms of its flora, Roraima-tepui (2810 m), through a floristic characterization of its different vegetation types. We also identified the environmental gradients underlying the major plant communities, and assessed the effects of human activities on the chemistry of soils and water at three field camps. Our results yielded five main community types: three meadows, one shrubland, and one forest, with their corresponding diagnostic species. The herbaceous communities were mainly influenced by the presence of flat sandy soils, with varying flooding capacity. Shrublands and forests were characterized by irregular organic soils with very low pH. Finally, pH values below 3 were measured on an organic soil of a field camp, although further studies will be necessary to attribute this deviation to human activities.     

Interactions of the chemotaxis signal protein CheY with bacterial flagellar motors visualized by evanescent wave microscopy

The chemotaxis signal protein CheY of enteric bacteria shuttles between transmembrane methyl-accepting chemotaxis protein (MCP) receptor complexes and flagellar basal bodies [1]. The basal body C-rings, composed of the FliM, FliG and FliN proteins, form the rotor of the flagellar motor [2]. Phosphorylated CheY binds to isolated FliM [3] and may also interact with FliG [4], but its binding to basal bodies has not been measured. Using the chemorepellent acetate to phosphorylate and acetylate CheY [5], we have measured the covalent-modification-dependent binding of a green fluorescent protein-CheY fusion (GFP-CheY) to motor assemblies in bacteria lacking MCP complexes by evanescent wave microscopy [6]. At acetate concentrations that cause solely clockwise rotation, GFP-CheY molecules bound to native basal bodies or to overproduced rotor complexes with a stoichiometry comparable to the number of C-ring subunits. GFP-CheY did not bind to rotors lacking FIiM/FliN, showing that these subunits are essential for the association. This assay provides a new means of monitoring protein-protein interactions in signal transduction pathways in living cells.     

Modern quantitative acid-base chemistry

Quantitative analysis of ionic solutions in terms of physical and chemical principles has been effectively prohibited in the past by the overwhelming amount of calculation it required, but computers have suddenly eliminated that prohibition. The result is an approach to acid-base which revolutionizes our ability to understand, predict, and control what happens to hydrogen ions in living systems. This review outlines that approach and suggests some of its most useful implications. Quantitative understanding requires distinctions between independent variables (in body fluids: pCO2, net strong ion charge, and total weak acid, usually protein), and dependent variables [( HCO-3], [HA], [A-], [CO(2-)3], [OH-], and [H+] (or pH]. Dependent variables are determined by independent variables, and can be calculated from the defining equations for the specific system. Hydrogen ion movements between solutions can not affect hydrogen ion concentration; only changes in independent variables can. Many current models for ion movements through membranes will require modification on the basis of this quantitative analysis. Whole body acid-base balance can be understood quantitatively in terms of the three independent variables and their physiological regulation by the lungs, kidneys, gut, and liver. Quantitative analysis also shows that body fluids interact mainly by strong ion movements through the membranes separating them.     

Mid-Mesozoic flea-like ectoparasites of feathered or haired vertebrates

Parasite-host associations among insects and mammals or birds are well attended by neontological studies [1]. An Eocene bird louse compression fossil [2, 3] and several flea specimens from Eocene and Oligocene ambers [4-8], reported to date, are exceptionally similar to living louse and flea taxa. But the origin, morphology, and early evolution of parasites and their associations with hosts are poorly known [9, 10] due to sparse records of putative ectoparasites with uncertain classification in the Mesozoic, most lacking mouthpart information and other critical details of the head morphology [11-15]. Here we present two primitive flea-like species assigned to the Pseudopulicidae Gao, Shih et Ren familia nova (fam. nov.), Pseudopulex jurassicus Gao, Shih et Ren genus novum et species nova (gen. et sp. nov) from the Middle Jurassic [16] and P. magnus Gao, Shih et Ren sp. nov. from the Early Cretaceous in China [17]. They exhibit many features of ectoparasitic insects. Large body size and long serrated stylets for piercing tough and thick skin or hides of hosts suggest that these primitive ectoparasites might have lived on and sucked the blood of relatively large hosts, such as contemporaneous feathered dinosaurs and/or pterosaurs or medium-sized mammals (found in the Early Cretaceous, but not the Middle Jurassic).     

Hox genes and the phylogeny of the arthropods

The arthropods are the most speciose, and among the most morphologically diverse, of the animal phyla. Their evolution has been the subject of intense research for well over a century, yet the relationships among the four extant arthropod subphyla - chelicerates, crustaceans, hexapods, and myriapods - are still not fully resolved. Morphological taxonomies have often placed hexapods and myriapods together (the Atelocerata) [1, 2], but recent molecular studies have generally supported a hexapod/crustacean clade [2-9]. A cluster of regulatory genes, the Hox genes, control segment identity in arthropods, and comparisons of the sequences and functions of Hox genes can reveal evolutionary relationships [10]. We used Hox gene sequences from a range of arthropod taxa, including new data from a basal hexapod and a myriapod, to estimate a phylogeny of the arthropods. Our data support the hypothesis that insects and crustaceans form a single clade within the arthropods to the exclusion of myriapods. They also suggest that myriapods are more closely allied to the chelicerates than to this insect/crustacean clade.     

Behavioural evidence for polarisation vision in stomatopods reveals a potential channel for communication.

Polarisation sensitivity (PS) - the ability to detect the orientation of polarised light - occurs in a wide variety of invertebrates [1] [2] and vertebrates [3] [4] [5], many of which are marine species [1]. Of these, the crustacea are particularly well documented in terms of their structural [6] and neural [7] [8] adaptations for PS. The few behavioural studies conducted on crustaceans demonstrate orientation to, or local navigation with, polarised sky patterns [9]. Aside from this, the function of PS in crustaceans, and indeed in most animals, remains obscure. Where PS can be shown to allow perception of polarised light as a 'special sensory quality' [1], separate from intensity or colour, it has been termed polarisation vision (PV). Here, within the remarkable visual system of the stomatopod crustaceans (mantis shrimps) [10], we provide the first demonstration of PV in the crustacea and the first convincing evidence for learning the orientation of polarised light in any animal. Using new polarimetric [11] and photographic methods to examine stomatopods, we found striking patterns of polarisation on their antennae and telson, suggesting that one function of PV in stomatopods may be communication [12]. PV may also be used for tasks such as navigation [5] [9] [13], location of reflective water surfaces [14] and contrast enhancement [1] [15] [16] [17] [18]. It is possible that the stomatopod PV system also contributes to some of these functions.     

An evaluation of the Lost World and Vertical Displacement hypotheses in the Chimantá Massif, Venezuelan Guayana

Aim To document the occurrence of vertical displacements of vegetation in the high plateaus of the Venezuelan Guayana (tepuis) over the last c. 6000 years, and to discuss their significance for the origin of their flora, especially the endemism patterns observed in their flat summits. Two hypotheses have been proposed for the origin of the summit flora. One (the Lost World hypothesis) proposes a long history of evolution in isolation from the surrounding plains, while the other (the Vertical Displacement hypothesis) suggests that vertical movements of vegetation during the Pleistocene glacial‐interglacial cycles would have resulted in floristic mixing within the lowlands, and genetic interchange among plateau summits. Location This work has been conducted on the flat summit of the Churí‐tepui, in the Chimantá massif, at 5°15′ Lat. N and 62°01′ Long. W, around 2250 m altitude. Methods Pollen analysis and radiocarbon dating of two peat outcrops, using modern analogue technique and numerical methods for palaeoecological interpretation were used. Results The replacement of a high‐altitude plant community (a paramoid Chimantaea shrubland) by a lower elevation (< 2300 m) Stegolepis meadow, occurred about 2500 years before present (yr bp ). This vegetation change is inferred to have resulted from a regional climatic shift to higher temperature and moisture. A subsequent decrease in temperature and moisture led to the establishment of present conditions after about 1450 yr bp . Main conclusions The highland vegetation of the tepuis responded to climate shifts with vertical displacements, supporting the hypothesis of vertical mixing. However, a physiographical analysis shows that around half of the tepuis would never have been connected by lowlands. Therefore, both hypotheses are needed to explain the origins of the summit flora in the tepuis.     

Sequence similarities among monkey ori-enriched (ors) fragments

Nucleotide sequences have been determined for eight ors (ori-enriched sequence) fragments isolated from monkey DNA by a method that was designed to enrich for origins of DNA replication [Kaufmann et al., Mol. Cell. Biol. 5 (1985) 721-727]. Evidence has been presented that some or possibly all of these sequences can serve, albeit inefficiently, as oris in vivo [Frappier and Zannis-Hadjopoulos, Proc. Natl. Acad. Sci. USA 84 (1987) 6668-6672]. Two of the fragments were found to contain the long terminal repeat-like elements of the 'O-family' of moderately repetitive sequences that are present in human DNA as a transposon-like element [Paulson et al., Nature 315 (1985) 359-361]. Extensive pair-wise comparisons of the sequences failed to detect any statistically significant common sequences, except for long asymmetrically distributed A + T-rich stretches. Nonetheless, when the ors fragments were examined for the presence of published consensus sequences, seven of eight were found to contain the control sequence described by Dierks et al. [Cell 32 (1983) 695-706], and the same seven of eight were found to contain both the scaffold attachment region T consensus [Gasser and Laemmli, Cell 46 (1986) 521-530] and the minimal Saccharomyces cerevisiae autonomously replicating sequence consensus [e.g., Palzkill and Newlon, Cell 53 (1988) 441-450].