Historical biogeography of the ancient lycophyte genus Selaginella: early adaptation to xeric habitats on Pangea

The ancient and cosmopolitan lycophyte genus Selaginella has living representatives around the world, but their historical biogeography has not been assessed with modern methods. We estimated a time‐calibrated phylogeny using DNA marker regions rbcL and ITS1‐5.8S‐ITS2 from 200 species. Node density analyses revealed that Selaginellaceae has significantly older median and mean node ages than other putative “ancient” families. We used statistical model comparison to assess different biogeographical models on our dated tree, and to estimate ancestral ranges. These revealed that Selaginella originated on Euramerica around 383 Ma in the Devonian period, while its peak diversification began with the formation of Pangea. The divergence of the two main species‐rich Selaginella lineages occurred approximately 318 Ma on the supercontinent. The major divergences within these main lineages of Selaginella took place in the Late Permian and Early Triassic, along with lineages highly adapted for xeric habitats on Pangea.     

Battenizyga, a new Early Triassic gastropod genus with a discussion of the caenogastropod evolution at the Permian/Triassic boundary

Battenizyga, a new Early Triassic gastropod genus from the Moenkopi Formation of Utah, is described and the speciesAnoptychia eotriassica Batten & Stokes, 1986 is placed in it. The new genus has an axially ribbed planktonic larval shell and a teleoconch with an angulated periphery. This character combination is unknown from the Palaeozoic. Therefore,Battenizyga represents additional evidence that recovery from the end-Permian mass extinction was connected with a faunal turnover. Additionally, the extinction of diverse Palaeozoic groups of the Caenogastropoda in the Permian (e.g., the Pseudozygopleuridae) suggest a turnover. All caenogastropod genera that hold Early Triassic species, have post-Palaeozoic type species and most were not reported from the Palaeozoic. This corroborates the view that there was an intense faunal turnover within the Caenogastropoda.Battenizyga is probably a caenogastropod that is closely related to the superfamily Zygopleuroidea which is abundant in the late Palaeozoic and early Mesozoic.      

Climate change and the spread of vector‐borne diseases: using approximate Bayesian computation to compare invasion scenarios for the bluetongue virus vector Culicoides imicola in Italy

Bluetongue (BT) is a commonly cited example of a disease with a distribution believed to have recently expanded in response to global warming. The BT virus is transmitted to ruminants by biting midges of the genus Culicoides, and it has been hypothesized that the emergence of BT in Mediterranean Europe during the last two decades is a consequence of the recent colonization of the region by Culicoides imicola and linked to climate change. To better understand the mechanism responsible for the northward spread of BT, we tested the hypothesis of a recent colonization of Italy by C. imicola, by obtaining samples from more than 60 localities across Italy, Corsica, Southern France, and Northern Africa (the hypothesized source point for the recent invasion of C. imicola), and by genotyping them with 10 newly identified microsatellite loci. The patterns of genetic variation within and among the sampled populations were characterized and used in a rigorous approximate Bayesian computation framework to compare three competing historical hypotheses related to the arrival and establishment of C. imicola in Italy. The hypothesis of an ancient presence of the insect vector was strongly favoured by this analysis, with an associated P ≥ 99%, suggesting that causes other than the northward range expansion of C. imicola may have supported the emergence of BT in southern Europe. Overall, this study illustrates the potential of molecular genetic markers for exploring the assumed link between climate change and the spread of diseases.     

The Devonian nekton revolution

Klug, C., Kröger, B., Kiessling, W., Mullins, G.L., Servais, T., Frýda, J., Korn, D. & Turner, S. 2009: The Devonian nekton revolution. Lethaia, 10.1111/j.1502‐3931.2009.00206.x Traditional analyses of Early Phanerozoic marine diversity at the genus level show an explosive radiation of marine life until the Late Ordovician, followed by a phase of erratic decline continuing until the end of the Palaeozoic, whereas a more recent analysis extends the duration of this early radiation into the Devonian. This catch‐all approach hides an evolutionary and ecological key event long after the Ordovician radiation: the rapid occupation of the free water column by animals during the Devonian. Here, we explore the timing of the occupation of the water column in the Palaeozoic and test the hypothesis that ecological escalation led to fundamental evolutionary changes in the mid‐Palaeozoic marine water column. According to our analyses, demersal and nektonic modes of life were probably initially driven by competition in the diversity‐saturated benthic habitats together with the availability of abundant planktonic food. Escalatory feedback then promoted the rapid rise of nekton in the Devonian as suggested by the sequence and tempo of water‐column occupation. □Devonian, diversity, ecology, food webs, nekton, plankton, radiation.     

Phylogeography of the pallid kangaroo mouse,Microdipodops pallidus: a sand‐obligate endemic of the Great Basin,western North America

Aim Kangaroo mice, genus Microdipodops Merriam, are endemic to the Great Basin and include two species: M. pallidus Merriam and M. megacephalus Merriam. The pallid kangaroo mouse, M. pallidus, is a sand‐obligate desert rodent. Our principal intent is to identify its current geographical distribution and to formulate a phylogeographical hypothesis for this taxon. In addition, we test for orientation patterns in haplotype sharing for evidence of past episodes of movement and gene flow. Location The Great Basin Desert region of western North America, especially the sandy habitats of the Lahontan Trough and those in south‐central Nevada. Methods Mitochondrial DNA sequence data from portions of three genes (16S ribosomal RNA, cytochrome b, and transfer RNA for glutamic acid) were obtained from 98 individuals of M. pallidus representing 27 general localities sampled throughout its geographical range. Molecular sequence data were analysed using neighbour‐joining, maximum‐parsimony, maximum‐likelihood and Bayesian methods of phylogenetic inference. Directional analysis of phylogeographical patterns, a novel method, was used to examine angular measurements of haplotype sharing between pairs of localities to detect and quantify historical events pertaining to movement patterns and gene flow. Results Collecting activities showed that M. pallidus is a rather rare rodent (mean trapping success was 2.88%), and its distribution has changed little from that determined three‐quarters of a century ago. Two principal phylogroups, distributed as eastern and western moieties, are evident from the phylogenetic analyses (mean sequence divergence for cytochrome b is c. 8%). The western clade shows little phylogenetic structure and seems to represent a large polytomy. In the eastern clade, however, three subgroups are recognized. Nine of the 42 unique composite haplotypes are present at two or more localities and are used for the orientation analyses. Axial data from haplotype sharing between pairwise localities show significant, non‐random angular patterns: a north‐west to south‐east orientation in the western clade, and a north‐east to south‐west directional pattern in the eastern clade. Main conclusions The geographical range of M. pallidus seems to be remarkably stable in historical times and does not show a northward (or elevationally upward) movement trend, as has been reported for some other kinds of organism in response to global climate change. The eastern and western clades are likely to represent morphologically cryptic species. Estimated times of divergence of the principal clades of M. pallidus (4.38 Ma) and between M. pallidus and M. megacephalus (8.1 Ma; data from a related study) indicate that kangaroo mice diverged much earlier than thought previously. The phylogeographical patterns described here may serve as a model for other sand‐obligate members of the Great Basin Desert biota.     

A new species of Macaranga from the middle Miocene of Fujian,China and its significance

A well-preserved Macaranga leaf fossil from the middle Miocene Fotan Group of Zhangpu County, Fujian, South-eastern China is examined and described as a new species, Macaranga zhangpuensis Z.X. Wang et B.N. Sun sp. nov. The present fossil species represents the highest latitudinal distribution of a reliable Macaranga fossil in the world, and we present the first Macaranga fossil described with detailed cuticular characteristics from China. Based on the global palaeogeographic distribution of Macaranga, we infer that the genus probably originated during the Oligocene in Africa and spread from Africa to India and then to South Fujian, China, further into the Mariana Islands and finally into the Philippine Islands and Malaysia, leading to its present distribution. In addition, the new leaf material described herein is the first Macaranga fossil record with three leaf tips globally. This finding demonstrates that Macaranga with three leaf tips already existed during the Miocene in Fujian, South China, and it provides new information for understanding climatic changes between the Miocene and the present -day.     

The Tetrapod Ichnogenus Protochirotherium Fichter and Kunz 2004, a Characteristic Early Triassic Morphotype of Central Pangea

Early Triassic chirotherian footprint assemblages from Poland, Germany, and Morocco are important for understanding archosaur evolution in the aftermath of the Permian-Triassic crisis. However, their ichnotaxonomy is confusing because various authors have interpreted their diversity differently. After an analysis and ichnotaxonomic re-assessment, the presence of the ichnogenera Brachychirotherium, Isochirotherium, and Chirotherium in these assemblages is not supported. Distant similarities with these ichnotaxa are functions of extra morphological variation and substrate-related factors. Instead, Early Triassic chirotherian footprints described under these names are assigned here to the ichnogenus Protochirotherium and to a more slender morphotype identified as Synaptichnium. In particular, Protochirotherium appears to be more widely distributed in central Pangea as a characteristic morphotype reflecting a distinct stage in archosaur evolution. Trackmakers were nonarchosaurian archosauriforms or, alternatively, stem-group crocodylians. Morphologically and temporally these footprints match the hypothetical ancestor of the Chirotherium barthii trackmaker. Chirotherium barthii appears by the beginning of the Middle Triassic. Because of its restricted stratigraphic range, and its wider distribution in central Pangea, Protochirotherium also has biostratigraphic significance for this region and can be considered as an indicator of Early Triassic-aged strata.     

Towards the conservation of the Mesozoic relict fern Christensenia: a fern species with extremely small populations in China

The Chinese occurrences of the marattioid fern genus Christensenia have been considered as requiring protection because of its extreme rarity and very small population size. Here, we explored different biological aspects to enable protection of these rare ferns, well known as Mesozoic living fossils. Firstly, we documented the cytology of the Chinese occurrences for the first time. This is the second tetraploid record of Christensenia worth for further studies to confirm its taxonomic status. Secondly, we obtained the first complete plastid genome of this genus, which confirmed the proposed conservatism of the plastid genome structure in marattioid ferns. By comparing the chloroplast genome with other marattioids, we identified several candidate regions to develop highly variable markers to investigate the intra-species diversity of marattioid ferns. Thirdly, phylogenetic analyses of rbcL sequences implied that there are at least two distinct species of Christensenia. Finally, we re-assessed the conservation status of Christensenia in the context of its local and global distribution by assessing specimen information extracted from publications and digitized voucher information. This assessment confirmed the need to obtain more accurate information about the distribution of this genus to assess the status incorporating the disjunct distribution from southern China and India in the North towards the Solomon Islands in the South.

     

Global conservation status assessment of the threatened aquatic plant genus Baldellia (Alismataceae): challenges and limitations

In this study, the aquatic monocot Baldellia (Alismataceae) is used as a model for evaluating the general hindrances and shortfalls in the global conservation status assessment of a threatened taxon. Our study clearly shows that Linnean shortfalls (uncertainty in the number of species and taxonomy) and the Wallacean shortfall (fragmentary knowledge regarding distribution) form the basis for all other hindrances. We demonstrate that even in Europe, which has traditionally been very well investigated, between 60 and 75% of regions or countries possess no detailed distribution maps and/or data banks for Baldellia spp. Furthermore, between 50 and 60% of regions do not have any published red list category. Thus, only general conclusions concerning the global conservation status of the three Baldellia taxa are possible—a global assessment of conservation status for B. ranunculoides subsp. repens is nearly impossible. Baldellia ranunculoides s.str. shows a strong decline in practically all regions of its natural range, and thus it is probably the most threatened species in the genus. Baldellia alpestris is the least threatened species in the genus, even though it is a narrow endemic. Our case study clearly shows the need for reinforced coordination of research and conservation activities as well as an urgent need for data accessibility regarding taxonomic, chorological and conservation studies of endangered species. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Silurian craniide brachiopods from Gotland

Seven craniide brachiopod genera are described from the Silurian (Wenlock–Ludlow) of Gotland, including one new genus and five new species. The new genus and species Thulecrania septicostata is unique among Silurian craniides as it possesses solid spines. The new species Lepidocrania multilamellosa is the first Silurian record of this poorly known Permian genus. The problematic North American Propatella Grubbs, 1939 , was originally described as a gastropod, but the new species Propatella palmaria from Gotland shows that it is a craniide with sutured hollow spines of a type not previously recorded from Silurian craniides. The dorsal valves of the new species Valdiviathyris? bicornis are remarkably similar to those of the type species and represent the first possible Palaeozoic record of this poorly known extant craniide. This first systematic study of craniide brachiopods from the Silurian of Gotland shows that the diversity is relatively high as compared to other known Silurian craniide faunas, but a more thorough comparison is not possible due to the lack of data from most parts of the world. The new data from Gotland support the view that the craniides were not affected by the end‐Ordovician extinction.     

Biogeographically dynamic genetic structure bridging two continents in the monotypic Central American rodent Ototylomys phyllotis

Central America is an ideal region in which to study patterns of historical divergence and population genetic differentiation, because of its extraordinarily dynamic biogeographical, tectonic, and climatic history. The rodent Ototylomys phyllotis is the only extant species of the genus Ototylomys and is distributed within this region from the Isthmus of Tehuantepec, Mexico, to central Costa Rica, offering an excellent opportunity to study spatial and temporal patterns of population structure of the species and to explain the ecological and evolutionary processes responsible for those patterns. We estimated the genetic diversity and structure within and between populations of O. phyllotis, times of divergence, and migration patterns using mitochondrial DNA and a comprehensive combination of phylogenetic and phylogeographical computational analyses. Our results support monophyly of the genus Ototylomys. We identified three major phylogeographical lineages within O. phyllotis that are linked to its diversification and coincide with the main geological features that shaped Middle America. The origin of the genus was before 3.35 Mya, prior to the Great American Biotic Interchange (GABI), and its initial occurrence was near the centre of its current distribution (Honduras/El Salvador), from which it later spread (3.20–2.84 Mya) following a series of GABI pulses. The species showed an initial northward dispersal to the Chiapas and Guatemala highlands (2.27 Mya) followed by diversification. A later dispersal (1.82 Mya) occurred toward both the south (Nicaragua, Costa Rica) and the north (Belize). The Yucatan peninsula was colonized (0.8 Mya) by individuals from Belize. Extremely high radiation and range expansion occurred throughout the entire range, the highest of which was in the Yucatan peninsula (0.125 Mya). © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, 107 , 593–610.     

Geographic distribution and migration pathways of Pistacia – present,past and future

The global distribution of Pistacia is correlated to its adaptability to environmental conditions and mechanisms that had driven the genus to the current unique narrow latitudinal belt in between 10° North and 45° North. The current geostatisitcal distribution maps of the genus are shown and the derived probability maps over a period between 121 Kyr before present and the year 2100 were calculated. The tolerance of Pistacia trees to harsh climate conditions was related to leaf phenology, evergreeness vs deciduousness, which has led to geographic classification of the genus in two corresponding sections that corroborate recent molecular genetic studies. The deciduous trees are more tolerant to extreme climate conditions (?26°C to 46°C) than the evergreen species (?8°C to 41°C), except Pistacia lentiscus, which occurs at a max. temperature of 45°C. The close spatial distribution of the later species and the deciduous ones may have been conducive in further evolution of the genus. Based on the long evolution of Pistacia (approx. 84 Ma), we suggested that the genus may have originated in the boreal forest and its migration pathways might have been evoked in relation to climate change, shifting the species distribution to evolving suitable environmental conditions. The fact that most of the genera in the family of Anacardiaceae and the whole genus Pistacia are dioecious raised questions about plausible relationships between the geographic distribution, environmental conditions and evolution of dioecy. The genus Pistacia was shown to be a good candidate for research about the relationships between environmental conditions, adaptation traits and geographic distribution.     

Redistribution of the lizardfish Harpadon nehereus in coastal waters of China due to climate change

Climate change has the potential to greatly alter species distributions and threatens biodiversity in marine ecosystems. Mapping changes in species distribution patterns under climate change will help facilitate management strategies to maintain ecosystem structure and function. The lizardfish Harpadon nehereus is an aggressive predator that has experienced rapid population growth along the coast of China in recent decades, compressing the ecological niches of other marine species and disrupting food webs. If this species’ range is shifting due to climate change, it could further impact the integrity of ecological communities. To map the distribution of H. nehereus, we developed an ensemble species distribution model and projected the present and future habitat suitability in Chinese coastal waters. Annual mean benthic water temperature was identified as the most important variable affecting the projected distribution of H. nehereus, followed by water depth and salinity. Currently suitable habitats are along the coast from Guangxi Province to the southern Jiangsu Province. As climate changes, the southern portion of its distribution is predicted to recede with habitat losses, and the overall suitable habitat will shift northward. To avoid the potential impacts of H. nehereus redistribution, precautionary management based on species distribution modeling would help to maintain healthy marine ecosystems in the newly invaded areas.

     

Non-lethal presence of predators modifies morphology and movement rates in <Emphasis Type="Italic">Euplotes</Emphasis>

Many species are able to modify aspects of their behaviour and morphology in the presence of predators. The aim of this study was to investigate the relationship between the expression of morphological and behavioural defences according to the framework proposed by DeWitt et al (1999). Experiments were carried out using hypotrich ciliates of the genus Euplotes as prey and turbellarians of the genus Stenostomum as predators. The smaller species Euplotes octocarinatus showed a greater proportional increase in width, a reduction in foraging movement rates and an increase in maximum movement rates following exposure to predator cues. The larger Euplotes aediculatus induced lesser changes in width, similar reductions in movement during foraging and no change in maximum speed following predator exposure. These results provide evidence of a cospecialised relationship between morphological and behavioural defences. Despite substantial differences in the absence of predators, movement rates and lateral body width were similar in both species following predator exposure. The observed changes may be considered adaptive, gape limited flatworm predators are unable to ingest large Euplotes and a reduction in movement rates during foraging reduces predator encounter rates, while an increase in maximal movement rates increases chances of predator evasion. Handling editor: S. I. Dodson     

Effect of climate change in lizards of the genus Xenosaurus (Xenosauridae) based on projected changes in climatic suitability and climatic niche conservatism

Accelerated climate change represents a major threat to the health of the planet's biodiversity. Particularly, lizards of the genus Xenosaurus might be negatively affected by this phenomenon because several of its species have restricted distributions, low vagility, and preference for low temperatures. No study, however, has examined the climatic niche of the species of this genus and how their distribution might be influenced by different climate change scenarios. In this project, we used a maximum entropy approach to model the climatic niche of 10 species of the genus Xenosaurus under present and future suitable habitat, considering a climatic niche conservatism context. Therefore, we performed a similarity analysis of the climatic niche between each species of the genus Xenosaurus. Our results suggest that a substantial decrease in suitable habitat for all species will occur by 2070. Among the most affected species, X. tzacualtipantecus will not have suitable conditions according to its climatic niche requirements and X. phalaroanthereon will lose 85.75% of its current suitable area. On the other hand, we found low values of conservatism of the climatic niche among species. Given the limited capacity of dispersion and the habitat specificity of these lizards, it seems unlikely that fast changes would occur in the distribution of these species facing climate change. The low conservatism in climatic niche we found in Xenosaurus suggests that these species might have the capacity to adapt to the new environmental conditions originated by climate change.     

Diversification and biogeographic history of the Western Palearctic freshwater flatworm genus Schmidtea (Tricladida: Dugesiidae), with a redescription of Schmidtea nova

The freshwater flatworm genus Schmidtea is endemic in the Western Palearctic region, where it is represented by only four species, thus contrasting with the high species diversity of the closely related genus Dugesia within Europe. Although containing an important model species in developmental and regeneration research, viz. Schmidtea mediterranea, no evolutionary studies on the genus Schmidtea have been undertaken. For the first time, we present a well‐resolved molecular phylogenetic tree of the four species of the genus, inferred on the basis of two molecular markers, and provide also the first detailed morphological account of Schmidtea nova. The phylogenetic tree generated corroborates an earlier speciation hypothesis based on karyological data and points to chromosomal rearrangements as the main drivers of speciation in this genus. The high genetic divergence between the four species, in combination with previous dating studies and their current geographic distribution, suggests that Schmidtea could have originated in Laurasia but lost most of its diversity during the Oligocene. Thus, its present distribution pattern may be the result of the expansion of three of its four relictual species over Europe, probably after the Pleistocene glaciations. Our detailed morphological study of S. nova revealed that it shows a number of remarkable features: interconnected testis follicles, parovaria, an ejaculatory duct exiting into the primary as well as the secondary seminal vesicle by means of a nipple, and the wall of the distal section of the ejaculatory duct being sclerotic or chitinized.     

末次盛冰期以来红豆树在不同气候变化情景下的分布动态

红豆树(Ormosia hosiei Hemsl. et Wils.)是中国的特有种,具有极高的经济价值、景观价值和药用价值。由于木材珍贵,人工盗伐严重,其种群数量和分布范围不断减少,被世界自然保护联盟(IUCN)濒危物种红色名录列为近危物种。气候变化会对物种的分布造成严重影响,理解该影响将有助于物种保护策略的制定,尤其是为濒危物种未来的保护提供重要参考。本研究基于红豆树在中国的地理分布数据,借助经相关性分析后筛选出的9个变量因子,利用最大熵模型(MaxEnt)模拟红豆树在末次盛冰期、全新世中期、当代和未来气候情景下的潜在分布区和影响其分布的主导环境因子,并且通过空间分析模拟其在不同气候变化情景下空间分布格局的变化。结果表明MaxEnt在各时期训练集和测试集的AUC(受试者工作特征曲线下的面积)均值均达到0.9以上,表明模型有很好的预测能力。刀切法(Jackknife)表明年均降水量、温度季节性变化标准差和昼夜温差月均值是主导其分布的三大因子,累计贡献率达到91.8%。将模拟结果导入到ArcGIS后,处理得到不同时期红豆树适生区空间分布格局变化。结果表明,自末次盛冰期以来,红豆树的适生区面积收缩且向北迁移;随着全球气候变暖,未来4种气候情景下红豆树适生区也有较大面积的减少,特别是在高浓度排放情景(RCP8.5)下,丧失率最高达到45.6%。重庆、江西及位于粤桂两省中南部的大面积适生区可能都将丧失。而新增区域仅存在于适生区的边缘,新增率仅为1%—2%。本研究表明红豆树的分布受气候变化的影响较大,该研究结果将对未来红豆树的迁地保护与栽培提供重要参考。     

Rapid radiation of Impatiens (Balsaminaceae) during Pliocene and Pleistocene: Result of a global climate change

Impatiens comprises more than 1000 species and is one of the largest genera of flowering plants. The genus has a subcosmopolitan distribution, yet most of its evolutionary history is unknown. Diversification analyses, divergence time estimates and historical biogeography, illustrated that the extant species of Impatiens originated in Southwest China and started to diversify in the Early Miocene. Until the Early Pliocene, the net diversification rate within the genus was fairly slow. Since that time, however, approximately 80% of all Impatiens lineages have originated. This period of rapid diversification coincides with the global cooling of the Earth’s climate and subsequent glacial oscillations. Without this accelerated diversification rate, Impatiens would only have contained 1/5th of its current number of species, thereby indicating the rapid radiation of the genus.     

The biogeographic and tectonic history of India

Aim To present an up to date account of the Mesozoic history of India and its relationship to the other Gondwana continents and to Eurasia. Location Continents surrounding the Western Indian Ocean. Methods Utilization of recent evidence of continental relationships based upon research in stratigraphy, palaeomagnetism, palaeontology, and contemporary biotas. Results The physical data revealed a sequence of events as India moved northward: (1) India–Madagascar rifted from east Africa 158–160 Ma (million years ago), (2) India–Madagascar from Antarctica c. 130 Ma, (3) India–Seychelles from Madagascar 84–96 Ma, (4) India from Seychelles 65 Ma, (5) India began collision with Eurasia 55–65 Ma and (6) final suturing took place c. 42–55 Ma. However, data from fossil and contemporary faunas indicate that, throughout the late Cretaceous, India maintained exchanges with adjacent lands. There is an absence in the fossil record of peculiar animals and plants that should have evolved, had India undergone an extended period of isolation just before its contact with Eurasia. Main conclusions The depiction of India in late Cretaceous as an isolated continent is in error. Most global palaeomaps, including the most recent one, show India, as it moves northward, following a track far out in the Indian Ocean. But the evidence now indicates that India's journey into northern latitudes cannot have taken place under such isolated circumstances. Although real breaks among the lands were indicated by the physical data, faunal links were maintained by vagile animals that were able to surmount minor marine barriers. India, during its northward journey, remained close to Africa and Madagascar even as it began to contact Eurasia.