Importance of demographic history for phylogeographic inference on the arctic–alpine plant Phyllodoce caerulea in East Asia

Arctic–alpine plants have enormous ranges in the Northern Hemisphere. Phylogeographic studies have provided insights into their glacial survival as well as their postglacial colonization history. However, our understanding of the population dynamics of disjunct alpine populations in temperate regions remains limited. During Pleistocene cold periods, alpine populations of arctic–alpine species in East Asia were either connected to an ice-free Beringia refugium or they persisted with prolonged isolation after their establishment. To estimate which of these scenarios is more likely, we elucidated the genetic structure of Phyllodoce caerulea (Ericaceae) in Beringia and northern Japan, East Asia. Sequence variation in multiple nuclear loci revealed that P. caerulea can be distinguished into northern and southern groups. A demographic analysis demonstrated that the north–south divergence did not predate the last glacial period and detected introgression from Phyllodoce aleutica, relative widely distributed in East Asia, exclusively into the southern group. Therefore, although there has been genetic divergence between northern Japan and Beringia in P. caerulea, the divergence is unlikely to have resulted from their prolonged geographic separation throughout several cycles of glacial and interglacial periods. Instead, our study suggests that the introgression contributed to the genetic divergence of P. caerulea and that the range of P. caerulea was plausibly connected between northern Japan and Beringia during the last glacial period. Overall, our study not only provides a biogeographic insight into alpine populations of arctic–alpine plants in East Asia but also emphasizes the importance of careful interpretation of genetic structure for inferring phylogeographic history.     

Flowering phenology influences seed production and outcrossing rate in populations of an alpine snowbed shrub, Phyllodoce aleutica: effects of pollinators and self-incompatibility

Background and Aims

Because of differences in snowmelt time, the reproductive phenologies of alpine plants are highly variable among local populations, and there is large variation in seed set across populations. Temporal variation in pollinator availability during the season may be a major factor affecting not only seed production but also outcrossing rate of alpine plants.

Methods

Among local populations of Phyllodoce aleutica that experience different snowmelt regimes, flowering phenology, pollinator availability, seed-set rate, and outcrossing rate were compared with reference to the mating system (self-compatibility or heterospecific compatibility with a co-occurring congeneric species).

Key Results

Flowering occurred sequentially among populations reflecting snowmelt time from mid-July to late August. The visit frequency of bumble-bees increased substantially in late July when workers appeared. Both seed set and outcrossing rate increased as flowering season progressed. Although flowers were self-compatible and heterospecific compatible, the mixed-pollination experiment revealed that fertilization with conspecific, outcrossing pollen took priority over selfing and hybridization, indicating a cryptic self-incompatibility. In early snowmelt populations, seed production was pollen-limited and autogamous selfing was common. However, genetic analyses revealed that selfed progenies did not contribute to the maintenance of populations due to late-acting inbreeding depression.

Conclusions

Large variations in seed-set and outcrossing rates among populations were caused by the timing of pollinator availability during the season and the cryptic self-incompatibility of this species. Despite the intensive pollen limitation in part of the early season, reproductive assurance by autogamous selfing was not evident. Under fluctuating conditions of pollinator availability and flowering structures, P. aleutica maintained the genetic composition by conspecific outcrossing.Key words: Alpine snowbed, autogamy, bumble-bee, cryptic self-incompatibility, flowering phenology, mixed pollination, outcrossing rate, Phyllodoce aleutica, pollination success, seasonality, self-pollination     

Comparative age-specific demography of four commercially important deep-water snappers: implication for fishery management of a long-lived lutjanid

Lutjanid snappers belonging to the subfamilies Apsilinae and Etelinae are commercially valuable components of tropical deep-water fisheries throughout much of the Indo-Pacific region. Based on age assessment using sagittal otoliths, the age-specific demographic characteristics of four deep-water snappers, Etelis coruscans, Paracaesio caerulea, Pristipomoides filamentosus and Pristipomoides sieboldii, in the Okinawa Islands, southwestern Japan, were examined, and the results were discussed for fishery management. Age validation using edge-type analysis demonstrated that opaque zones in all species were formed once per year and were considered valid annual growth increments. The von Bertalanffy growth equations were also determined for each species. These snappers are long-lived (>50 years for E. coruscans and P. caerulea and >30 years for the two species of Pristipomoides) and relatively slow-growing. The age of acquiring sexual maturity in females was relatively later in E. coruscans, P. caerulea and P. filamentosus than in P. sieboldii. The results revealed differences in the biological traits among these four species; E. coruscans and P. caerulea with long life spans and late maturation are particularly more vulnerable to fishing impact than the two Pristipomoides species. Therefore, further approaches to decrease and control fishing intensity, such as networking of marine-protected areas and regulation to control the numbers of boats and/or the total allowable catch, are necessary for the management of the stock of these species, especially for E. coruscans and P. caerulea.     

Assessment of genetic diversity in <Emphasis Type="Italic">Trigonella foenum-graecum</Emphasis> and <Emphasis Type="Italic">Trigonella caerulea</Emphasis>using ISSR and RAPD markers

Background  

Various species of genus Trigonella are important from medical and culinary aspect. Among these, Trigonella foenum-graecum is commonly grown as a vegetable. This anti-diabetic herb can lower blood glucose and cholesterol levels. Another species, Trigonella caerulea is used as food in the form of young seedlings. This herb is also used in cheese making. However, little is known about the genetic variation present in these species. In this report we describe the use of ISSR and RAPD markers to study genetic diversity in both, Trigonella foenum-graecum and Trigonella caerulea.     

Floral morphogenesis in choripetalous and sympetalous members of Ericaceae s.l. (Ericales)

Choripetaly of Rhododendron tomentosum is the result of secondary reduction. Our data support earlier observations of P. Leins [4] that a rudimentary corolla tube occurs during early stages of floral development. Choripetaly of Empetrum may be a plesiomorphic condition, because no vestiges of corolla tube were detected throughout all stages of flower development. However, a complete loss of corolla tube cannot be excluded. The type of congenital petal fusion varies within a group of closely related taxa of Ericaceae: Phyllodoce caerulea has late sympetaly while Loiseleuria procumbens has early sympetaly. In contrast, in euasterids, the type of sympetaly appears to characterize taxa of higher rank.     

Plant population patterns in a glacier foreland succession: pioneer herbs and later-colonizing shrubs

Glacier forelands provide valuable sites for the study of patterns and processes of primary succession The Storbreen glacier foreland, south-central Norway, has previously been the subject of a suite of community-level approaches Here the focus is population-level studies of six key pioneer plants Arabis alpina, Deschampsia alpina, Oxyria digyna, Poa alpina, Saxifraga cespitosa and Trisetum spicatum and four heath shrubs Betula nana, Empetrum hermaphroditum, Phyllodoce caerulea and Salix complex (Salix glauca and S lanata) Size-frequency distributions are used as indicators of population phenomena The optimal conditions for growth and fecundity of Arabis alpina and Saxifraga cespitosa appear to be almost immediately upon deglaciation (< 7 yr), although both achieve their maximum cover on slightly older terrain A similar pattern was noted for the grasses, although their optima come at slightly later times Population phenomena in Oxyria digyna could not be related to terrain age, indicating a differing role of this species within the succession Among the shrubs a number of demographic and behavioural patterns are identifiable as successional features, and other patterns are explicable in relation to changing histories of disturbance and site moisture relationships Establishment and build-up in Empetrum hermaphroditum and Salix complex do not appear to be environmentally constrained, in contrast to Betula nana and Phvllodoce caerulea Betula nana is the slowest species to colonize, but occasional early colonists appear healthy and fecund, pointing to a strong environmental sieve at the point of establishment The population patterns varied considerably among the shrub species, the most striking common feature being in mode of establishment Establishment of ramets by vegetative means occurs increasingly on older ground within the foreland (maximum terrain age 230 yr) but only in sites of the mature heath outside the foreland is it the characteristic mode of establishment In exception to this pattern, vegetative establishment of Phyllodoce caerulea is very rare in all conditions studied Discussion focuses upon the importance of population phenomena, seed dispersal and life-history characteristics in successional change It is concluded that processes involved in species turnover include both allogenic and autogenic elements and that although the two classes are difficult to separate in practice, autogenic factors become more significant whilst species-environment relations become tighter in later phases of vegetation development     

Identification of multiple nursery habitats of skates in the eastern Bering Sea

The use of more than a single nursery habitat type is examined for oviparous elasmobranchs using data summarized from studies conducted on the Alaska skate Bathyraja parmifera and the Aleutian skate Bathyraja aleutica in the eastern Bering Sea. The eastern Bering Sea skate species use two discrete areas as nurseries, one for egg deposition and a second for newly emergent juveniles. Egg deposition sites were located along the outer shelf and upper slope near canyons in the eastern Bering Sea. Newly emergent juveniles were found along the outer and middle shelf for B. parmifera and deep‐slope for B. aleutica, suggesting that habitat used by newly emergent juvenile skates is distinct from habitat used for egg deposition and embryo development. In reviewing many studies on oviparous elasmobranchs, similar patterns emerge of habitat use during their early life history. To distinguish these distinct habitats, appropriate terminology is proposed. Egg case nursery is suggested for areas of egg deposition and juvenile nursery is suggested for areas where juveniles aggregate after emergence. Criteria to describe each habitat type are outlined.     

Reproductive biology of skates, Bathyraja(Ishiyama), along the eastern Bering Sea continental slope

Eight Bathyraja skate species, Bathyraja aleutica, Bathyraja interrupta, Bathyraja lindbergi, Bathyraja maculata, Bathyraja minispinosa, Bathyraja parmifera, Bathyraja taranetzi and Bathyraja trachura, collected along the eastern Bering Sea continental slope were examined for reproductive features. The sex ratio for six of eight Bathyraja skate species studied was c. 1 : 1 with males dominating for B. interrupta and B. parmifera. Five of eight species examined, B. aleutica, B. lindbergi, B. parmifera, B. taranetzi and B. trachura approximately equalled or exceeded the maximum reported total length, L_T. Maximum L_T for three species (B. interrupta, B. minispinosa and B. parmifera) revealed that the largest individuals were males and that in three other species (B. lindbergi, B. maculata and B. trachura) both females and males grew to approximately the same size. Both females and males matured at approximately the same L_T for seven of eight species. Only female B. aleutica matured at a much larger size than males. All species matured within a narrow size range following a protracted juvenile stage and once mature these skates appeared to grow very little. First maturity for all species occurred at ≥80% of their maximum L_T. The number of oocytes did not significantly differ between the left and right ovary for any species. The total number of mature oocytes increased with L_T for six of eight species examined. The number of oocytes for one species, B. aleutica, increased in number until 145 cm L_T, then declined in those individuals greater in length. The egg cases of all species are described, of which four (B. lindbergi, B. parmifera, B. taranetzi and B. trachura) have never been previously reported. Evidence is presented that suggest skates may be limited to as little as one and as many as 13 spawning seasons, depending on the species, and that senescence may occur in older individuals.     

Speciation via hybrid dysgenesis: negative evidence from the Drosophila affinis subgroup

Differences in karyotypic structure are compared with reported isozyme differences in three Mediterranean species of Patella. In addition, the karyotypic structure of Patella is discussed in terms of the karyotypic variability of Archaeogastropoda. Both P. lusitanica and P. caerulea have a haploid complement of n=9 (6 metacentric, 1 submetacentric, 1 subtelocentric, 1 telocentric chromosome in P. lusitanica and 6 metacentric, 1 submetacentric, 2 telocentric chromosome in P. caerulea). P. aspera, although regarded as morphologically more closely related to P. caerulea, has a haploid complement of only n=8 (7 metacentric and 1 submetacentric chromosomes).     

Sewage pollution and extinction risk: an endangered limpet as a bioindicator?

The mollusc Patella ferruginea, endemic to the Mediterranean, is the most endangered marine species on the list of the European Council Directive 92/43/EEC and it is under serious risk of extinction. In spite of the low abundances and restricted distribution of this limpet, important populations have been found in the harbour of Ceuta, north Africa. The main objective of the present study was to characterise, for the first time, the effects of sewage pollution on P. ferruginea and related limpet species, and to evaluate the potential value of these limpet assemblages as bioindicators, using univariate and multivariate analyses. Physicochemical parameters and limpets were sampled in nine stations located at 0, 1, 2, 4, 8, 16, 32, 64 and 128 m away from the discharge point of a sewage effluent in Ceuta harbour. The stations closer to the outfall (0, 1, 2, 4 and 8) were characterised by higher values of turbidity, phosphate and ammonia in the water column, and organic matter, faecal coliforms and faecal Streptococci in sediments. A total of six limpet species were found and studied (Patella ferruginea, P. caerulea, P. nigra, P. rustica, P. ulyssiponensis and Siphonaria pectinata); the number of limpet species increased with increasing distance from the outfall, while diversity and evenness reached the highest values at intermediate sites. Siphonaria pectinata and P. caerulea were the most resistant and abundant species, while P. ferruginea was the most sensitive species to sewage pollution, only found at stations from 32 to 128 m. The distribution of this endangered limpet seems mainly affected by the pollution gradient, and not by the competition with the remaining limpets. The results of this study should be taken into account in future programmes of management and conservation of P. ferruginea.     

Intrinsic and extrinsic factors acting on the reproductive process in alpine‐snowbed plants: roles of phenology,biological interaction,and breeding system

Pollinator activity and competition for pollinators lead to quantitative and qualitative pollen limitations on seed production and affect the reproductive success of plant species, depending on their breeding system (e.g., self‐compatibility and heterospecific compatibility) and genetic load (e.g., inbreeding depression and hybrid inviability). In alpine ecosystems, snowmelt regimes determine the distribution and phenology of plant communities. Plant species growing widely along a snowmelt gradient often grow with different species among local populations. Their pollinators also vary in their abundance, activity, and behavior during the season. These variations may modify plant–pollinator and plant–plant interactions. We integrated a series of our studies on the alpine dwarf shrub, Phyllodoce aleutica (Ericaceae), to elucidate the full set of intrinsic (species‐specific breeding system) and extrinsic factors (snow condition, pollinator activity, and interspecific competition) acting on their reproductive process. Seasonality of pollinator activity led to quantitative pollen limitation in the early‐blooming populations, whereas in the late‐blooming populations, high pollinator activity ensured pollination service, but interspecific competition for pollinators led to qualitative and quantitative pollen limitation in less competitive species. However, negative effects of illegitimate pollen receipt on seed‐set success might be reduced when cryptic incompatibility systems (i.e., outcross pollen grains took priority over self‐ and heterospecific pollen grains) could effectively prevent ovule and seed discounting. Our studies highlight the importance of species‐specific responses of plant reproduction to changing pollinator availability along environmental gradients to understand the general features of pollination networks in alpine ecosystems.     

Reproductive biology and population structure of white warehou Seriolella caerulea and silver warehou Seriolella punctata in the austral zone off Chile

Using an extensive database compiled by scientific observers aboard commercial fishing operations between 1984 and 2014, we describe the maturity and size structures of white warehou Seriolella caerulea and silver warehou Seriolella punctata from by-catch of the trawling industrial fisheries operating in the austral zone off Chile. Macroscopic maturity stages and gonadosomatic (I_GS) index show mature females throughout the year and a pronounced spawning period in both austral autumn and winter seasons, with an I_GS peak in July for S. punctata and August for S. caerulea. Reproductive patterns in both species show an extended spawning season (July to September) across the area between 43 and 47° S. Length–mass relationships showed significant differences between sexes in both species, where females reach a larger size. Fork length at 50% maturity was 43.5 cm for S. caerulea and 37.2 cm for S. punctata. Female catch composition is dominated by adult fish (96% for S. caerulea and 86% for S. punctata). Currently, both species are exploited with no assessment and management-decision framework. Therefore, information regarding reproductive biology and demographic traits becomes an important baseline to ensure adequate fisheries management for both species.     

A multilocus sequencing approach reveals the cryptic phylogeographical history of Phyllodoce nipponica Makino (Ericaceae)

Discordant phylogeographical patterns among species with similar distributions may not only denote specific biogeographical histories of different species, but also could represent stochastic variance of genealogies in applied genetic markers. A multilocus investigation representing different genomes can be used to address the latter concern, allowing robust inference to biogeographical history. In the present study, we conducted a multilocus phylogeographical analysis to re‐examine the genetic structuring of Phyllodoce nipponica, in which chloroplast (cp)DNA markers exhibited a discordant pattern compared to those of other alpine plants. The geographical structure of sequence variation at five nuclear loci was not consistent with that of cpDNA and showed differentiation between the northern and southern parts of the range of this species. Its demographic history inferred from the isolation‐with‐migration model suggests that the north–south divergence originated from Pleistocene vicariance. In addition, the demographic parameters showed a lack of chloroplast‐specific gene flow, suggesting that stochastic variance in genealogy resulted in the discordant geographical structure. Thus, P. nipponica probably experienced Pleistocene vicariance between its southern and northern range parts in concordance with other alpine plants in the Japanese archipelago. The findings of the present study demonstrates the importance of using a multilocus approach for inferring population dynamics, as well as for reconciling discordant phylogeographical patterns among species. © 2013 The Linnean Society of London, Biological Journal of the Linnean Society, 2013, 110 , 214–226.     

The diet of field volesMicrotus agrestis at low population density in upland Britain

Studies on field volesMicrotus agrestis Linnaeus, 1758 in lowland grasslands have shown them to be unselective grazers. The diet of the field vole in upland Britain was investigated using feeding trials with four of the dominant British upland monocots,Molinia caerulea,Nardus stricta, Deschampsia flexuosa and Eriophorum vaginatum. The suitability of faecal analysis was assessed and then used to analyse the diet of wild voles from faecal samples. Percentages of plant species in the faeces were compared to percentages on the ground in sites dominated byMolinia caerulea, Eriophorum vaginatum,Nardus stricta andCalluna vulgaris. Significant preferences for the grassDeschampsia flexuosa were observed in feeding trials and in the wild while the sedgeEriophorum vaginatum was avoided in both. There was no clear preference forMolinia caerulea andNardus stricta. Preference for plant species was related to palatability and nutrient content. The low nutrient conditions in British uplands mean that voles that live in these environments must be selective feeders to maximise nutrient intake.     

Genetic relationship of myrmecophyte (Anthorrhiza caerulea) individuals within and among territories of the arboreal ant (Dolichoderus sp.) detected using random amplified polymorphic DNA markers

The polymorphic bands of random amplified polymorphic DNA (RAPD) markers of the population of epiphytic myrmecophyte, Anthorrhiza caerulea Huxley & Jebb (Rubiaceae: Hydnophytinae), and those of its obligatory occupant ant, Dolichoderus sp. (Formicidae: Dolichoderinae), were analyzed in this study in a Papua New Guinean secondary forest. All the reproducible polymorphic bands were used as RAPD markers. Euclidean distances were calculated as the similarity indices of the band patterns, and similarities of the band patterns were analyzed using cluster analysis (group average method). It was frequently observed that the occupant ants gathered the seeds of A. caerulea and buried them into their carton trails on the bark of host trees. The colony boundaries of Dolichoderus sp. were assessed using RAPD analysis, and were clearly detected by this method. From the analyses of genetic relationships in the population of A. caerulea, it was revealed that the myrmecophytes within a certain ant territory consisted of close relatives. Therefore, it was inferred that the descendants of A. caerulea were dispersed only within the territory of a certain ant colony.     

Karyotypes of two Australian hylids

Karyotypes of the Australian frogs Hyla caerulea (White) and Hyla phyllochroa Günther were prepared and analysed from colchicine-treated, primary cultures of adult heart, lung and kidney. The diploid chromosome number for both species is 26, which resembles that of most known karyotypes of Papuan hylids but differs from that of the genus Hyla in other regions. A statistical comparison, involving data from arm ratios, centromere indices and relative lengths of metaphase chromosomes of H. caerulea incubated at 31 and 37° C respectively, showed that the differences were non-significant at the 5% level. Similar treatment of data from cells of Hyla caerulea and Hyla phyllochroa incubated at 31°C failed to show a significant species difference. Absolute lengths of haploid chromosome complements of the two species at the same temperature were found to differ significantly. No evidence of sexual heteromorphism was found. The occurrence of a terminally situated, heterochromatic zone associated with the longer arm of the 11th pair of chromosomes is described and compared with the known condition in other hylids.     

Genetic analysis of Pinus strobus and Pinus monticola populations from Canada using ISSR and RAPD markers: development of genome-specific SCAR markers

Pinus is the largest genus of conifers, containing over 100 species and is also the most widespread genus in the Northern Hemisphere. Pinus monticola and P. strobus are two closely related and economically important species in Canada. Morphological and allometric characteristics have been used to assess genetic variation within these two species but these markers are not reliable due to ecological variations. The purpose of the present study was to determine the level of genetic diversity within and among Canadian populations from the two species using molecular markers and to identify and characterize genome-specific inter-simple sequence repeats (ISSR) and random amplified polymorphic DNA (RAPD) markers. The level of genetic variation among populations was much lower for P. monticola than P. strobus. For both species, the among population variation values were smaller than within population variation. The populations from P. monticola were more closely genetically related than populations from P. strobus based on ISSR and RAPD analyses. Six ISSR and four RAPD markers specific to either P. monticola or P. strobus were cloned and sequenced. Primer pairs flanking these specific sequences were designed and genome specific SCAR markers for P. monticola and P. strobus were developed and characterized.     

Experimental manipulation of succession in heathland ecosystems

An experiment was carried out in two heathland ecosystems, one dominated by Calluna vulgaris and the other by Molinia caerulea, to analyse the effects of soil organic matter accumulation and nutrient mineralization on plant species dynamics during succession. The experiment included one treatment that received nutrient solution and two treatments where the rate of soil organic matter accumulation was reduced by removing litter or accelerated by adding litter. In a fourth treatment the C. vulgaris litter produced in the C. vulgaris-dominated plots was replaced by litter of M. caerulea and vice versa. Treatments were applied over 8 years. Addition of nutrient solution caused C. vulgaris to decline, and grass species to increase sharply, compared to the control plots. Addition of litter enhanced both N mineralization and the biomass of M. caerulea and Deschampsia flexuosa but reduced the biomass of C. vulgaris. The effects of replacing C. vulgaris litter by M. caerulea litter, or vice versa, on N mineralization and species dynamics could not be attributed to differences between the decomposability of the different litter materials that were transferred. The results confirm the hypothesis that increased litter inputs accelerate the rate of species replacement during succession.     

<Emphasis Type="Italic">Molinia caerulea</Emphasis> responses to N and P fertilisation in a dry heathland ecosystem (NW-Germany)

In the present study we analysed whether airborne N pollution may constitute one important driver for the encroachment of Molinia caerulea in dry heathland ecosystems. Based on full-factorial field experiments (in 2006 and 2008) and complementary greenhouse experiments (in 2008), we quantified growth responses of Molinia caerulea to N and P fertilisation (50 kg N ha⁻¹ year⁻¹, 20 kg P ha⁻¹ year⁻¹). Aboveground biomass production of Molinia caerulea was limited by P in 2006, but by N in both experiments in 2008. In the greenhouse experiment, N addition caused a sixfold increase of the biomass of vegetative tillers, and in all experiments the biomass and numbers of flowering tillers showed a significant increase due to fertilisation. Our experiments indicated that growth of Molinia caerulea was primarily limited by N, but in dry heaths the kind of nutrient limitation may be mediated by other factors such as water availability during the vegetative period. Shifts in biomass allocation patterns resulting from N fertilisation showed that Molinia caerulea encroachment in dry heaths is not only attributable to increased leaf biomass, but also due to higher investments in reproductive tissue that allow for increased seed production and thus accelerated encroachment of seedlings in places where the dwarf shrub canopy has been opened after disturbance.     

Use of sod cutting for restoration of wet heathlands: revegetation and establishment of typical species in relation to soil conditions

Abstract. Field observations in sod-cut wet Molinia caerulea dominated heath lands revealed information on the revegetation process in relation to plot age and soil variables. Data on the most common species; Erica tetralix, Molinia caerulea, Drosera intermedia and Rhynchospora fusca show that Molinia tends to dominate again within a few years. Whereas the other species are affected in their development by either groundwater regime, soil acidity, nutrient availability or cut depth, Molinia caerulea is highly competitive in all situations. Soil acidity is the major factor affecting species diversity.