Some species of the genus Phyllodoce (Polychaeta) from Southern California

Five species of the genus Phyllodoce from Southern California are reviewed. A new subgenus (Aponaitides) is proposed and a new species, Phyllodoce cuspidata , is described. A key to the Southern California species is included.     

Phylogeny and classification of the Phyllodocidae (Polychaeta)

The phylogenetic relationships of the benthic phyllodocid genera are estimated by a parsimony analysis based on 26 morphological characters. It is concluded that the cephalization within Eteone has led to a reduction of segment 1 and loss of the first pair of tentacular cirri. Biramous parapodia (like in Austrophyllum and Notophyllum ) constitute an ancestral state, which has evolved independently also within Phyllodoce . A new subfamily, Notophyllinae, is erected, and the two additional subfamilies, Phyllodocinae and Eteoninae, are redefined. Pareteoninae is treated as a junior synonym to Phyllodocinae, and Lugiinae as a nomen dubium. A number of genera are redefined and new combinations introduced. Based on type material Zverlinum and Phyllouschakovius are synonymized with Phyllodoce, Nipponophyllum with Notophyllym, Vitiaziphyllum with Eumida, Pareteone, Eulalia (Euphylla) and Compsanaitis with Paranaitis. More tentatively proposed synonymizations include Bergstroemia with Austrophyllum, Prophyllodoce with Phyllodoce , and Pseudeulalia with Protomystides. Eulalia lapsus is proposed as a replacement name for E. anoculata Fauchald, 1972, junior homonym to E. anoculata Hartman & Fauchald, 1971, and a number of species are synonymized and new combinations introduced. A checklist of phyllodocids together with information on the location of their type specimens is included.     

Eight microsatellite markers for sympatric alpine shrubs,Phyllodoce aleutica and P. caerulea (Ericaceae)

Phyllodoce aleutica and Phyllodoce caerulea are sympatric alpine‐snowbed plants in northern Japan. They compete for pollinators (bumblebees) each other and the competitive situation varies depending on snow conditions. We isolated and characterized eight microsatellite loci in these species. Additionally, one of 13 primers developed for Rhododendron metternichii was available in these species. The expected heterozygosity of these nine markers ranged from 0.06 to 0.93 in P. aleutica and from 0.09 to 0.96 in P. caerulea. These markers may be useful to reveal the mating system evolution, patterns of pollen flow and the process of natural hybridization in these Phyllodoce species.     

Variations in bumble bee preference and pollen limitation among neighboring populations: comparisons between Phyllodoce caerulea and Phyllodoce aleutica (Ericaceae) along snowmelt gradients

Two alpine-snowbed shrubs, Phyllodoce caerulea and P. aleutica (Ericaceae), co-occur in locales in northern Japan with early to late snowmelt, but they have different mating systems. Phyllodoce caerulea is an obligate outcrosser in any population, whereas the selfing ability of P. aleutica is highly variable among neighboring populations along snowmelt gradients: it shows high self-compatibility in early to middle snowmelt populations but low self-compatibility in late snowmelt populations. We investigated the relationships between pollinator availability and mating systems of these species along three snowmelt gradients. Relative abundance of flowers and nectar standing crop of P. caerulea decreased from early to late snowmelt plots. Bumble bees preferred P. caerulea to P. aleutica in early and middle snowmelt plots, while their preference shifted to P. aleutica in late snowmelt plots. Pollen limitation was severe in P. aleutica in early to middle snowmelt plots but it was severe in P. caerulea in late snowmelt plots. Seed-set success under natural conditions of P. aleutica was higher than that of P. caerulea in all plots. Thus, we infer that the selfing ability of P. aleutica under pollinator limitation acts as a reproductive assurance. We conclude that the interaction through pollination between the sympatric species is strong enough to cause a phenotypic change in mating system even within a local area.     

A chemotaxonomic survey of flavonoids and simple phenols in leaves of the Ericaceae

Leaves of 344 species of the Ericaceae and of 37 species in three related families have been surveyed for flavonoids and simple phenols. The yellow flavonol gossypetin was found in 158 ericaceous species and is of chemotaxonomic interest within the Rhododendroideae (where it occurs in 11 of 19 genera) and in the Vaccinioideae (in 3 of 22 genera). 5-0-methylflavonols appear in 81 species, again mainly in the Rhododendroideae (eight genera); whereas 3,5-di-O-methylquercetin, caryatin, is present only in Bejaria, Phyllodoce and Rhododendron. Dihydroflavonols. common in Rhododendron , show a sporadic distribution elsewhere in the family. Of the simpler phenols surveyed, gentisic acid is common but methyl salicylate is present mainly in Gaultheria. Hydroquinone appears in ten related genera, methylhydroquinone in three and orcinol in seven. The chemical evidence generally supports Stevens' recent classification of the family, particularly his inclusion of Epigaea in the Rhododendroideae and his transfer of Diplarche from the Diapensiaceae to the Ericaceae. There is also chemical evidence to support the separation of Calluna and Cassiope , the inclusion of the Aibutoideae within the Vaccinioideae and the retention of the Pyroloideae and Monotropoideae within the Ericaceae. In addition, chemistry indicates that Phyllodoce should be placed nearer to the Rhodoreae than in Stevens' system and that Cassiope and Harrimanella , placed together in the Cassiopeae by Stevens, are distinctly different.     

Pollination Ecology and Endemism of Pedicularis pulchella Pennell (Scrophulariaceae)

Abstract Pedicularis pulchella Pennell, endemic to sparsely vegetated areas of alpine tundra scree in southwestern Montana and northwestern Wyoming, was found to be roat-hemiparasitic and obligately dependent for its pollination upon bumblebees of three non-tundra species that forage primarily in an inverted position for pollen scraped from anthers concealed in the galea, This sternotribic pollination syndrome is considered related to the short (5–6mm) proboscides of pollinators on flowers with deep (12.4mm), nectariferous corolla tubes and favoring pollen as an insect attractant. Photographic and spectropholomelric analysis of the red-purple floral color included prominent blue corolla and ultraviolet calyx reflections corresponding to dominant colors of bumblebee visual spectra. Nectar analysis by thin-layer chromatography and refractomctry indicated, respectively, the presence of fructose and sucrose, and a sucrose-equivalent concentration of dissolved solids exceeding 50%; both coincide with the general pattern of the genus. Analysis of pollen from corbicular loads of P. pulchella pollinators indicated relatively low pollen-constancy of insects and substantial foraging association with Vaccinium in the montane-subalpine zone. By contrast, pollinators on P. cystopteridifolia and Phyllodoce glanduliflora in a diverse tundra plant community on rich alpine turf revealed broad pollinator spectra (7–8 species), high pollen-foraging constancy on Pedicularis, and high nectar-foraging frequency on Phyllodoce : The pollination ecology and endemism of Pedicularis pulchtella are interpreted in terms of resource sharing, spatial competition of plant species, and selective stress of the physical environment.     

Phylogeography and refugia of the Japanese endemic alpine plant, Phyllodoce nipponica Makino (Ericaceae)

Aim  This study aims to elucidate the phylogeography of the Japanese endemic alpine plant, Phyllodoce nipponica Makino (Ericaceae) and to infer the location of refugia of alpine plants in Japan during climatic oscillations.
Location  Alpine zone in the Japanese archipelago.
Methods  We determined the chloroplast (cp) DNA haplotypes of 155 individuals (22 populations) based on sequence data from the trnL-F and trnT-L intergenic spacers and the trnL intron, whose phylogenetic relationships were analysed using the program tcs . To examine the genetic structure, analysis of molecular variance ( amova ) was carried out and the population differentiation was shown by the parameters G _ST and N _ST.
Results  The haplotype composition and the results of amova showed that populations in the Japanese Central Mountain Region (JCMR) and in the westernmost region were highly divergent (18.8%). The diversity within populations was very high in the JCMR ( h _S = 0.421); less variation was found within populations located in other regions at lower elevations.
Main conclusions  Phyllodoce nipponica survived climatic changes during the Quaternary in the JCMR and the westernmost region. Most of the distribution range was colonized during only one range expansion. The source location from which the range expansion occurred was unclear.     

Novel, posterior sensory organ in the trochophore larva of Phyllodoce maculata (Polychaeta)

A new posterior sensory organ (PSO), located at the dorsal midline of the hyposphere, is described by immunocytochemical detection of acetylated alpha tubulin and serotonin (5-HT) in a laser-scanning microscope, as well as three-dimensional reconstructions after optical serial sectioning in the trochophore larva of the polychaete Phyllodoce maculata (Phyllodocidae). The unpaired PSO consists of five bipolar sensory cells, two of them being 5-HT immunopositive, which send axons to the cerebral ganglion and prototroch nerve. The dendrites of these cells project to the surface and bear one cilium each. A single neuronal fibre from the apical sensory organ innervates the PSO.     

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.     

东北杜鹃花科4属植物花粉形态的研究

利用光学显微镜和扫描电子显微镜对东北杜鹃花科甸杜属(Chamaedaphne Moench.)、杜香属(Ledum Linn.)、松毛翠属(Phyllodoce Salisb.)、越橘属(Vaccinium Linn.)4属6种, 1变种的花粉形态进行了观察和比较研究。以上4属花粉粒均为四合花粉。四合花粉呈正四面体排列。花粉粒近球形。每粒花粉均具3沟,在四合花粉粒上,相邻花粉粒上的沟相接,有些种可见沟内的萌发孔。光镜下可见花粉外壁为内外两层,厚度约相等。不同属种的花粉粒在大小上有区别,花粉粒平均直径为34.0~43.7μm。在扫描电镜下观察可见不同的种花粉表面具有不同的纹饰。为分类学提供了孢粉学方面的证据。     

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.     

Cladistic analysis of Enkianthus with notes on the early diversification of the Ericaceae

The genus Enkianthus (Ericaceae) has been subject to a cladistic analysis of morphological, anatomical, embryological, and cytological data, using two species of Clethra as outgroup. In order to evaluate the monophyly of Enkianthus , the genera Epigaea and Phyllodoce from the subfamily Rhododendroideae, and the genus Andromeda from the subfamily Vaccinioideae were also included in an initial analysis which yielded 20 equally parsimonious cladograms. The results indicated that Enkianthus is monophyletic, and a subsequent analysis with only the outgroup taxa and 14 species of Enkianthus yielded two equally parsimonious cladograms with a more resolved topology. In order to obtain strictly monophyletic taxa, the current subgeneric classification of Enkianthus is revised, recognizing four sections: Enkianthus (9 spp.), Andromedina (2 spp.), Meisteria (2 spp.), and Enkiantella (4 spp.). The paper provides a key, illustrations of representative taxa, a cladogram, and strict consensus trees.     

Sphinctocystis phyllodoces gen. n., sp. n. (Eugregarinida: Lecudinidae)--a new gregarine from Phyllodoce citrina (Polychaeta: Phyllodocidae)

A new species of aseptate gregarine, Sphinctocystis phyllodoces gen. n., sp. n., from the gut of a polychaete Phyllodoce citrina Malmgren, 1865 from White Sea is described. The electron and light microscopic data on trophozoits are presented. Taxonomy of the described species is discussed. Certain ultrastructural characters are included in generic and specific diagnoses. Order Eugregarinida Leger, 1900; suborder Aseptata Chakravarty, 1960; family Lecudinidae Kamm, 1922. GENUS: Sphinctocystis gen. n. TYPE SPECIES: Sphinctocystis phyllodoces sp. n. DIAGNOSIS: Characters of the family. Free trophozoits elongated, often with several annular constrictions. Anterior end asymmetric, without hooks, not separated from the body, with small apical papilla encircled by smooth area. Epicyte "classical", without additional axial formations at the tops of folds; epicytic folds high, monomorphic in cross sections, finger-shaped, with parallel sidewalls. In the gut of polychaetes. DIFFERENTIAL DIAGNOSIS: The new genus differs from Lecudina by having asymmetric anterior end, developed smooth area around the apical papilla, and monomorphic epicytic folds looking finger-shaped in cross sections. It also differs from Lankesteria by the absence of additional axial formations at the tops of the epicytic folds. It differs from both named genera by presence of annular constrictions on the trophozoit body. Sphinctocystis phyllodoces sp. n. DIAGNOSIS: Characters of the genus. Free trophozoits elongated, large, up to 617 x x 77 um. The average height of epicytic folds 976 nm, thickness 194 nm; there are 6-8 apical filaments and rippled dense structures per fold. Nucleus spherical (ellipsoid after fixation), 24-52 microm along longest axis, localised in anterior third of the body, carries several karyosomes of various size; 25-30 nm thick fibrils (possible fragments of nucleolonema) may be present in karyolymph. Other stages unknown. TYPE SERIES: Microscope preparation with 7 trophozoits, Karacci's haematoxylin stained, is kept in the Zoological museum of the Moscow State University (collection number: Z-1). TYPE HOST: Phyllodoce citrina Malmgren, 1865 (Polychaeta: Phyllodocidae). LOCALISATION: Mid-gut. TYPE LOCALITY: White Sea Biological Station of the Moscow State University, Yeremeyevsky Rapid, Velikaya Salma Strait, Kandalaksha Bay, White Sea.     

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     

A hybrid zone dominated by fertile F1s of two alpine shrub species, Phyllodoce caerulea and Phyllodoce aleutica, along a snowmelt gradient

In alpine ecosystems, the steep environmental gradients produced by the difference in snowmelt timing create a dynamic selective regime for alpine plants. As these gradients directly alter flowering phenology, they can affect pollen-mediated gene flow among populations of single and related species. In northern Japan, we found a hybrid zone dominated by fertile F(1)s of two alpine shrub species, Phyllodoce caerulea and P. aleutica, along a snowmelt gradient. Seed germination confirmed the fertility of F(1) hybrid, making the rarity and absence of backcross and F(2) plants puzzling. The long-term clonal perpetuation of F(1) hybrids (at least a few thousand years ago) contributes the maintenance of this unique hybrid zone. The distribution patterns of chloroplast DNA haplotypes suggest that F(1) formation might be caused by directional pollen flow between parental species along the snowmelt gradient. Based on these results, we discuss the ecological and evolutionary significance of this unique hybrid zone.     

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     

Suivi pluriannuel du peuplement à Abra alba dans la partie méridionale de la Mer du Nord (région de Dunkerque-France)

The Abra alba community is considered as a faunistical unity well established on coastal areas of North Sea. This community shows, since several years, very important recruitments of certain species. Since 1978, recruitments of Abra alba, Tellina fabula, Phyllodoce (Anaïtides) mucosa, Lagis koreni or Lanice conchilega have been observed. However these demographic phenomena does not affect the structure of the community, which shows, through rank-frequency diagrams, the characteristic of a mature population, principally constituted of 7 species.An annual cycle study shows a succession of recruitments of several species. A correspondance factorial analysis was realised from grain size datas as well as abundances of 18 main species. This analysis shows an annual cycle which seems to be linked to the clay part. Maximal values of clay part appears in summer and minimal values at the end of winter or early springtime.The hydrological parameters studied, allowed to make the difference between external factors (continental contributions, phytoplanctonic activity etc...) and benthic activity.

     

Snowbeds trapping seed rain—a comparison of methods

The possibility of assessing seed rain by deposits in snowbeds was investigated in the Abisko area in northernmost Swedish Lapland. In late summer the "grey blanket" on the snowbed surface was skimmed off, melted and sieved through a filter paper. The debris contained on average 741 seeds m², making up 6.5% of the total debris dry mass. A total of 11,909 seeds, representing 69 species, were encountered in the study. Phyllodoce coerulea, Carex bigelowii and Juncus trifldus were the most abundant. The efficiency of the snowbeds as diaspore traps was compared with a set of artificial traps. Data from two years were compared. The snowbed proved to be more efficient than artificial seed traps, in particular for trapping seeds of graminoids and herbs. Both methods revealed considerable among-year variation in seed output as a possible result of differences in summer climate. The results also suggest that snowbed can be used for assessing aeolian inorganic deposition, in the present study on average 9.0 g m².