PERIPHYTON COLONIZATION OF ROCK SURFACES IN A BOREAL FOREST STREAM STUDIED BY SCANNING ELECTRON MICROSCOPY AND TRACK AUTORADIOGRAPHY1

Periphyton colonization of natural rock surfaces and granite tiles was followed experimentally in the Matamek River, an acidic (pH 5.5) sixth order boreal stream in northeast Quebec, Canada. Accumulation of chlorophyll a and freshweight algal biomass was logarithmic over a 25 day colonization period. The major colonizers were Tabellaria flocculosa (Roth) Kütz., T. fenestrata (Lyngb.) Kütz., Mougeotia sp., and Eunotia pectinalis (Kütz) Rabh., and its varieties. The microcolonization sequence on granite tiles, followed over 27 days with scanning electron microscopy, showed an initial accumulation of algal cells on the upstream and downstream margins and in depressions, followed by a gradual filling-in of the flat surfaces. It is hypothesized that the observed slow rate of colonization was due to the high surface tension of the granite substratum and the absence of preconditioning by an organic film. It is further hypothesized that the increase in cellular carbon fixation rates of T. flocculosa measured over a 23 day period using nuclear track autoradiography parallels the development of an algal-detrital microcosm on the granite tile, and is evidence for the establishment of localized tight nutrient spiralling. It is suggested that the periphyton community may be regarded as a nutrient recycling system at a microenvironmental level which may be especially important in oligotrophic systems.     

POLYMORPHISM OF THE DIATOM PINNULARIA BREBISSONII IN CULTURE AND A FIELD COLLECTION1,2

Two clones of Pinnularia brebissonii (Kütz.) Rabh. var. brebissonii were established and maintained in logarithmic phase of growth. Initial length of the cells was 37 μm. As cell division occurred, the mean length of cells in each population decreased as predicted by the MacDonald-Pfitzer hypothesis; however, the decrease in mean length was not uniform throughout the growth period. This nonuniformity is probably caused by nonrandom division of cells in the population or by a changing increment of size reduction due to division. The initial increment of size reduction was calculated as 0.7 μm/division. The smallest, cells observed were 8 μm long. As cells decrease in length, cell volume decreases and the proportion of cells with aberrant valve structure increases. More than 90% of the valves were abnormal in a population with mean length of 14 μm. The abnormalities of structure involved the raphe, the central area and the striae.     

PHYSIOLOGICAL MODIFICATIONS RELATED TO DENSITY INCREASE IN PERIPHYTIC ASSEMBLAGES 1

Periphytic communities in running waters were examined as they developed on granite rocks, concrete balls and glass slides. At equivalent cell densities, no differences in pigment concentrations, species diversity or production levels were found among the different substrata examined. Development of the assemblage appeared to result from the elongation of short algal filaments which had initially settled on the surface. As these communities matured, a distinct canopy and understory developed. Cellular metabolisms were comparable among the communities. In the understory of the communities, even though the cellular content of chl a and b did not differ, chl c and carotenoid pigment concentrations were higher than those in the over-story. Bicarbonate assimilation of Tabellaria fenestrata (Lyng.) Külz. and Eunotia pectinalisi var. pectinalis (O. F. Müll?) Rabh. was higher than that of the more abundant Tabellaria flocculosa (Roth.)Kütz. var. flocculosa IV (sensu Koppen) at both high and low cell densities. This probably reflects a seasonal succession of colonizing species. Glucose assimilation appeared to be mainly attributable to bacterial activity, and algal cells of the upper layer were less active than those of the bottom. The small amount of glucose that was incorporated by the algal cells was probably absorbed passively since its amount was in direct proportion to cell volumes.     

VALVE AND BAND MORPHOLOGY OF SOME FRESHWATER DIATOMS I. FRAGILARIA CAPUCINA VAR. MESOLEPTA1,2

Acid cleaned cells from clonal cultures of Fragilaria capucina var. mesolepta Rabh. were examined with light and scanning electron microscopy. Recently isolated cells are linear-lanceolate in shape with a median constriction. After several transfers over 25 mo, cells exhibit size diminution resulting in small elliptically shaped valves. Adjacent valves are united to one another by interlocking marginal spines. Every valve has an apical pore field at each apex. A single labiate process is present infrequtently, appearing underdeveloped most often in size-reduced cells. The girdle region consists of two cingula, each composed of a series of underlapping bands. Each pleura in the series is a discontinuous ring with a central ligula. A survey of past ultrastructure studies on the freshwater Fragilariaceae reveals that the occurrence of the apical pore field and labiate process are likely key characteristics for the family. The apical pore field of Diatoma, Asterionella and Tabellaria is positioned on the valve face, whereas the apical pore field of F. capucina var. mesolepta is located on the valve mantle, the girdle region of F. capucina var. mesolepta is basically similar to that of Gomphonema parvulum (Kütz.) Grun.     

OCCURRENCE OF THREE NITZSCHIA (BACILLARIOPHYCEAE) TAXA WITHIN COLONIES OF TUBE-FORMING DIATOMS1

In benthic samples from the unchannelized Missouri River, the diatoms Nitzschia dissipata (Kütz.) Grun., N. filiformis (W.Sm.) Schütt and N. pseudofonticola Hust. were observed within the mucilage tubes of four tube-forming diatoms: Cymbella prostrata (Berk.) Cl., C. prostrata var. auerswaldii (Rabh.) Reim., Navicula tripunctata var. schizonemoides (V.H.) Patr., and Nitzschia filiformis. Microscopical observations of live and preserved specimens indicated that “invasion” by Nitzschia occurred primarily in older tubes. Data are presented on the environmental conditions in which the tube-formers and their cohabitants have been found.     

COMMUNITY STRUCTURE DYNAMICS OF EPILITHIC AND EPIPHYTIC DIATOMS IN A MISSISSIPPI STREAM1

Epilithic and epiphytic diatoms were collected monthly from 18 June 1979 through 19 May 1980 from both shaded and unshaded sites of Sessums Creek, a shallow, oligotrophic and strongly alkaline stream in northeastern Mississippi. The dominant macroalgae were filamentous green algae belonging to the non-mucilage producing genera Cladophora and Rhizoclonium. Ninety taxa were identified with Navicula minima Grun. and Nitzschia dissipata (Kütz.) Grun. accounting for 50% of all valves counted. Other important taxa were Navicula menisculus var. upsaliensis (Grun.) Grun., N. cryptocephala var. veneta (Kütz.) Rabh., Achnanthes minutissima Kütz. and Cymbella turgida (Greg.) Cl. With few exceptions, the more dominant taxa were equally abundant in the shaded and unshaded sites and also in the epilithon and epiphyton. Species diversity (H') and the number of taxa in a sample in all four habitats showed an identical pattern, being lowest in winter and highest in fall. No one pair of habitats was more structurally similar than any-other pair when compared by a similarity index (SIMI). Apparently, differences in light intensity and substrate type had little effect on diatom distribution in Sessums Creek and it was concluded that the locality sampled supported a single, nearly homogeneous diatom community.     

Preliminary results of a classification of fifty-one selected northeastern Wisconsin Lakes (USA) using indicator diatom species

Diatom species lists were generated for 51 lakes in northeastern Wisconsin, and then classified by two-way indicator species analysis (TWINSPAN). The lakes were initially divided into two main groupings: Group I lakes which were alkaline lakes of moderate to high productivity, and Group II lakes which were acid lakes of low productivity. Group I lakes were further divided into two subgroupings, and four levels. In total, twelve indicator diatom species were recognized during the TWINSPAN lake classification, these were: Achnanthes exigua Grun., A. lanceolata var. dubia Grun., Amphora ovalis var. Affinis (Kütz.) V.H. ex Det., A. perpusilla (Grun.) Grun., Cymbella cistula (Ehrenb.) Kirch., Diploneis elliptica (Kütz.) Cl., Eunotia pectinalis (O.F. Mull.) Rabh., Fragilaria vaucheriae (Kütz.) Peters., Frustulia rhomboides var. crassinervia (Bréb. ex W. Sm.) Ross, Navicula capitata Ehrenb., N. decussis Østr., and N. scutelloides W.Sm. ex Gregory.     

Identification of Naemacyclus minor hyphae within needle tissues of Pinus sylvestris by immunoelectron microscopy

Cultural investigations revealed that Naemacyclus minor, Lophodermium seditiosum and Cenangium ferruginosum were the most frequent colonizers of asymptomatic and symptomatic Pinus sylvestris needles. Since ultrastructural observations showed that morphological features were not suitable to differentiate hyphae of N. minor from hyphae of other isolates, the on-section immunogold labelling technique was applied in combination with an anti-N. minor specific immunoserum. The specificity of this serum was tested against culture hyphae of all isolates. Anti-N. minor specific immunoserum was then used to identify N. minor hyphae in thin sections of green P. sylvestris needles. The infection loci identified were restricted to small tissue areas located in the vicinity of stomata. In the hypodermis, hyphae and endocell-containing hyphae were located within the lumina of host cells but outside the protoplast. The growth of hyphae from cell to cell occurred through pits. The hyphae spreat into the mesophyll intercellularly and continued with the intracellular colonization of moribund and dead mesophyll cells in a later stage of infection. The observed host-parasite interactions at cellular and ultrastructural level are discussed in connection with the still controversial interpretation of the pathogenicity of N. minor.     

SEXUAL DIMORPHISM IN RELATION TO WINTER FORAGING AND TERRITORIAL BEHAVIOUR OF THE THREE-TOED WOODPECKER PICOIDES TRIDACTYLUS AND THREE DENDROCOPOS SPECIES

A comparison of morphological characters shows that Picoides tridactylus is more markedly sexually dimorphic than Dendrocopus leucotos, D. major or D. minor. This is correlated with intersexual differences in foraging behaviour in P. tridactylus, lacking in the Dendrocopus spp. Apparently as a consequence, the sexes are able to share winter territories among P. tridactylus but not among D. major or D. minor; conclusive observations of D. leucotos were not possible. It is suggested that these differences may reflect the distributional history of the species, and the extent of interspecific competition.     

棉铃虫蛹和成虫形态特征的数值分析

通过对棉铃虫Helicoverpa armigera(Hubner)及其近缘种的蛹和成虫进行了形态特征变异的数值分析,结果表明：棉铃虫种群内个体变异较大,但种群间相对分化较小,无明显的地理分化,也没有发现棉铃虫种群有稳定变异的地理特征;但发现雄性抱器冠的长、宽、抱器瓣长宽比及抱器瓣长与冠宽比4个特征与纬度有一定的相关性,其相关系数分别为：-0.612,-0.699,0.693和0.691。尽管如此,棉铃虫还是相对比较单一的一个种,而近缘种在阳茎刺和抱器瓣的形态及阳茎端囊上的大刺数等方面有明显而稳定的差异和间断。     

Characteristics of softwater streams in Rhode Island II. Composition and seasonal dynamics of macroalgal communities

Forty stream segments in Rhode Island, U.S.A., were examined seasonally from June 1979 to March 1982. Thirty-nine species of macroalgae were collected, respresenting 25 genera. The composition of the lotic flora was 54% green algae, 31% red algae, 5% blue-green algae, 5% xanthophytes, 3% chrysophytes and 3% diatoms. The majority of these taxa (85%) were filamentous. From a biweekly examination of five stream segments, macroalgal communities could be grouped according to light regime. Species in unshaded streams exhibited little seasonality, whereas in streams shaded by one or more layers of riparian canopy, maxima in species numbers and abundance occurred during colder seasons. The most widespread and abundant species were the blue-green alga Phormidium retzii, the green alga Draparnaldia acuta, and the diatom Eunotia pectinalis. P. retzii and E. pectinalis were aseasonal annuals, while D. acuta was primarily a winter-spring form. It appears that pH is a major factor affecting broad geographic distribution patterns of stream macroalgae, whereas the light regime established by overhanging canopy is an important factor which influences localized abundance and seasonality of lotic macroalgal communities. Niche pre-emption appears to be a common mode of resource space division among stream macroalgae in Rhode Island. E. pectinalis is the strongly developed dominant in this drainage system.     

SCANNING ELECTRON MICROSCOPY OBSERVATIONS OF DEFORMITIES IN SMALL PENNATE DIATOMS EXPOSED TO HIGH CADMIUM CONCENTRATIONS1

Different types of malformations are likely to affect the morphology of diatoms when exposed to particularly unstable environmental conditions, the most easily identifiable being distortion of the whole frustule. In the present study, we investigated, by means of SEM, valve abnormalities induced by high cadmium contamination (100 μg · L^?1) in small pennate diatoms. Changes in the shape of Amphora pediculus (Kütz.) Grunow and anomalous sculpturing of the cell wall of many species, such as Encyonema minutum (Hilse) D. G. Mann, Mayamaea agrestris (Hust.) Lange‐Bert., Gomphonema parvulum (Kütz.) Kütz., or Eolimna minima (Grunow) Lange‐Bert., were observed, which were not, or almost not, noticeable in the LM. With consideration to current knowledge of diatom morphogenesis, metal uptake by the cell would induce, directly or indirectly, damage to many cytoplasmic components (e.g., microtubules, cytoskeleton, Golgi‐derived vesicles) involved in the precisely organized silica deposition. This study confirms that many species, whatever their size, are likely to exhibit morphological abnormalities under cadmium stress, and that this indicator may be valuable for the biomonitoring of metal contamination, even if SEM observations are not necessary for routine studies.     

Integrated analyses of chromosome, molecular and morphological variability in the Andean mice Eligmodontia puerulus and E. moreni (Rodentia, Cricetidae, Sigmodontinae)

Patterns of evolution and systematics of sigmodontine rodents are matters of continuous revision and debate. The silky mouse, Eligmodontia, is a phyllotine rodent adapted to arid environments. Chromosomal and molecular data have identified six species in this genus. Among these E. puerulus and E. moreni are sister taxa from the high Andean and lowland deserts, respectively, with large chromosomal differences and intermediate levels of molecular divergence. The purpose of our study was to quantify the degree of variability (morphological, cytogenetic, and molecular) and to analyze its evolutionary implications within, and between, these sister species in the Monte and Puna biomes of Argentina. Our results show a high variability at the chromosomal and molecular level, but low morphological differentiation among populations of E. puerulus. Diploid numbers vary from 31 to 37 due to a complex Robertsonian system, whereas cytochrome-b distances range from 0.15% to 5.75%. On the other hand, E. moreni shows high morphological differentiation between populations, but low intraspecific differentiation at the molecular (from 0.73% to 1.4%) and chromosomal level (2n = 52). Comparison of E. puerulus with E. moreni reveals high morphological and chromosomal distinction between them, but absence of molecular differentiation. Our results suggest that: (1) the high genetic variability of E. puerulus could be associated to its geographic distribution in the complex topography of the high Andean Puna; (2) the high morphological differentiation between E. moreni and E. puerulus could be the result of natural selection; and (3) molecular polyphyly between E. puerulus and E. moreni could be due to introgression or a recent split of these taxa. Finally, our results emphasize the need to integrate different datasets in the analysis of species variability and diversification, as tools for understanding their evolutionary histories.     

STUDIES IN PENNATE DIATOMS: VALVE MORPHOLOGIES OF LICMOPHORA AND CAMPYLOSTYLUS1

Using scanning electron and light, microscopy, several hundred specimens of Campylostylus normanianus (Grev.) Gerloff, Licmophora abbreviata Agardh, L. gracilis var. anglica (Kütz) Per. et Per., and L. flabellata (Carm.) Agardh were examined to elucidate their valve morphology. These species were found to be heterovalvar with respect to the presence of the labiate process in the basal apices of the cell, although one was always present at the head pole of each valve. This form of heterovalvar may be one basis for partitioning the large and variable family Diatomaceae. Because of the similar valve morphologies exhibited by C. normanianus and Licmophora species examined, it is recommended that Campylostylus normanianus be placed in the older genus Licmophora, as L. normaniana (Grev.) Wahrer.     

SPECIES BOUNDARIES IN THE TOXIC DINOFLAGELLATE PROROCENTRUM LIMA (DINOPHYCEAE,PROROCENTRALES), BASED ON MORPHOLOGICAL AND PHYLOGENETIC CHARACTERS1

Wild and cultured specimens of Prorocentrum lima (Ehrenb.) F. Stein from 26 widely different areas in 13 countries were examined in order to determine consistent characters for delimiting species boundaries in this taxon. The morphological characters valve shape, valve size, valve ornamentation, number and shape of valve pores, number and shape of marginal pores, and periflagellar platelets were observed using LM and SEM, and two molecular genetic regions were sequenced. We identified stable morphological characters that were consistent among wild specimens and all cultures, which were valve shape, valve ornamentation, and number and arrangement of periflagellar platelets. All cultures of P. lima identified by these characters formed a monophyletic group in phylogenetic analyses based on the two genes, which, however, included the species Prorocentrum arenarium. P. arenarium was determined to be within the range of morphological variation of P. lima, and therefore we synonymize the two taxa. Within this monophyletic group, P. lima was divided into several subclades in the all phylogenetic analyses. There were no morphological characters specifically related to any one subclade. The subclades appeared to correlate broadly to sample collection regions, suggesting that geographically separated populations may have become genetically distinct within this epi‐benthic species. We have emended species boundaries in P. lima.     

DEFINING TAXON BOUNDARIES IN MEMBERS OF THE MORPHOLOGICALLY AND GENETICALLY PLASTIC GENES CAULERPA (CAULERPALES,CHLOROPHYTA)1

Numerous attempts to capture the morphological variability of the genus Caulerpa have resulted in an unstable classification of numerous varieties and formae. In the present study we attempted to test taxon boundaries by investigating morphological and genetic variation within and between seven taxa of Caulerpa, supposedly belonging to four species, sampled at different sites in a Philippine reef system. Using both field and culture observations, we described the relation between the variability of a set of morphological characters and ecological parameters, such as wave exposure, light intensity, and substrate type. Statistical analyses showed that the limits between two (out of three) ecads of the C. racemosa (Forsskål) J. Agardh complex were obscured by the presence of morphological plasticity. Other studied taxa of Caulerpa (i.e. C. cupressoides [Vahl] C. Agardh, C. serrulata [Forsskål] J. Agardh, and two formae of C. sertularioides [S. Gmelin] Howe) could be grouped based on morphology despite the presence of morphological plasticity. Our results indicated a strong association between light intensity and several quantitative morphological variables. Genetic diversity of these taxa was assessed by partial sequencing chloroplast rbcL and tufA genes and the ycf10‐chlB chloroplast spacer. In all phylogenetic analyses, C. serrulata, C. cupressoides, C. sertularioides, and the three ecads of C. racemosa emerged as distinct genetic units. Despite the presence of morphological plasticity and morphological convergence, a subset of morphological characters traditionally used in taxonomic delimitation still had sufficient discriminative power to recognize the terminal phylogenetic clades.     

LIPID BIOMARKER DIVERSITY IN THE COCCOLITHOPHORID EMILIANIA HUXLEYI (PRYMNESIOPHYCEAE) AND THE RELATED SPECIES GEPHYROCAPSA OCEANICA1

Twenty-four strains of Emiliania huxleyi and two strains of Gephyrocapsa oceanica were grown at 15°C under identical culture conditions to assess genetic variability in key lipid biomarker profiles (C₃₇-C₃₉ alkenones, C₃₆ and C₃₇ alkyl alkenoates, and C₃₁-C₃₈ alkenes). Under our culture conditions, little divergence an biomarker composition was detected between E. huxleyi strains from different oceanic regions or between E. huxleyi and G. oceanica even though the strains originated from biogeographical regions as diverse as the subpolar North Atlantic and subtropical Western Pacific. The major differences observed were in tetraunsaturated alkenone abundance and alkene profiles, which tended to separate neritic from open ocean strains. Different strains from the same locality were as different as strains originating from widely separated ocean basins, indicating extreme genotypic diversity within a population. Replicate cultures of the same strain showed significant variability in their biomarker profiles even though the culture temperature varied by only ±0.3°C, indicating that their synthesis ratios are influenced by environmental and/or physiological variable(s), as yet unidentified, in addition to temperature. Strong covariance in C₃₇ and C₃₈ methyl alkenone unsaturation ratios (U^k₃₇ and U^k_38Mρ respectively) and, in coastal strains, C₃₃, alkene and alkenone unsaturation ratios indicates that these compounds are biochemically linked.     

The great tit (Parus major) – a misclassified ring species

The great tit (Parus major) has been considered to be the most typical example of an avian ring species. The terminal taxa of the ring (major and minor sectors) are supposed to be reproductively isolated in a zone of secondary contact in the middle Amur valley, Siberia. Our study combines molecular markers (cytochrome‐b), bioacoustic analyses and morphological characters to judge the ring species status of the great tit complex. Despite a notable percentage of intermediately coloured birds in the mixed population of middle Amur, a lack of mitochondrial introgression between the major and minor sectors and a small number of true hybrids among voucher specimens from this area suggest at least a partial reproductive barrier between both sectors. In contrast, variation of morphological and especially acoustic characters along the ring‐shaped area and the phylogenetic structure of the P. major group do not match the ring species concept. Bioacoustic and molecular data (cytochrome‐b sequences) reveal two large and closely related subspecies blocks, the sectors major and bokharensis in the Western Palaearctic and central Asia, and the sectors minor and cinereus in the Eastern Palaearctic and South‐east Asia, respectively. The two western sectors diverged only recently (0.5 Mya) and they were separated from the eastern group by Pleistocene events about 1.5 Mya. Songs from allopatric regions of the two subspecies blocks differ distinctly in frequency parameters and element composition. In the area of secondary contact, males of all phenotypes share the same frequency range of song, close to the range of the typical minor song. Hybrids and major males sing mixed repertoires of typical major and minor strophe types as well as mixed strophes. In contrast, phenotypic minor males display only pure minor strophes. Differences in mate choice and mating success based on repertoire size are believed to uphold the reproductive barrier between major and minor birds in the area of sympatry. Taxonomic consequences suggest three separate species in the Parus major complex: Parus major s.s. (including the very closely related bokharensis sector), Parus minor and Parus cinereus. © 2005 The Linnean Society of London, Biological Journal of the Linnean Society, 2005, 86 , 153–174.     

Loxocaudinae: A new subfamily in the ostracod family loxoconchidae

A new Ostracoda subfamily, Loxocaudinae subfam. nov., separated from the subfamily Loxoconchinae of the family Loxoconchidae with five genera (Loxocauda Schornikov, 1969, Glacioloxoconcha Hartmann, 1990, Phlyctocythere Keij, 1958, Pseudoloxoconcha Müller, 1894 and Sarmatina Stancheva, 1984) is described in the paper. The new subfamily differs from Loxoconchinae Sars, 1926 s. str. by the presence of a compact eye, the absence of an eye tubercle, an originally adont hinge with a tendency to formation of an anterior tooth on the left valve and a posterior one on the right valve, a pronounced caudal process and reduction of fossa-mural sculpture. The volume of the genera that are included in the subfamily is determined. A review of 55 various species and forms mentioned in the literature in open nomenclature, whose taxonomic position has not been determined yet is presented. A total of 45 species close to Loxocauda are attributed to the new subfamily, but proper morphological investigations are necessary for their classification. Ten species that were referred earlier to the genera included in Loxocaudinae are excluded from this subfamily. Issues of functional morphology of Loxocaudinae shells and morphological evolution of their sculpture are considered.