Spread of Elodea canadensis in Lake Baikal

Elodea canadensis Michx. was first observed in Lake Baikal at the end of the 70's. E. canadensis was probably introduced to Baikal by aquarists or by transport vehicles and transport together with fishes taken from lakes of the Urals, where mass development of the plant was observed at the beginning of the present century. The spread of E. canadensis along lake Baikal may be due to navigation, but natural factors also affect its spread.The fast spread of E. canadensis, its high competitive quality and metabolic properties for purifying sewage waters are of interest.The expansion of E. canadensis may be considered a catastrophic phenomenon; its effect now covers practically the whole perimeter of Lake Baikal.     

Dynamics of submerged macrophyte populations in response to biomanipulation

1. A 6‐year study (1992–97) of changes in submerged vegetation after biomanipulation was carried out in the eutrophicated Lake Finjasjön, Southern Sweden. Ten sites around the lake were revisited each year. At each site five samples of above‐ground biomass were taken at 10 cm water depth intervals. An investigation of the seed bank at the 10 sites, and a grazing experiment where birds and large fish were excluded was also conducted. 2. Between 1992 and 1996, in shallow areas (water depth < 3 m), vegetation cover increased from < 3 to 75% and above‐ground biomass from < 1 to 100 g DW m^–2. Mean outer water depth increased from 0.3 to 2.5 m. Elodea canadensis and Myriophyllum spicatum accounted for > 95% of the increase in biomass and plant cover. The following year (1997), however, cover and above‐ground biomass decreased, mainly attributable to the total disappearance of E. canadensis. Secchi depth increased after biomanipulation until 1996, but decreased again in 1997. 3. Total and mean number of submerged species increased after biomanipulation, probably as a result of the improved light climate. However, after the initial increase in species number there was a decrease during the following years, possibly attributed to competition from the rapidly expanding E. canadensis and M. spicatum. The lack of increase in species number after the disappearance of E. canadensis in 1997 implies that other factors also affected species richness. 4. A viable seed bank was not necessary for a rapid recolonization of submerged macrophytes, nor did grazing by waterfowl or fish delay the re‐colonization of submerged macrophytes. 5. Submerged macrophytes are capable of rapid recolonization if conditions improve, even in large lakes such as Finjasjön (11 km²). Species that spread by fragments will increase rapidly and probably outcompete other species. 6. The results indicate that after the initial Secchi depth increase, probably caused by high zooplankton densities, submerged vegetation further improved the light climate. The decrease in macrophyte biomass in 1997 may have caused the observed increase in phosphorus and chlorophyll a, and the decrease in Secchi depth. We suggest that nutrient competition from periphyton, attached to the macrophytes, may be an important factor in limiting phytoplankton production, although other factors (e.g. zooplankton grazing) are also of importance, especially as triggers for the shift to a clear‐water state.     

Phenotypic plasticity as a clue for invasion success of the submerged aquatic plant Elodea nuttallii

• Two closely related alien submerged aquatic plants were introduced into Europe. The new invader (Elodea nuttallii) gradually displaced E. canadensis even at sites where the latter was well established. The aim of the study was to evaluate the combined effects of environmental factors on several phenotypic characteristics of the two Elodea species, and to relate these phenotypic characteristics to the invasion success of E. nuttallii over E. canadensis.
• In a factorial design, Elodea plants were grown in aquaria containing five different nitrogen concentrations and incubated at five different light intensities. We used six functional traits (apical shoot RGR), total shoot RGR, relative elongation, root length, lateral spread, branching degree) to measure the environmental response of the species. We calculated plasticity indices to express the phenotypic differences between species.
• Light and nitrogen jointly triggered the development of phenotypic characteristics that make E. nuttallii a more successful invader in eutrophic waters than E. canadensis. The stronger invader showed a wider range of phenotypic plasticity. The apical elongation was the main difference between the two species, with E. nuttallii being more than two times longer than E. canadensis. E. canadensis formed dense side shoots even under high shade and low nitrogen levels, whereas E. nuttallii required higher light and nitrogen levels.
• We found that under more eutrophic conditions, E. nuttallii reach the water surface sooner than E. canadensis and through intensive branching outcompetes all other plants including E. canadensis. Our findings support the theory that more successful invaders have wider phenotypic plasticity.

     

Establishment of the Baikalian endemic amphipod Gmelinoides fasciatus Stebb. in Lake Ladoga

Gmelinoides fasciatus Stebb., a small amphipod from the Lake Baikal basin, was discovered in July 1988 in Lake Ladoga, the largest European lake. G. fasciatus likely invaded Lake Ladoga as a consequence of its intentional introduction, aimed at enhancing fish production, in some Karelian Isthmus lakes close to Lake Ladoga's western shore in the early 1970's. Benthos studies conducted in 1989 and 1990 revealed that G. fasciatus was well established in littoral communities along the western and northern shores of Lake Ladoga. G. fasciatus was the dominant species in these littoral communities and contributed over 70% of the macroinvertebrate biomass. The species was abundant in different macrophyte beds and stony littoral areas, both in heavily polluted and undisturbed sites. The maximum abundance and biomass of G. fasciatus was about 54,000 ind m^–2 and 160 g (wet wt.) m^–2. Negative impacts of G. fasciatus on native species have been observed. Studies are needed to identify the effects of G. fasciatus on the functioning of littoral communities and to predict and control the spread of this amphipod.     

Investigation of distribution,species composition,and degree of resistance to antibiotics of the bacteria of the <Emphasis Type="Italic">Enterococcus</Emphasis> genus in Lake Baikal

The paper presents the result of investigation of species composition, distribution and degree of resistance to antibiotics of the Enterococcus bacteria in Lake Baikal. It is shown that they live in littoral zones of the lake. In the deep-water part of the lake the studied microorganisms are not found. We isolated 120 strains identified as E. faecium, E. avium, E. faecalis, E. mundii, E. hirae, E. durans, E. gallinarum. On the whole, the strains of enterococcus isolated from Lake Baikal are characterized as antibiotic-sensitive. Nevertheless, the antibiotic-resistant strains of enterococci are found in the studied collection, including those with typical intermediate level of resistance to vancomycin.     

GENOME RELATIONS AMONG ELYMUS CANADENSIS,ELYMUS TRITICOIDES,ELYMUS DASYSTACHYS,AND AGROPYRON SMITHII

Hand-emasculated Elymus canadensis pollinated by E. triticoides, E. dasystachys, and Agropyron smithii yielded 15, 21, and 1 viable hybrid seeds from 56, 52, and 52 florets, respectively. The 28-chromosome species—E. canadensis, E. triticoides, and E. dasystachys—behaved meiotically as allotetraploids and consistently formed 14 bivalents at metaphase I. Octoploid A. smithii, 2n = 56, averaged 0.41^I, 27.72^II, and 0.03^IV in 87 metaphase-I cells. Agropyron smithii is apparently an allooctoploid or a segmental autoallooctoploid. Meiosis was similar in the E. canadensis X E. triticoides and E. dasystachys hybrids. Chromosome pairing was very low in both hybrids, about two loosely connected open-ended bivalents per cell, and may not represent genuine homologies. The genomes of E. canadensis are distinctly different from those of E. triticoides and E. dasystachys. The E. canadensis X A. smithii hybrid averaged 13.37^I and 14.31^II in 76 metaphase-I cells. More than half of the bivalents were closed at both ends. Inability to distinguish between auto- and allosyndesis resulted in two interpretations of genome relations. Either A. smithii is an alloploid with two of its four genomes similar to those of E. canadensis, or it is a segmental autoalloploid genomically unrelated to E. canadensis. The first interpretation is favored. Agropyron dasystachyum, or one of its close relatives, and E. triticoides are suggested as possible parents of A. smithii.     

中国淡水硅藻新记录——偏肿内丝藻兴凯变种

于2016年5月对浙江省仙居国家公园进行藻类多样性调查时,发现了中国淡水硅藻的1个新记录种——偏肿内丝藻兴凯变种[Encyonema ventricosum var.hankensis(Skvortzov) RodionovaPomazkina]。该文采用光镜和扫描电镜对偏肿内丝藻兴凯变种浙江种群的形态特征进行了观察和描述,并将其与该种的其他地理种群及内丝藻属其他相似物种的形态特征进行比较,同时讨论了该物种的生境特征。结果表明:(1)偏肿内丝藻兴凯变种壳面均具明显背腹之分,背缘明显拱起,腹缘轻微波曲;端部呈头状至喙状。(2)偏肿内丝藻兴凯变种浙江地理种群在背部中央区域具一孤点,而其兴凯种群对孤点这一特征未有描述。(3)偏肿内丝藻兴凯变种在俄罗斯的兴凯湖和贝加尔湖等地有分布,该种在中国是首次报道。     

Reproductive allocation of Solidago canadensis L. plays a key role in its invasiveness across a gradient of invasion degrees

Reproductive allocation (RA) plays a vital role in the development of ecological strategies during the life cycle of plant species. Invasive alien plants (IAP) may exist at various invasion degrees across a gradient of the colonization process with several grades of relative abundances in the occupied environments. The progressive variation in the invasion degree of IAP has the potential to modify their RA strategy. This study purposes of estimating the RA strategy of the IAP Solidago canadensis L. and the correlations among RA of S. canadensis, the invasion intensity of S. canadensis, the invasiveness of S. canadensis, and the community invasibility across a gradient of invasion degrees by using the field sampling experiment. The height and relative abundance of S. canadensis did not remarkably affect its RA. The RA of S. canadensis was positively related to its reproductive biomass and total biomass. The key reason may be that plant individuals with higher total biomass can allocate more resources into sexual reproduction. The RA of S. canadensis was positively related to its invasiveness. Thus, the RA of S. canadensis may be crucial to its invasiveness.     

SYNTHETIC HYBRIDS OF HORDEUM BOGDANII WITH ELYMUS CANADENSIS AND SITANION HYSTRIX

Seven viable hybrid seeds were obtained from 48 hand-emasculated Elymus canadensis L., 2n = 28, florets pollinated by Hordeum bogdanii Wilensky, 2n = 14. The hybrids were large, vigorous, and completely sterile plants that bore a closer morphological resemblance to E. canadensis than to H. bogdanii. Chromosome associations in the 21-chromosome hybrids averaged 9.98^I, 5.40^II, and 0.08^III in 264 metaphase-I cells. Chromosome pairing was attributed to allosyndetic pairing between E. canadensis and H. bogdanii chromosomes. The H. bogdanii genome appears to be partially homologous with one of the two E. canadensis genomes. One Sitanion hystrix (Nutt.) J. G. Smith X H. bogdanii hybrid was obtained from a cross involving 37 emasculated S. hystrix florets. This triploid, 2n = 21, hybrid was morphologically intermediate between the parents and totally sterile. Averages of 9.09^I, 5.72^II, and 0.16^III were observed in 106 cells at metaphase I. A modified form of the H. bogdanii genome appears to occur in S. hystrix as well as in E. canadensis. Many allotetraploid Agropyron, Elymus, and Sitanion species apparently contain a genome derived from Hordeum.     

Experiments on growth interactions between two invasive macrophyte species

Abstract. The success of invasive species has been attributed to the ability to displace other species by direct competition. We studied growth and possible competition between the two macrophyte species Elodea nuttallii and E. canadensis, because the former has been observed to replace the latter in the field. Additional experiments were conducted in aquaria with mixed plantings of Elodea species. Species growth was measured and competitive abilities of each species determined by applying the reciprocal yield model to mean plant weight and length. In monocultures the growth rates of the two species were similar, while in mixtures the growth rate of E. canadensis was significantly lower than that of E. nuttallii. E. canadensis was more sensitive to intraspecific than to interspecific neighbours, whereas E. nuttallii was indifferent to the presence of neighbours. Differential growth characteristics of Elodea species can explain the displacement of E. canadensis by E. nuttallii under eutrophic field conditions.     

Competition between Invasive and Indigenous Species: Impact of Spatial Pattern and Developmental Stage

Elodea canadensis (indigenous) and Elodea nuttallii (invasive) were grown in experimental tanks in monocultures and mixtures in two spatial patterns (aggregated or mixed) and two developmental stages (small or large plants of E. canadensis, and small plants of E. nuttallii). Competitive interactions between the two species were assessed by monitoring the area colonised by each species, the number of rootings and biomass after 10 weeks. In monocultures the growth of E. canadensis was significantly lower than that of E. nuttallii. In mixtures the number of rootings and biomass of E. canadensis were always significantly less than those of E. nuttallii. The tank surface area colonised by E. canadensis was always significantly less than that occupied by E. nuttallii, but it was higher in the aggregated treatment, where the colonisation of E. nuttallii was lower. Therefore both spatial pattern and developmental stage of an indigenous species (E. canadensis) may influence the outcome of competition with potential invaders (E. nuttallii).     

Expression patterns and organization of the hsp70 genes correlate with thermotolerance in two congener endemic amphipod species (Eulimnogammarus cyaneus and E. verrucosus) from Lake Baikal

We studied various aspects of heat‐shock response with special emphasis on the expression of heat‐shock protein 70 (hsp70) genes at various levels in two congener species of littoral endemic amphipods (Eulimnogammarus cyaneus and E. verrucosus) from Lake Baikal which show striking differences in their vertical distribution and thermal tolerance. Although both the species studied demonstrate high constitutive levels of Hsp70, the thermotolerant E. cyaneus exhibited a 5‐fold higher basal level of Hsp70 proteins under normal physiological conditions (7 °C) and significantly lower induction of Hsp70 after temperature elevation compared with the more thermosensitive E. verrucosus. We isolated the hsp70 genes from both species and analysed their sequences. Two isoforms of the cytosolic Hsp70/Hsc70 proteins were detected in both species under normal physiological conditions and encoded by two distinct hsp/hsc70 family members. While both Hsp70 isoforms were synthesized without heat shock, only one of them was induced by temperature elevation. The observed differences in the Hsp70 expression patterns, including the dynamics of Hsp70 synthesis and threshold of induction, suggest that the increased thermotolerance in E. cyaneus (compared with E. verrucosus) is associated with a complex structural and functional rearrangement of the hsp70 gene family and favoured the involvement of Hsp70 in adaptation to fluctuating thermal conditions. This study provides insights into the molecular mechanisms underlying the thermal adaptation of Baikal amphipods and represents the first report describing the structure and function of the hsp70 genes of endemic Baikal species dwelling in thermally contrasting habitats.     

The species composition and distribution of free-living nematodes (Nematoda) in an area of natural oil and gas seeps in Lake Baikal

The species composition and quantitative parameters of free-living nematode fauna have been studied in an area of natural oil seeps off Cape Gorevoi Utes (Middle Baikal). A total of 34 species of 17 genera, 10 families, and 8 orders have been found, 18 of them endemic to Lake Baikal. The nematode fauna of bitumen volcanoes was found to be represented by three specialized species (Monhystera naphthera Gagarin et Naumova, 2010; Eumonhystera abyssalis Gagarin et Naumova, 2010; and Eutobrilus mirandus Gagarin et Naumova, 2011). Areas of the bottom with scattered oil seeps are dominated by eurybathic species endemic to Lake Baikal. The maximal values of nematode population density and maximum proportion of nematodes in the meiozoobenthos were recorded in areas directly adjacent to oil seeps.     

Distribution of dissolved organic carbon in Lake Baikal and its watershed

 Concentrations of dissolved organic carbon (DOC) in Lake Baikal ranged from 90 to 110 μM C, considerably higher than those in oceanic environments. The DOC concentrations in the epilimnion were higher than those in the hypolimnion. Since particulate organic carbon (POC) concentrations in the pelagic waters of Lake Baikal were <10–40 μM C in the epilimnion and 2–5 μM C in the hypolimnion, DOC constitutes a major component of the organic carbon pool in Lake Baikal, especially in the deep layers. The DOC concentrations downstream of the Barguzin and Selenga Rivers were quite high (400–500 μM C). Probably because of the high concentrations of DOC in these rivers, the DOC levels in Barguzin Bay and offshore at the mouth of the Selenga River were higher than those in the pelagic regions of the central and south basins of Lake Baikal. The relationship between DOC and electric conductivity revealed the transport of DOC from rivers to the pelagic area in Lake Baikal. The spatial distribution of DOC suggested that a major part of DOC in the lake was allochthonous (land-derived). Received: July 26, 2002 / Accepted: September 16, 2002 Present address:Research Institute for Humanity and Nature, 335 Takashima-cho, Marutamachi Kawaramachi, Kamigyo-ku, Kyoto 602-0878, Japan Tel. +81-75-229-6167; Fax +81-75-229-6150 e-mail: yos@chikyu.ac.jp Acknowldgments The authors wish to thank Director Prof. M. Grachev and Dr. O. Timoshkin of the Limnological Institute, Siberian Branch, Academy of Science, Russia, for arranging the cruise on Lake Baikal. We are also indebted to Drs. V. Sinyukovich, I. Khanaev, and A. Zhdanov for their kind assistance during the expeditions. We wish to thank Ms. Y. Ito for measuring the DOC concentrations. This work was supported and financed by Grants-in-Aid for Scientific Research from the Ministry of Education, Culture, Sports, Science and Technology (MEXT), Japan (No. 09041159) and for Scientific Research of Priority Area B (No. 11213101) and by the International Geosphere-Biosphere Programme (IGBP) at Nagoya University. Correspondence to:T. Yoshioka     

Heat shock proteins in the mechanisms of stress adaptation in Baikal amphipods and palearctic <Emphasis Type="Italic">Gammarus lacustris</Emphasis> Sars. I. HSP70 family

The participation of HSP70 family proteins in the mechanisms of stress adaptation under thermal and toxic stress in Baikal endemic and palearctic organisms was investigated. The four Lake Baikal endemic species Gmelinoides fasciatus (Stebb.), Eulimnogammarus cyaneus (Dyb.), E. vittatus (Dyb.), Ommatogammarus fl avus (Dyb.) were compared with the palearctic fauna representative Gammarus lacustris Sars. The type of synthesis of HSP70 family heat shock proteins under thermal (exposure at a temperature of 20, 25, and 30°C) and toxic (exposure in CdCl₂ solutions with the concentrations 50, 10, 5, 0.5, and 0.05 mg/l) stress was determined. All species under investigation were shown to have a common trend towards HSP70 content increase; therewith some species-specific characteristics of the type of synthesis of the protein under investigation were observed. The conclusion has been made on HSP70 participation in the mechanisms of thermal and toxic resistance in the studied amphipod species.     

Food composition and feeding rate of the lake Baikal invader <Emphasis Type="Italic">Gmelinoides fasciatus</Emphasis> (Stebbing, 1899) in Lake Ladoga

The feeding of the Baikal invader Gmelinoiudes fasciatus (Stebbing, 1899) in Lake Ladoga has been studied based on field and experimental data. The food spectrum of this amphipod was revealed; it includes many plant and animal species. Differences in the feeding of G. fasciatus living in different littoral biotopes were shown. The ecological and physiological rations of this species were determined. The effect that the bottom substrate has on the feeding rate of the amphipod was estimated. The introduction of G. fasciatus into Lake Ladoga provided a more adequate utilization of the energy accumulated by the primary producers and its transfer to higher trophic levels. This Baikal amphipod occupied a niche that had been, in effect, vacant. This made the invader capable of maintaining the quantitative parameters of its population at a very high level, and it largely reorganized the flow of matter and energy in the littoral zone of the lake.     

Finding a new form of the grayling Thymallus arcticus (thymallidae) in the basin of lake baikal

A dwarf form of the Arctic grayling Thymallus arcticus, inhabiting a group of small lakes at the origin of the Yakchii stream, which flows into the Verkhnyaya Angara River (basin of Lake Baikal), has been found in the northeastern watershed of the Baikal and Lena basins. The form is similar to upper Lena populations in body coloration and the pattern of dorsal fin. Our comparison by meristic characters indicated that the grayling of the Yakchii Lakes is more similar to the fish from Kutima River (basin of the upper courses of the Lena River) than to the black Baikal grayling T. arcticus baicalensis. The presence in Lake Baikal of a population similar to upper Lena graylings may have three causes: (1) possible drainage from Baikal to the pra-Lena via ancient valleys of the Barguzin and the Upper Angara; (2) appearance of transit zones as a result of glacial or tectonic events during the periods of watershed development between the Baikal and Lena basins; (3) the upper Lena grayling could be an endemic of the Baikal basin which was replaced by graylings penetrating from the Yenisei basin and remained in the form of relic populations in the upper courses of certain tributaries of the northern and northeastern parts of Lake Baikal.     

Presence and genetic diversity of microcystin-producing cyanobacteria (<Emphasis Type="Italic">Anabaena</Emphasis> and <Emphasis Type="Italic">Microcystis</Emphasis>) in Lake Kotokel (Russia,Lake Baikal Region)

A survey was conducted for the presence of cyanobacteria toxins in Lake Kotokel due to a few cases of Haff disease registered in 2008–2009 caused by consumption of fish from Lake Kotokel, and wildlife mortality including large fish kill. The aims of this study were to determine what cyanotoxins (if any) were present in the lake, to describe phytoplankton composition including morphology, density, and species diversity of cyanobacteria, as well as to evaluate the trophic state of the lake. Samples were collected from both nearshore and central sites in August of 2009. Aphanocapsa holsatica dominated the phytoplankton. The presence of toxigenic genotypes of Microcystis spp. and Anabaena lemmermannii was detected by sequencing of PCR-amplified aminotransferase domain of microcystin synthetase gene. LR, RR, and YR microcystin (MC) variants were detected with liquid chromatography-UV mass spectrometry. The data do not shed light on the etiology of Haff disease in Lake Kotokel region, nevertheless taking into account the recreational importance of the lake and its direct connection to Lake Baikal, a necessity to monitor cyanobacteria in these water bodies is evident. This is the first report on simultaneous detection of MC-producing genotypes and MCs in the Lake Baikal region.     

Regeneration dynamics of Tsuga canadensis in the southern Appalachian Mountains, USA

Tsuga canadensis is often considered a foundation species because of its influence on the biological and ecological processes of upland forest systems. With hemlock woolly adelgid (Adelges tsugae; HWA)-induced mortality of T. canadensis in southern Appalachian upland forests, there is a crucial need to understand the regeneration dynamics of this tree species. This research was conducted within mature T. canadensis forests of Great Smoky Mountains National Park (GSMNP) prior to widespread HWA infestation. Tsuga canadensis regeneration abundance is lower than has been reported in more northern locales, but varies widely among sites in the southern Appalachian Mountains. Ordination results suggest landscape-scale patterns of regeneration abundance that are influenced by gradients in understory Rhododendron maximum cover, soil sand and nutrient content. Regression models reinforce the observed gradients in regeneration abundance. Sites with abundant T. canadensis regeneration have less R. maximum cover, soil sand and litter depth, but more soil nutrient availability. Tsuga canadensis seedlings and saplings tend to be spatially aggregated at the 14 × 21 m plot scale. Sites with spatially aggregated seedlings and saplings have greater canopy openness and more species with gap affinity. This research elucidates associations between successful T. canadensis regeneration and site factors such as soil nutrient availability and canopy dynamics. More forest surveys are required to understand the conditions necessary for successful T. canadensis regeneration. Land managers involved in forest restoration efforts may have to utilize appropriate silvicultural techniques conjoined with R. maximum removal operations to regenerate T. canadensis in southern Appalachian upland forests.     

SYNTHETIC AGROPYRON-ELYMUS HYBRIDS: II. ELYMUS CANADENSIS X AGROPYRON DASYSTACHYUM

Pollination of 40 emasculated E. canadensis florets with A. dasystachyum pollen gave rise to 12 shriveled seeds, one of which was viable. The hybrid seedling developed into a vigorous plant whose vegetative and spike characteristics were for the most part intermediate between those of the parents. Both parents behaved cytologically as allotetraploids, 2n = 28, and formed 14 bivalents regularly at metaphase I. Chromosome associations in the hybrid, 2n = 28, averaged 0.42 I, 13.70 II, 0.02 III, and 0.03 IV in 142 metaphase-I cells interpreted. The hybrid produced 165 seeds under open-pollination during its second year in the field. E. canadensis and A. dasystachyum apparently contain closely homologous genomes whose major structural differences consist of several inverted segments and possibly a small reciprocal translocation. The close homology between the E. canadensis and A. dasystachyum genomes was unexpected in view of the wide morphological, ecological, and reproductive differences that separate the two species.