Two Corbicula (Corbiculidae: Bivalvia) mitochondrial lineages are widely distributed in Asian freshwater environment

We investigated the biogeography of Asian Corbicula using mitochondrial gene sequence variation for Corbicula members sampled from 24 localities of eight Asian regions. A total of 210 individuals were genetically characterized by examining sequence variations of a 614 bp fragment of the mitochondrial cytochrome oxidase I (COI) gene. Phylogenetic analyses of the COI dataset revealed that Corbicula members are subdivided into two well-supported clades: estuarine and freshwater. A robust dichotomy between the Japanese/Korean (Corbicula japonica) and Chinese (Corbicula fluminalis) estuarine forms was evident, suggesting that these two regional populations represent a deep phylogeographic split. Our mitochondrial gene tree showed that among the freshwater members, two Corbicula mitochondrial lineages are the most common, having attained extensive geographic distribution in the Asian freshwater environment. While the present study provides significant biogeographic information on Asian Corbicula, a comprehensive phylogenetic study by cross-referencing the mitochondrial-based Corbicula phylogeny with nuclear gene data is required to fully understand the evolutionary origin(s) of triploidy/clonality in this genus.     

The effect of phytoplankton and suspended sediment on the growth of Corbicula fluminea (Bivalvia)

Juvenile Corbicula fluminea Müller (1774) were cultured at 15.3 °C in the laboratory on eight combinations of suspended sediment and phytoplankton. Sediment concentrations were 2.6, 25, 50, and 150 mg l^–1. Chlorophyll a levels were 15.6 and 62.5 µg l^–1. Clam tissue growth was found to be independent of silt concentration but increased at the higher chlorophyll level (p < 0.05). The growth experiment was repeated at 24 °C with chlorophyll a concentrations of 18.9 and 112.6 µg l^–1. Growth was again greater at the higher phytoplankton level (p < 0.05). These results demonstrate that Asiatic clam populations are food-limited most of the growing season in the Northern and Western portions of California's eutrophic Sacramento-San Joaquin Delta where chlorophyll a levels average less than the lower of these values. Comparisons of clam growth in the laboratory and estuary support the food limitation hypothesis as at the higher food concentration laboratory tissue growth was 2.3 and 3.8 times greater during the high and low temperature evaluations than in the estuary.     

Growth of Corbicula fluminea (bivalvia) fed artificial and algal diets

Laboratory experiments were conducted to assess the growth of Corbicula fluminea Müller (1774) fed mixed diets of the green alga Ankistrodesmus and five artificial foods (nine-grain cereal, rice flour, rye bran, denatured brewer's yeast and Ralston Purina trout chow). Clams lost weight after 30 days on each artificial diet; however, weight losses were significantly less than those of control clams which were not fed (p < 0.05). In contrast, clam growth was positive in each treatment which included Ankistrodesmus. There was no significant difference in the tissue growth of clams fed on an Ankistrodesmus monoculture and Ankistrodesmus supplemented with each artificial diet (p > 0.2). Therefore, we conclude that all of the artificial diets evaluated were of nutritional value. However, the alga Ankistrodesmus appears to be a superior food item. In subsequent experiments, clams were fed on diets consisting of various combinations of six genera of green algae (Pedinomonas, Ankistrodesmus, Chlamydomonas, Chlorella, Scenedesmus, and Selenastrum). All trialgal combinations of Chlamydomonas, Chlorella, Scenedesmus, and Ankistrodesmus and a dialgal culture of Chlamydomonas and Chlorella resulted in strong positive tissue growth. No other dialgal or monoalgal culture resulted in similar growth. Cultures of Selenastrum were found to be toxic, and their filtrate was determined to inhibit Corbicula filtration.     

Reproduction and development of the freshwater clam Corbicula australis in southeast Australia

The freshwater clam Corbicula australis is an important component of the macrobiota of the river systems of southeast Australia. Reproduction of two populations of this clam in the Nepean River at Douglas Park and Menangle was investigated to document the gametogenic cycle, larval morphology and to determine when they incubate embryos. C. australis is a simultaneous hermaphrodite and broods its young in the inner demibranchs. The gonads are ovotestes with oogenic and spermatogenic regions in each ascinus. The sperm are biflagellate, a condition unique in the Bivalvia to clonal corbiculids. Gametogenesis was continuous and did not exhibit a seasonal pattern. In contrast, spawning and incubation of embryos was limited to the warmer months of the year. Embryos were present in the gills from October to May. C. australis develops through a modified veliger larva with a vestigially ciliated velum which is not used for swimming or particle capture. The velum is covered by microvilli and it is suggested that the velar epithelium may be specialised for nutrient uptake in the marsupial environment. C. australis produces several clutches each year and the young are released as advanced juveniles with a well-developed foot. Reproductive output differed between the two populations. This was in part due to the larger size of the clams from Menangle and may also reflect the enhanced productivity at this site. The suite of life history traits exhibited by C. australis: hermaphroditism, potential for self-fertilization/androgenesis, brooding progeny to the crawl-away juvenile stage and a high reproductive output, provide for rapid colonization and population growth in this clam which typically inhabits disturbance prone sandy lotic habitats.     

Karyotypes of Cerastoderma glaucum (Bivalvia) from Baltic and Mediterranean populations

Chromosomes of C. glaucum from Baltic (Gdansk Bay) and Mediterranean (Thau lagoon, Sète) populations were studied using karyometric analysis and silver-staining. The karyotype of the Gdansk population consists of three metacentric, ten submetacentric and six subtelocentric chromosome pairs. The karyotype of the Thau population shows four metacentric, nine submetacentric and six subtelocentric pairs. Nucleolus organizer regions were found terminally on the second largest submetacentric chromosome pair of the Gdansk population and on the second largest metacentric chromosome pair of the Thau population. This suggests a cytotaxomomic difference between these two geographically isolated populations. Striking differences exist between C. glaucum and C. edule karyotypes, probably resulting from a long divergence time.     

Corbicula fluminea (Bivalvia: Corbiculidae): a possible second molluscan intermediate host of Echinostoma cinetorchis (Trematoda: Echinostomatidae) in Korea.

More than 1,500 clams of Corbicula fluminea, the most favorable food source of freshwater bivalves in Korea, were collected from 5 localities to examine cercarial and metacercarial infection with Echinostoma cinetorchis. Although 3 clams infected with suspicious E. cinetorchis metacercariae out of 200 specimens collected at Kangjin, Chollanam-do were detected, no cercarial and metacercarial infections with E. cinetorchis were observed in field-collected Corbicula specimens. In the susceptibility experiments with laboratory-reared clams, those infected with miracidia of E. cinetorchis did not release their cercariae up to 60 days after infection. To confirm the identity of second intermediate host of E. cinetorchis experimentally, a total of 30 clams were exposed to the cercariae from Segmentina hemisphaerula that had been infected with miracidia of E. cinetorchis. The clams were susceptible to cercariae of E. cinetorchis with an infection rate of 93.3%. Metacercariae from clams taken more than 7 days after cercarial exposure were fed to rats (S/D strain), and adult worms of E. cinetorchis, characterized by 37-38 collar spines on the head crown, were recovered from the ileocecal regions. This is the first report of C. fluminea as a possible second intermediate host of E. cinetorchis.     

Brooding in Corbicula madagascariensis (Bivalvia, Corbiculidae) and the repeated evolution of viviparity in corbiculids

The limnic bivalve genus Corbicula Megerle von Mühlfeld, 1811 is a hyper-invasive neozoon in North and South America as well as in Europe, where currently some taxa are rapidly extending their range. In addition to its extraordinarily invasive potential, the 'Asiatic clam' is remarkable for its recently discovered wide spectrum of reproductive strategies comprising oviparity, ovoviviparity and euviviparity. It renders Corbicula an ideal model for studying evolutionary transformations of reproductive features, in particular with respect to intrabranchial incubation (brooding) of embryos and shelled larvae in freshwater lineages. Based on rare material from Madagascar we here present evidence for prolonged incubation interpreted as being indicative of euviviparous reproduction in C. madagascariensis Smith, 1882. This mode is not only novel for corbiculids from the Ethiopian biogeographical region, but suggests — in combination with a mtDNA phylogeny — a more complicated pattern of the evolution of reproductive modes in corbiculids than previously assumed. We find an independent origin of viviparity and even euviviparity in the South American Neocorbicula Fischer, 1887 and the Afro-Asian Corbicula , representing a remarkable example of parallel evolution in New and Old World corbiculids.     

Spawning periodicity and annual breeding seasons of Tilapia nilotica (L.) in the White Nile

Two annual breeding seasons are indicated for populations of T. nilotica in the White Nile: a major autumnal season (mid-July–September) coinciding with the annual rainfall and a smaller back-up season in February–March. During these seasons reproducing females constituted 80% and 40% of the total population of mature females compared to 10–12% in the intervening summer period. Enhanced gonadal development, fecundity and GSI values characterized the breeding seasons. Gonadal maturation proceeded uninterruptedly even during the summer and repeated fractional spawning is suggested for this species within and beyond the breeding seasons. Most of the spring breeders were either very young or relatively old fish and most medium-sized females bred in the main autumnal season. Older fish were less fecund (by 70–75%) compared to younger fish. In the latter, fecundity increased progressively with body growth (r² = 0.732; p<0.001); the proportionality was lost in medium-sized fish and in older females (0.9–1.5 kg) fecundity correlated negatively with increased body size.     

The karyotype of Nodipecten nodosus (Bivalvia: Pectinidae)

Earlier karyotypical work on Nodipecten nodosus embryos indicated that this species has a diploid number of 38, with six pairs respectively of metacentric and submetacentric chromosomes and seven pairs of subtelocentric chromosomes, although there were some difficulties in obtaining complete metaphases. The present work provides additional results on specific regions of the chromosomes in N. nodosus and, by meiotic studies, confirms the chromosome number with more reliability. Active nucleolar organizer regions (NOR), detected in mitotic metaphases from embryos, can be characterized in N. nodosus by a high level of heteromorphism of NOR-sites, indicating that these regions are not appropriate as chromosomal markers in this species. The procedure for detecting constitutive heterochromatin of chromosomes allowed us to observe most of the heterochromatic blocks at a pericentromeric position and some at telomeric and interstitial positions. The analysis of meiotic chromosomes from gonad tissue revealed the presence of 19 bivalents during metaphase I, all homomorphic and isopicnotic, confirming the previously described diploid chromosomal number of 38 for N. nodosus. From these results, some evolutionary aspects of the Pectinidae are briefly discussed.     

Secondary production of introduced Asiatic clam,Corbicula fluminea,in a central Arizona canal

Asiatic clam, Corbicula fluminea, is often a serious pest where introduced and established outside its native range. This includes many canals of the southwestern U.S.A. Because of its potential role in organic matter processing, nutrient dynamics, and sedimentation, the clam is an important component of the benthic community and of the aquatic ecosystems which it inhabits. To better understand the ecology of Corbicula in canals, secondary production and life history of an introduced population of clams in a small, earthen canal in the Phoenix metropolitan area, central Arizona, were studied over a 12-month period in 1981–1982. Clams had a monthly mean density of 2 255 m^–2, single annual spawning, 2 year life span, and overlapping cohorts. Annual secondary production (size-frequency method) was 25.62 g DM m^–2 and cohort production (composite Allen curve) was 29.32 g DM m^–2, with turnover ratios of 2.70 and 5.81, respectively. These are among the highest known single-species estimates for molluscs, and illustrate the importance of clams in these unique southwestern aquatic systems.     

Reproduction and sperm structure in Galeommatidae (Bivalvia, Galeommatoidea)

Reproduction and/or sperm structure was studied in 18 species belonging to five genera of Galeommatidae from Phuket Island, Andaman Sea, Thailand, and Hong Kong. Incubation of the ova occurs in the inner and outer demibranchs of both gills, and sperm is most probably transferred to the ctenidial brood chamber as masses of agglutinated and non-encapsulated spermatozoa. The smallest specimens are males, but change into the female sex and there is a strong indication that some of the species are alternate hermaphrodites. Dwarf males occur in one species (Galeomma layardi). The sperm of all 18 species studied are of the ent-aquasperm type with a more or less hemispherical acrosome that is tilted with respect to the long axis of the moderately elongated nucleus. It is suggested that this type of sperm affords the best autapomorphy for the family Galeommatidae.     

Scale-dependent associations between native freshwater mussels and invasive Corbicula

In North America there is conflicting evidence concerning whether the invasive Asian clam, Corbicula fluminea, and native mussels (Unionidae), can successfully co-exist. One reason underlying disparate conclusions may be the different spatial scales at which data have been collected. We compared the distribution and abundance of native unionid mussels and Corbicula at two spatial scales, stream reaches and 0.25 m² patches, within one biogeographic region, the Ouachita Highlands, of the south central U.S. We found that Corbicula abundance was negatively related to native mussel abundance at small spatial scales. While Corbicula densities varied widely in patches without native mussels, and in patches where mussels occurred at low abundance, Corbicula density was never high in patches where mussels were dense. We hypothesize that the likelihood of successful Corbicula invasion decreases with increasing abundance of adult native mussels. Several mechanisms may potentially drive this pattern including lack of space for Corbicula to colonize, physical displacement by actively burrowing mussels, and locally reduced food resources in patches where native mussels are feeding. In addition, Corbicula may be unable to withstand environmental bottlenecks as readily as unionids. When patch-scale density and biomass information were pooled to represent entire stream reaches, the negative relationship between native mussels and Corbicula was no longer as apparent, and there was not a significant relationship between native mussels and Corbicula. These results point to the importance of appropriate sample scale in examining potential associations between species.     

Seasonal abundance,breeding and sex-structure of populations of Tilapia nilotica (L.) and Labeo niloticus (F.) in the Jebel Aulia Dam area of the White Nile

Seasonal abundance, maturation and sex-structure were studied in populations of T. nilotica and L. niloticus in the Jebel Aulia reservoir and in the White Nile below the dam. L. niloticus showed a single annual breeding cycle from mid-July to December and was most abundant in October in the lake, and in early November in the river. T. nilotica showed two distinct annual seasons of abundance: summer (April to June) and autumn (November to October). Spawning activity was highest in summer (April to June) and autumn (late July to September). The sex-ratio varied inversely with age (size); males were predominant in populations of older fish, perhaps because of their greater endurance to adverse conditions as evidenced by their better survival in hypoxic water.     

The macroinvertebrate fauna of water hyacinth fringes in the Sudd swamps (River Nile,southern Sudan)

The water hyacinth now forms an almost ubiquitous fringe to open-water habitats in the permanent wetland of the Sudd. In a typical river-lake the fringe varied between 9 and 16 m in breadth with highest plant biomasses in the centre. Largest quantities of macroinvertebrates in hyacinth root-mats were obtained in drop-trap samples within 6 m of the open water. They were dominated by coleopterans, odonatans, gastropods andCaridina nilotica. Outer edges of the fringe offer easy access, good dissolved oxygen concentrations, and a variety of potential food resources for aquatic invertebrates. By contrast the less hospitable landward zone contained a reduced fauna in which coleopterans, hydracarines and gastropods were prominent. A marked reduction in the numbers of invertebrates in lift-net samples of detached hyacinth rafts was attributed to browsing by fish. Hyacinth root-mats appear to have replaced the niches formerly provided by those of the Nile cabbage for aquatic invertebrates in the Sudd.     

Variation in wild populations of annual beet (Beta,Chenopodiaceae)

Among the morphological variation in wild annual populations of sect.Beta only tepal characters show a geographic pattern, and hence can be used to distinguish different taxa. Two morphological types correspond with taxa already described:B. macrocarpa (incl.B. bourgaei) has a Macaronesian and W. to E. Mediterranean andB. adanensis an E. Mediterranean distribution area. A third type in the Aegean region is not well known yet and possibly has to be included inB. macrocarpa. Both diploid and tetraploid (x = 9) cytotypes are found withinB. macrocarpa, the latter exclusively on the Canary Islands.     

Auxospore formation inLicmophora (Bacillariophyta)

InLicmophora gracilis var.anglica two auxospores are produced per pair of mother-cells, through the allogamic fusion of migratory and stationary gametes. Both active gametes are produced from the same mother-cell and hence both zygotes are formed in the other mother-cell. Pairing can occur between two stalked cells, or between a stalked cell and a detached cell; in the latter case the migratory gametes derive from the detached cell. The auxospores expand parallel to one another and to the apical axis of the donor mother-cell. Behavioural anisogamy of this kind, which may be termed thecis-type, seems to be characteristic of most araphid pennates and contrasts with thetrans-type exhibited byCymbella, Gomphonema and some other raphid taxa, where each mother-cell produces one migratory and one stationary gamete.     

The effect of environmental conditions on distribution,growth and life span of <Emphasis Type="Italic">Protothaca euglypta</Emphasis> (Mollusca: Bivalvia)

The distribution and growth of the bivalve mollusk Protothaca euglypta in Peter the Great Bay and at the northern boundary of its area were investigated. It is shown that in some biotopes P. euglypta biomass reaches 1–2 kg/m², which exceeds the estimated values of abundance obtained for most associated species of mollusks. It has been established that local variability of P. euglypta growth rates can be compared with latitudinal variability, and the life span increases from the southern part of the area towards its northern boundary. It has been concluded that P. euglypta occupies a more prominent position in the malacofauna of shallow waters in its abundance, than it was considered earlier.     

Pappus part number in annual species ofMicroseris (Compositae,Cichoriaceae)

All North American annual species of the genusMicroseris have a five-part pappus, the one South American annual,M. pygmaea, has ten pappus parts. The pappus develops over a constant number of ten provascular bundles with or without inhibition between alternate sites of pappus development. Each natural population contains a predictable proportion of achenes with aberrant pappus part numbers. Hybridization betweenM. bigelovii (5 parts) andM. pygmaea results in F 1 and F 2 plants with many aberrant achenes. In each plant either five or ten can be shown to be the basic number with aberrant numbers following a Poisson distribution for numbers added to 5 or deleted from 10. Occasional plants show no basic number but have a random distribution of numbers about an intermediate mean. The evolutionary genetics of this character is discussed.