Birds in the trophic web of Lake Esrom,Denmark

Using an extended method for calculating food consumption by diversified bird communities taking into account variability of diet, energy content and digestibility of different food items within a diet, it was possible to calculate the energy consumption of the bird community in eutrophic Lake Esrom. As the production of food items and energy flow in Lake Esrom have been well studied, it was possible to relate the bird community to the trophic web. Birds consumed approximately 20% of the annual macrophyte production (excluding roots and rhizomes), and 4% of the annual zoobenthos production. The annual consumption of fish by birds corresponded to 22% of the annual consumption by piscivorous fish measured in kJ, and was almost the same as the commercial fishery catch. However, the consumption of fish by birds consisted mainly of non-piscivorous fish, in contrast to the commercial catch. In Lake Esrom, the birds have a significant role in the energy flow of consumers within the ecosystem. This result indicates that bird communities should be included in future studies of lake ecology. Also, competition between fish and birds is potentially significant.     

Life-history strategies and production of Tinodes waeneri (L.) (Trichoptera) in Lake Esrom,Denmark

The life cycle and production of Tinodes waeneri (L.) was examined at five shallow littoral localities in Lake Esrom, Denmark, during 1979/80. Five larval instars were demonstrated. The differences in temporal composition of instars indicated three different life cycles. A bivoltine strategy was shown for the most exposed locality which also had the highest average abundance equal to 11 500 ind. m^–2. A univoltine population was found in the more sheltered NW part of the lake with an average density of 1 500 ind. m^–2. At the three remaining stations the life cycles were partly bivoltine with wintering populations of mixed cohorts and densities between 10 500 and 3 000 ind. m^–3. Two distinct flight periods with maxima in June and August were demonstrated for bi- and partly bivoltine populations. Differences in rates of growth and elimination added to the variability in life cycle. Growth rates peaked in August (5.0–7.4% d^–1) at an average temperature of 20°C. Estimates of production ranged from 1.9 to 17.5 g AFDW m^–2 a^–1 with an average of 8.0 g AFDW m^–2. The P/B ratios, which were within the expected range for uni- and bivoltine populations of trichopterans, were 5.73 for the bivoltine population and 3.31 for the univoltine population.Estimates of consumption revealed that the populations could be sustained by autochthonous epilithic production in the habitats. Variability in the quality and seasonal availability of food were judged to influence the type of life cycle.Lake Esrom Littoral Research Publ. No. 8. Publication No. 385 from the Freshwater Biological Laboratory, University of Copenhagen, 51, Helsingørsgade, DK-3400 Hillerød, Denmark.Lake Esrom Littoral Research Publ. No. 8. Publication No. 385 from the Freshwater Biological Laboratory, University of Copenhagen, 51, Helsingørsgade, DK-3400 Hillerød, Denmark.     

Species delimitation through DNA barcoding of freshwater leeches of the Glossiphonia genus (Hirudinea: Glossiphoniidae) from Eastern Siberia,Russia

The study of biodiversity is a priority task of biological science. The structural unit of biodiversity is a species that has a clear identification in a taxonomic system. Morphological features are traditionally the main criteria for species discrimination in zoological studies. However, the presence of inter- and intraspecific polymorphism and phenotypic plasticity makes it difficult to identify species in many groups of invertebrates. To solve this problem, in this research, we analyzed morphological and genetic data in combination to delimit species among the Eastern Siberia Glossiphonia leeches using different approaches. Morphology analysis revealed phenetically distinct groups, suggesting the existence of at least two species in the region, G. verrucata, a rare Palaearctic species, and a potentially new species Glossiphonia sp. Moreover, sequence-based species delimitation methods congruently supported eight distinct species groups (including two Siberian species) within the available molecular dataset of the Glossiphonia world fauna, using phylogenetic (ML and BI), coalescent (ABGD and GMYC) methods, and pairwise analysis of sequences. The detected p-distances (modal value of 0.11) between these 8 groups and the level of genetic polymorphism (max. 0.0041) within groups indicate that the groups are 8 independent species according to the DNA barcoding. Our results once again proved the usefulness of molecular systematics. At the same time, we detected several inaccuracies in the leech species identification, as well as many ambiguous sites in sequences uploaded on GenBank, which affects the analysis and impedes progress of DNA barcoding technology.     

The Life History and Occurrence of Sialis lutaria L. (Megaloptera) in the Littoral Zone of Lake Esrom,Denmark

The life history and distribution of Sialis lutaria L. in the exposed littoral zone of Lake Esrom, Denmark were examined during 1979/80. Sialis showed an annual life cycle with mean annual abundance of 20 and 40 ind m⁻² at 0.4 and 2.0 m depth, respectively. Growth peaked during July with rates of about 10 % d⁻¹. Net annual production averaged 0.2 g DW at 0.4 m depth and 0.8 g DW at 2.0 m depth. The annual P/B-ratios were estimated in the range of 4.6-7.1.     

Structure of the germinal vesicle during oogenesis in leech Glossiphonia heteroclita (Annelida, Hirudinea, Rhynchobdellida)

Oogenesis in the glossiphoniid leech Glossiphonia heteroclita (Hirudinea, Rhynchobdellida) is nutrimental, i.e., the growing oocyte is supported by specialized germline cells, the nurse cells. The main function of the nurse cells is to provide oocytes with cell organelles and RNAs (mainly rRNA). However, in studied leech species, irrespective of the nutrimental mode of oogenesis, the germinal vesicle (GV = oocyte nucleus) seems to be very active in rRNA production. As shown in the present study, during early previtellogenesis in the GV the meiotic chromosomes and prominent primary nucleoli occur. In late previtellogenesis the chromosomes condense and occupy a limited space of nucleoplasm in close vicinity to primary nucleolus, forming a karyosome. At the onset of vitellogenesis several prominent extrachromosomal DNA bodies appear in close association with the karyosome. At the same time, the primary nucleolus is no longer visible in the GV. As vitellogenesis proceeds the extrachromosomal DNA bodies undergo fragmentation and numerous spherical, RNA- and AgNOR-positive inclusions occur in the nucleoplasm. They are regarded as multiple nucleoli. Finally, in late oogenesis numerous accessory nuclei are formed in close proximity to the nuclear envelope. They usually contain one dense body, morphologically similar to multiple nucleoli. The amplification of rDNA genes, the occurrence of extrachromosomal DNA bodies, as well as the presence of multiple nucleoli and accessory nuclei are described for the first time in the phylum Annelida.     

Population dynamics and energy budget of Marionina southerni (Cernosvitov) (Enchytraeidae,Oligochaeta) in the shallow littoral of Lake Esrom,Denmark

Marionina southerni (Cernosvitov) was numerically the dominant oligochaete in a shallow, strongly exposed surf zone locality (0–1 m depth) in the mesotrophic Lake Esrom. It comprised 21,500 ind m^–2 or nearly 50% of the total oligochaete community, which otherwise was dominated by Nais spp. and the lumbriculid Stylodrilus heringianus Claparède. M. southerni appears to have a 1-y life cycle. Individual biomass ranged from 3 to 48 µg ash free dry weight. Annual net production at the site was 5.1 kcal m^–2 (size-frequency method) with a P/B ratio of 2.5. This is about 0.5% of the estimated mean zoobenthic production in the littoral zone from 0 to 2 m depth in Lake Esrom. At 5° C M. southerni showed a constant oxygen uptake down to 25% oxygen saturation, but practically no regulatory respiration was found at 20 °C. In the field M. southerni was never found in sediment with less than 60% oxygen saturation. The annual community respiration of M. southerni was 14.2 kcal m^–2, and annual assimilation thus made up 19.3 kcal m^–2 with a net production efficiency of 26.5%.     

Reproduction and Larval Morphology of Five Ophryotrocha Species (Poly cha eta, Dorvilleidae)

Reproduction and larval morphology of Ophryotrocha puerilis, O. labronica, O. nologlandulata, O. hartmanni , and O. bacci are described. All species have been cultivated in the laboratory. The morphology of adults and larvae is uniform, but there is a considerable variation in the shape of the egg masses. The interrelations between members of the genus Ophryotrocha are discussed: Gonochoric species have the same type of egg mass and the same chromosome number. O. puerilis is unique as a protandrous hermaphrodite. Among contemporary hermaphrodites with male and female segments, fusiform egg masses with a rigid surface are correlated with one larval type, gelatinous egg masses with another larval type.     

Oogenesis in the leech Glossiphonia heteroclita (Annelida, Hirudinea, Glossiphoniidae) II. Vitellogenesis, follicle cell structure and egg shell formation

By the end of previtellogenesis, the oocytes of Glossiphonia heteroclita gradually protrude into the ovary cavity. As a result they lose contact with the ovary cord (which begins to degenerate) and float freely within the hemocoelomic fluid. The oocyte's ooplasm is rich in numerous well-developed Golgi complexes showing high secretory activity, normal and transforming mitochondria, cisternae of rER and vast amounts of ribosomes. The transforming mitochondria become small lipid droplets as vitellogenesis progresses. The oolemma forms microvilli, numerous coated pits and vesicles occur at the base of the microvilli, and the first yolk spheres appear in the peripheral ooplasm. A mixed mechanism of vitellogenesis is suggested. The eggs are covered by a thin vitelline envelope with microvilli projecting through it. The envelope is formed by the oocyte. The vitelline envelope is produced by exocytosis of vesicles containing two kinds of material, one of which is electron-dense and seems not to participate in envelope formation. The cortical ooplasm of fully grown oocytes contains many cytoskeletal elements (F-actin) and numerous membrane-bound vesicles filled with stratified content. Those vesicles probably are cortical granules. The follicle cells surrounding growing oocytes have the following features: (1) they do not lie on a basal lamina; (2) their plasma membrane folds deeply, forming invaginations which eventually seem to form channels throughout their cytoplasm; (3) the plasma membrane facing the ovary lumen is lined with a layer of dense material; and (4) the plasma membrane facing the oocyte forms thin projections which intermingle with the oocyte microvilli. In late oogenesis, the follicle cells detach from the oocytes and degenerate in the ovary lumen.     

Morphology and ultrastructure of the midgut in Piscicola geometra (Annelida, Hirudinea)

This paper presents information on the organization of the midgut and its epithelium ultrastructure in juvenile and adult specimens of Piscicola geometra (Annelida, Hirudinea), a species which is a widespread ectoparasite found on the body and gills and in the mouth of many types of fish. The analysis of juvenile nonfeeding specimens helped in the explanation of all alterations in the midgut epithelium which are connected with digestion. The endodermal portion (midgut) of the digestive system is composed of four regions: the esophagus, the crop, the posterior crop caecum, and the intestine. Their epithelia are formed by flat, cuboidal, or columnar digestive cells; however, single small cells which do not contact the midgut lumen were also observed. The ultrastructure of all of the regions of the midgut are described and discussed with a special emphasis on their functions in the digestion of blood. In P. geometra, the part of the midgut that is devoid of microvilli is responsible for the accumulation of blood, while the epithelium of the remaining part of the midgut, which has a distinct regionalization in the distribution of organelles, plays a role in its absorption and secretion. Glycogen granules in the intestinal epithelium indicate its role in the accumulation of sugar. The comparison of the ultrastructure of midgut epithelium in juvenile and adult specimens suggests that electron-dense granules observed in the apical cytoplasm of digestive cells take part in enzyme accumulation. Numerous microorganisms were observed in the mycetome, which is composed of two large oval diverticles that connect with the esophagus via thin ducts. Similar microorganisms also occurred in the cytoplasm of the epithelium in the esophagus, the crop, the intestine, and in their lumen. Microorganisms were observed both in fed adult and unfed juvenile specimens of P. geometra, which strongly suggests that vertical transmission occurs from parent to offspring.     

Population dynamics and production of Daphnia galeata (Crustacea, Cladocera) in Lake Esrom

The population dynamics and production of D. galeata Sars were investigated from April 1974 to April 1975. The population density and biomass was maximum in June. Population dynamics indicated that food supply was better in spring and early summer than mid-summer. Net production was calculated by method of daily addition of biomass for eight size groups. The total net production of biomass for the year of investigation was 26.5 g dw m⁻². The P/B yr⁻¹ was 35 8.     

Diffuse Coevolution between Two Epicephala Species (Gracillariidae) and Two Breynia Species (Phyllanthaceae)

The diffuse coevolution between two moth species (Epicephala lativalvaris and E. mirivalvata) and two plant species (Breynia fruticosa and B. rostrata) is reported based on field observations and indoor experiments conducted in Hainan and Fujian, China. Study results showed that the two Epicephala species jointly pollinated the two Breynia species, which led to a unique obligate pollination mutualism of two-to-two species specificity. A single Epicephala larva exclusively fed on seeds of host plants and developed to maturity by consuming all six seeds of each fruit, whereas a fraction of intact fruits were left to ensure the reproduction of plants within the whole population. Larvae of the two Epicephala species are competitive for resources; the population of E. mirivalvata is much smaller than that of E. lativalvaris, which has resulted from the differences in the female ovipositor structures and oviposition mode. The life history of Epicephala species highly coincides with the phenology of Breynia plants, and different phenology of B. fruticosa resulted in the different life history of the two Epicephala species in Hainan and Fujian. The natural hybridization of two host plants, possibly induced by the alternate pollination of two Epicephala species, is briefly discussed.     

The effect of wind exposure and filamentous algae on the distribution of surf zone macroinvertebrates in Lake Esrom,Denmark

Distribution and composition of the epilithic macroinvertebrate fauna was studied on 21 different sites situated in the surf zone of Lake Esrom. A relative degree-of-exposure (windinduced wave activity) was calculated for each station from the specific number of wind days, wind velocity and a measured wind fetch. Macroinvertebrate abundance averaged 20 200 ind. m^–2 stone bottom, and was composed primarily of six taxa (70% of the mean densities). The distribution patterns of the 48 most common taxa were compared to exposure, the amount of filamentous algae and shore slope, by regression, cluster analysis and ordination analysis (PCA and RDA). The abundance of 25 taxa was significantly correlated to the amount of filamentous algae, and a protectional effect of algae growth was suggested for the exposed sites. Seventeen taxa showed significant difference in abundance between sheltered western stations and exposed eastern stations. Stations with a gentle slope were characterised by sand covered stones and a fauna that qualitatively was similar to the fauna at greater depth in the littoral. Redundancy analysis (RDA) showed filamentous algae and slope to be the most important factors controlling the species distribution, whereas the gradient of exposure in Lake Esrom was too short to explain the species tolerance range.     

Oogenesis in the leech Glossiphonia heteroclita (Annelida, Hirudinea, Glossiphonidae). I. Ovary structure and previtellogenic growth of oocytes

Glossiphonia heteroclita has paired ovaries whose shape and dimensions change as oogenesis proceeds: during early previtellogenesis they are small and club-shaped, whereas during vitellogenesis they broaden and elongate considerably. During early oogenesis (previtellogenesis), each ovary is composed of an outer envelope (ovisac) that surrounds the ovary cavity and is filled with hemocoelomic fluid, in which a single and very convoluted ovary cord is bathed. The ovary cord consists of germline cells, including nurse cells and young oocytes surrounded by a layer of elongated follicle cells. Additionally, follicle cells with long cytoplasmic projections occur inside the ovary cord, where they separate germ cells from each other. The ovary cord contains thousands of nurse cells. Each nurse cell has one intercellular bridge, connecting it to a central anucleate cytoplasmic mass, the cytophore (rachis); it in turn is connected by one intercellular bridge with each growing oocyte. Numerous mitochondria, RER cisternae, ribosomes, and Golgi complexes are transported from the nurse cells, via the intercellular bridge and cytophore, to the growing oocytes. Oogenesis in G. heteroclita is synchronous with all oocytes in the ovary in the same stage of oogenesis. The youngest observed oocytes are slightly larger than nurse cells, and usually occupy the periphery of the ovary cord. As previtellogenesis proceeds, the oocytes gather a vast amount of cell organelles and become more voluminous. As a result, in late previtellogenesis the oocytes gradually protrude into the ovary cavity. Simultaneously with oocyte growth, the follicle cells differentiate into two subpopulations. The morphology of the follicle cells surrounding the nurse cells and penetrating the ovary cord does not change, whereas those enveloping the growing oocytes become more voluminous. Their plasma membrane invaginates deeply, forming numerous broad vesicles that eventually seem to form channels or conducts through which the hemocoelomic fluid can easily access the growing oocytes.     

The Natural Feeding and Resource Partitioning of Erpobdella octoculata L. and Erpobdella testacea SAV. in Lake Esrom,Denmark

The natural feeding and competitive interactions between populations of Erpobdella ostoculata and Erpobdella testacea were investigated in Lake Esrom, Denmark, during 1979/80. Statistical analysis revealed that the size of the leeches was the main factor involved in resource utilization. Size-specific feeding included differences in the numerical composition of diets and the size of individual prey. Five weight classes of Erpobdella spp. were distinguished. The number of prey per leech increased from 1.63-2.84 for the first to the fifth weight class, whereas diversity in feeding averaged 1.71 prey per leech. Quantification of competitive interactions indicated that the temporal differences in weight class composition of the two species were sufficient to explain coexistence, although both species have an annual life cycle, and neither species showed taxonomic food refuges.     

Energy metabolism of Chironomus anthracinus (Diptera: Chironomidae) from the profundal zone of Lake Esrom,Denmark, as a function of body size,temperature and oxygen concentration

The profundal zone of Lake Esrom, Denmark has a dense population of Chironomus anthracinus, which survives 2–4 months of oxygen depletion each summer during stratification. The metabolism of 3rd and 4th instar larvae was examined in regard to variation in biomass and temperature. Respiration at air saturation was described by a curvilinear multiple regression relating oxygen consumption to individual AFDW and temperature. At 10 °C and varying oxygen regimes the O₂ consumption and CO₂ production of 4th instar larvae were almost unaltered from saturation to about 3 mg O₂ l^–1, but decreased steeply below this level. The respiratory quotient increased from 0.82 at saturation to about 3.4 at oxygen concentrations near 0.5 mg O₂ l^–1. This implied a shift from aerobic to partially anaerobic metabolism. At 0.5 mg O₂ l^–1 the total energy production equalled 20% of the rate at saturation of which more than one third was accounted for by anaerobic degradation of glycogen. This corresponded to a daily loss of 12 µg mg AFDW^–1 or approximately 5% of the body reserves. At unchanged metabolic rate the glycogen store would last three weeks, but long term oxygen deficiency causes a further suppression of the energy metabolism in C. anthracinus.     

Morphology,Reproduction and Diet in Australian and Papuan Death Adders (Acanthophis,Elapidae)

Death adders (genus Acanthophis) differ from most other elapid snakes, and resemble many viperid snakes, in their thickset morphology and ambush foraging mode. Although these snakes are widely distributed through Australia and Papua New Guinea, their basic biology remains poorly known. We report morphological and ecological data based upon dissection of >750 museum specimens drawn from most of the range of the genus. Female death adders grow larger than conspecific males, to about the same extent in all taxa (20% in mean adult snout-vent length,  =  SVL). Most museum specimens were adult rather than juvenile animals, and adult males outnumbered females in all taxa except A. pyrrhus. Females have shorter tails (relative to SVL) than males, and longer narrower heads (relative to head length) in some but not all species. The southern A. antarcticus is wider-bodied (relative to SVL) than the other Australian species. Fecundity of these viviparous snakes was similar among taxa (mean litter sizes 8 to 14). Death adders encompass a broad range of ecological attributes, taking a wide variety of vertebrate prey, mostly lizards (55%), frogs and mammals (each 21%; based on 217 records). Dietary composition differed among species (e.g. frogs were more common in tropical than temperate-zone species), and shifted with snake body size (endotherms were taken by larger snakes) and sex (male death adders took more lizards than did females). Overall, death adders take a broader array of prey types, including active fast-moving taxa such as endotherms and large diurnal skinks, than do most other Australian elapids of similar body sizes. Ambush foraging is the key to capturing such elusive prey.