Coexistence with similar life cycles in two species of freshwater copepods (Crustacea)

Two species of freshwater cyclopoid copepods, Cyclops scutifer Sars and Mesocyclops leuckarti (Claus), with different ecological requirements, coexist in three humic ponds near the timberline in boreal forests in south-central Norway. The life cycle in both species is basically regulated by complete depletion of oxygen during the long winter. There is a short period of reproduction in spring followed by a rapid development during summer and autumn to advanced instars, capable of diapause when oxygen depletion starts in early winter. Both species have an exclusive 1-yr cycle.The long winter under anoxic conditions is passed in diapause as large copepodids (late developmental instars) with usually no individuals left in the plankton.Even if co-occurring in the same habitats with similar life cycles, there is a temporal and spatial segregation.     

The life cycles and growth of Carabus glabratus and C. violaceus in Budalen, central Norway

Abstract. 1. The seasonal activity patterns of adults and larvae, and the seasonal development of female gonads showed that Curubus glabrafus Payk. and Curubus violaceus L. living in a subalpine habitat are spring (summer) breeders with biennial life cycles.
2. As a rule, the adults and third stage larvae hibernate, but C.glubrutus at least can hibernate also at the second larval stage, particularly in years in which unfavourable climatic conditions in the spring delay breeding.
3. The adaptive strategies underlying these life history patterns and their possible significance for species diversification and coexistence are discussed.     

Life history and production of Caenis luctuosa (Burmeister) (Ephemeroptera, Caenidae) in two nearby reaches along a small stream

Population dynamics and production of C. luctuosa were compared in two reaches of the Agüera stream (northern Spain). This species showed univoltine winter life history in both sites. However, the start of the recruitment period, and the cohort production interval differed in 1 month between reaches. Secondary production of C. luctuosa ranged from 76 mg m^–2 year^–1 (upper site) to 93 mg m^–2 year^–1 (lower site). Although annual production seemed to be mainly influenced by the biomass found at each site, changes in life history may have also been important. The need to have accurate information about life history of the analysed species at the study sites when assessing secondary production is highlighted.     

Aspects of the study of the life cycles of Antarctic copepods

Different approaches to the study of life cycle strategies of Antarctic copepods are described in an attempt to shed new light on our present knowledge. To date, most studies were carried out on abundance, horizontal and vertical distribution and stage composition during different seasons and in various regions. Hence, the seasonal pictures had to be compiled from different years and sampling regions. The physiological method includes measurements on e.g. egg production, feeding, respiration and excretion rates, C:N and O:N ratios, lipid and protein contents. However, both physiological and biochemical data are still rare. Results of field observations are given in this paper for investigations conducted within the last 15 years in the eastern Weddell Sea, while data of physiological parameters are based on a broader geographical region. In the eastern Weddell Sea, eight copepod species account for about 95% of copepod abundance and for more than 80% of copepod biomass. Within the calanoids, the small species Microcalanus pygmaeus dominates by numbers with 66%, while the large species Calanoides acutus and Calanus propinquus comprise together 52% of the biomass. Species abundance is lowest in winter and highest in summer/autumn, however, seasonal changes in the abundance of M. pygmaeus are small and this species occurs in similar quantities throughout the year. All copepod species show a distinct seasonal vertical distribution pattern and they occur in upper water layers in summer, in contrast to the other seasons. However, the depth layers of maximum concentration differ between species. The ontogenetic vertical migration is most pronounced in C. acutus and relatively weak in C. propinquus. The age structure also shows seasonal differences with the youngest population observed in summer for C. acutus, C. propinquus, Ctenocalanus citer or autumn for Metridia gerlachei, whereas the M. pygmaeus population is oldest during summer. The youngest copepodite stage and the males are not always present in C. acutus and C. propinquus. In contrast, all developmental stages and both sexes occur throughout the year in M. gerlachei, M. pygmaeus and C. citer. Gonad maturation in the dominant calanoid species proceeds well before the onset of phytoplankton production in the eastern Weddell Sea. However, the highest portion of females with ripe gonads and hence highest egg production rates coincide with the productive period in spring and summer. In autumn, ovaries of the three larger species C. acutus, C. propinquus and M. gerlacheiare all spent. In contrast, the percentage of ripe females of the two smaller species, C. citer and M. pygmaeus, stays high in autumn. Egg production rates are highly variable within one region and species. Many copepods accumulate large depots of lipid, mainly wax esters. In contrast, five species (C. propinquus, C. simillimus, Euchirella rostromagna, Stephos longipes and Paralabidocera antarctica) almost exclusively synthesise triacylglycerols and not wax esters. The lipid content exhibits distinct seasonal patterns, and is highest in autumn. A seasonal difference is also obvious in metabolic activities with lowest rates during the dark season. The adaptation to the pronounced seasonality in the Southern Ocean differs greatly between copepod species, and most Antarctic copepods stay active during the dark season. Calanoides acutus seems to be the only true diapause species. Calculations of summer developmental rates and winter mortality rates of the large species C. acutus and C. propinquus suggest that both species have a 1-year life cycle with few females overwintering and probably spawning a second time. In contrast, a 2-year life cycle is more likely in R. gigas. However, life cycle durations of all species studied are still uncertain and regional differences are very probable.     

Three new species of Austrophlebioides Campbell and Suter (Ephemeroptera: Leptophlebiidae: Atalophlebiinae) from the Wet Tropics bioregion of north-eastern Australia

Abstract  Three new species of the mayfly genus Austrophlebioides Campbell and Suter are described from the Wet Tropics bioregion of north-eastern Australia: A . rieki sp. n., A . wooroonooran sp. n. and A . porphyrobranchus sp. n. The three species are similar, and are characterised in the male imago by the presence of a prominent ventral projection on each lobe of the penes and segment one of the claspers narrowing at about one-third length, and in the nymph by the absence of fine setae along the outer margin of the mandible between the median setal tuft and the outer incisor. The generic diagnosis of the nymphal stage is modified slightly to accommodate the three new species.     

Comparative Study of the Nymphal Biology of Two Coexisting Species of Mayflies (Insecta: Ephemeroptera) in a Mediterranean Stream in Southern Europe

We study the life history, nymphal feeding and secondary production of two leptophlebiid mayfly species (Habrophlebia eldae and Paraleptophlebia submarginata). They cohabit in a Mediterranean stream and present a very high niche overlap in terms of trophic resources. The life cycle was estimated using size‐frequency analysis of samples taken throughout a year. Both species have a similar but displaced period of the nymphal development. Secondary production was calculated by means of the size‐frequency method. Annual secondary production of P. submarginata is much higher than that of H. eldae (1.95 g DW m^–2 year^–1 vs. 0.17 g DW m^–2 year^–1), and presents a quite similar annual P/B ratio, but slightly higher in ^{P. submarginata} (6.97 in ^{P. submarginata} and 9.21 in ^{H. eldae}). The study of the gut contents revealed that they are mainly detritivores but, when larger they feed also on CPOM from leaves fallen in the stream. They present an almost total niche overlap in terms of food acquisition. However the previously mentioned shift in trophic resources utilization with size makes it possible that, because no similar size classes of each species are present at the same time, niche segregation exists between the two species. Though further studies are needed to confirm it, this could be the consequence of previous episodes of competition between them. (© 2010 WILEY‐VCH Verlag GmbH & Co. KGaA, Weinheim)     

Evolution of complex life cycles of amphibians: bridging the gap between metapopulation dynamics and life history evolution

Current evolutionary models for amphibian life cycles reflect tradeoffs in size-specific growth and mortality rates between the aquatic and terrestrial stages. A limitation of these models is that they do not incorporate evolutionary phenomena that are associated with metapopulation structure. In this work I address components of the evolution of complex life cycles (CLCs) that are tied to the metapopulation dynamics of amphibians that use seasonal wetlands that vary in hydroperiod. In particular, I describe how selection for the minimum length of the larval period affects metapopulation viability and the selection/migration equilibrium. Selection to increase the minimum length of the larval period functionally reduces the number of viable breeding sites on the landscape, increases the average distance between neighboring sites, and increases the risk of metapopulation extinction. Within a metapopulation, asymmetric gene flow between populations that are adapted to different hydroperiods tends to swamp local selection for long larval periods at sites with long hydroperiods. The evolutionary stability of CLCs of many species with metapopulation structure may reflect the fact that extremely small metamorphs cannot survive on land, while lineages with long larval periods incur a high risk of metapopulation extinction. I encourage theorists to more carefully consider how life history traits and metapopulation viability are related for these and other taxa.     

Mayfly fauna (Ephemeroptera) and the biology of the species Potamanthus luteus (L.) in a warmed stretch of the Oslava River

Cooling waters of the Oslavany power station show a depressive effect on the species diversity of mayfly nymphs. Only seven of 35 taxons found in the study area occurred in all localities. In the warmed stretch of the river, the abundance and biomass of mayfly nymphs as well as the production of the most important speciesPotamanthus luteus decreased; a gradual regeneration of the disturbed mayfly population could be observed downstream. In Locality 5, situated 3.5 km downstream from the mouth of cooling waters, the species diversity, abundance, and biomass of mayflies were 69, 30, and 66 per cent, resp., of values found in Locality 1 with natural temperatures; the production ofPotamanthus luteus was 70 per cent of that found in Locality 1.     

Animated life cycles of fungi and plants with spores for teaching

The conveyance of knowledge of the life cycles performed by fungi and plants with spores is a challenge for teaching in university education. The life cycles of fungi, in particular, can be very complex and difficult to understand. This paper presents the development and implementation of a German and English educational software program, including seven animated life cycles of fungi, two animated developments of fungal meiosporangia, and two examples each for ferns, mosses, and algae (cryptogams). We could show that the use of the software supported the study of fungi and spore-producing plants. It was possible to identify attributes of usability that contributed to a high quality of the educational material in concrete teaching situations. Notably, an easy handling of the software and an appropriate amount of information per animation step are crucial. In addition to the didactic insights, the conversations between the animation designers and the mycologist on correct details of developmental processes in the animations lead to interesting questions concerning cell biology. Some of the resulting open questions are at the edge of currently available knowledge, e.g. concerning the transformation of a plasmodium into a high number of small fruiting bodies in myxomycetes or the dikaryotisation of monokaryotic hyphae after spermatisation in rust fungi.     

Temperature-dependent growth and life cycle of Deleatidium (Ephemeroptera: Leptophlebiidae) in two high-country streams in New Zealand

1. Measurements of larval growth rates were used to produce an empirical model for examining factors influencing patterns of size structure and emergence period for populations of the leptophlebiid mayfly Deleatidium ( lillii group) in two high-country streams (South Island, New Zealand). Growth rates were measured in situ by enclosing groups of larvae and natural stream substrata in growth chambers.
2. Multiple regression analysis showed that temperature explained most variation among growth rates in both streams. Growth rates were also significantly related to larval size, although the effect was minor compared with temperature. A significant relationship between larval biomass per chamber and growth rate was shown at only one site.
3. Growth models based on multiple regression equations ( R ² = 0.70–0.84) simulated the extended emergence period and complex size structure observed for populations of Deleatidium in the field. Larvae hatching from eggs deposited before mid-February (austral summer) probably emerge as adults before May (minimum cohort duration ≈ 3 months). Larvae hatching after mid-February probably do not emerge as adults until the following summer because of low growth rates during winter (maximum cohort duration ≈ 11 months).
4. On average, there are probably two generations of Deleatidium per year (bivoltine) at the study sites. The presence of numerous overlapping cohorts throughout summer, however, results in an extended emergence period and complex size structure.     

Nymphal biology of Ephoron virgo (Olivier, 1791) (Ephemeroptera,Polymitarcyidae) in an Apenninic river (NW Italy)

Life cycle, feeding habits and nymphal density of a population of Ephoron virgo from the Bormida river (Northwestern Italy) were studied. Nymphs were present in the river from the beginning of May until August, with the highest density of nymphs recorded at the end of May–beginning of June. The life cycle was univoltine and the nymphal growth was fast. During the life cycle, an obligatory egg diapause occurs and individuals remain in this stage for up to nine months. Nymphs fed mainly on detritus, although mineral matter constituted a considerable percentage of the gut contents, predominantly in smaller nymphs.     

Temperature-dependent growth and life cycle of Deleatidium (Ephemeroptera: Leptophlebiidae) in two high-country streams in New Zealand

1. Measurements of larval growth rates were used to produce an empirical model for examining factors influencing patterns of size structure and emergence period for populations of the leptophlebiid mayfly Deleatidium ( lillii group) in two high-country streams (South Island, New Zealand). Growth rates were measured in situ by enclosing groups of larvae and natural stream substrata in growth chambers.
2. Multiple regression analysis showed that temperature explained most variation among growth rates in both streams. Growth rates were also significantly related to larval size, although the effect was minor compared with temperature. A significant relationship between larval biomass per chamber and growth rate was shown at only one site.
3. Growth models based on multiple regression equations ( R ² = 0.70–0.84) simulated the extended emergence period and complex size structure observed for populations of Deleatidium in the field. Larvae hatching from eggs deposited before mid-February (austral summer) probably emerge as adults before May (minimum cohort duration ≈ 3 months). Larvae hatching after mid-February probably do not emerge as adults until the following summer because of low growth rates during winter (maximum cohort duration ≈ 11 months).
4. On average, there are probably two generations of Deleatidium per year (bivoltine) at the study sites. The presence of numerous overlapping cohorts throughout summer, however, results in an extended emergence period and complex size structure.     

Relationship between benthic insects (Ephemeroptera, Plecoptera, Coleoptera, Trichoptera) and temperature in small and medium-sized streams in Germany: A multivariate study

Benthic insect communities (Ephemeroptera, Plecoptera, Coleoptera, Trichoptera) were studied together with water temperature and environmental parameters in streams between June 2000 and June 2001. The sampling area consisted of 20 sites in small and medium-sized streams located in the lower mountainous area of Central Europe. Temperature was recorded nearly continuously and several physicochemical and environmental variables were assessed. Macroinvertebrates were sampled both in spring and summer. Data-sets of species abundance and occurrence were analysed using multivariate techniques and were correlated to the thermal and environmental conditions of the streams. The temperature preferences of the species were compared to published data-sets on their autecological characteristics. Up to 29% of the variability in the Ephemeroptera, Plecoptera, Trichoptera and Coleoptera community was explained by summer temperature variation in the data-sets for both small and medium-sized streams. A smaller, but significant part of the variability in species distribution was explained by conductivity, substratum type, and the percent coverage of local riparian forest. Compared to small streams, temperature was less important for the macroinvertebrate composition in medium-sized streams. This result is likely due to the more tolerant, eurythermic species composition in larger streams. A total of 33 Ephemeroptera, Plecoptera, Coleoptera and Trichoptera taxa were positively correlated and 28 taxa were negatively correlated to summer temperature patterns. The temperature preferences of taxa considered in this study were related to species traits, such as egg dormancies and life cycle plasticity.     

Longitudinal zonation and life cycles of macrozoobenthos in the Mauerbach near Vienna,Austria

A total of 12277 benthic invertebrates were caught from November 1990 to October 1991 at the Mauerbach, a first to fourth order forest brook near Vienna, Austria, using a Surber sampler and a hand net. Five taxa comprised 96.6% of the catch: Diptera larvae (45.6%), Amphipoda (37.6%) and larvae of Ephemeroptera (6.4%), Plecoptera (5.3%) and Trichoptera (1.7%). Mean values of macrozoobenthos density ranged from 39514 specimens m⁻² at upstream sampling site 5 in December to only 286 specimens m⁻² at sampling site 3 in May, shortly after a severe flood. Based on benthic invertebrate population structure, sites 5 and 6 (situated near the source) were clearly separated from the downstream sites 1 to 4. At upstream sites, shredders comprised up to 71.7% of the total, whereas at downstream sites collectors were most abundant. In addition, the proportion of eucrenal species decreased from 12% at site 6 near the source to only 2% at site 1 near the mouth. Among the insect species studied in detail, most were univoltine except Ephemera danica (Ephemeroptera) and Sericostoma personatum (Trichoptera), which had a two-year life cycle.     

Aflagellate sperm in three species of leptophlebiidae (Ephemeroptera)

Immotile spermatozoa of 3 species of Ephemeroptera, Habroleptoides umbratilis, Habrophlebia eldae and Choroterpes picteti (Leptophlebiidae) are described. In all 3 species, sperm present a coccoidal shape and lack flagella or microtubule systems. Habrophlebia eldae shows very atypical sperm with a roundish nucleus and scarce cytoplasm, including an apical acrosome. By contrast, a fibrillar perforatorium and a mitochondrion are present in the sperm of H. umbratilis and C. picteti. This latter species also shows electron-dense bodies located in the space between the nucleus and the cell membrane. Our findings suggest that sperm/egg interaction should depend in leptophlebiids on the contraction of the genital duct muscles.     

The Ecology of Baetis muticus and Baetis rhodani (Insecta,Ephemeroptera), with Special Emphasis on Acid Water Backgrounds

The two mayflies (Insecta, Ephemeroptera) Baetis muticus and Baetis rhodani are absent from the acid water (pH 4.8–5.2) streams of the Upper Duddon drainage basin. Field observations and laboratory experiments were used in an attempt to explain this absence. Baetis muticus could not tolerate acid water; in addition it was deduced that available foods in the Upper Duddon streams would not sustain it there, even if the water could be tolerated. Baetis muticus feeds by browsing on decaying, allochthonous leaves, implying an input of micro-organisms, especially fungi, to the diet. Well-decayed, allochthonous leaves do not occur in the acid water, Upper Duddon streams. Baetis rhodani proved to be acid water tolerant, but only if the background waters had high ionic loadings. Since Upper Duddon waters have only low ionic loadings, Baetis rhodani is apparently excluded from them on this account. The evidence was that if the Upper Duddon waters were chemically suitable, then Baetis rhodani could survive in them, because a suitable food, the alga Hormidium subtile is present.     

Host relations, life cycles and multiparasitism in some parasitoids of aphidophagous Syrphidae (Diptera)

Abstract. 1. Records of parasitism from the field and host choice experiments suggest that most parasitoids of syrphids are monophagous. A few are oligophagous.
2. To elicit egg release, females require a stimulus from the host haemolymph. Differential response to haemolymph cues by females may account for the observed pattern of host relations.
3. Differences in host ranges and timing of female flight periods probably characterize most of the parasitoids studied.
4. In one case of two monophagous parasitoids, Diplazon pectoratorius (Thunberg) and Syrphophilus tricinctorius (Thunberg) attacking the same host Syrphus ribesii (L.) no partitioning was found. Stricinctorius is, however, a superior intrinsic competitor.     

Two New Species of Afronurus (Ephemeroptera: Heptageniidae) from Vietnam

ABSTRACT Two new species of the heptageniid mayfly genus Afronurus Lestage, Afronurus meo sp. n. and Afronurus mnong sp. n., are described from Vietnam with line-drawings of key characters. The larvae of A. meo can be characterized by rowed minute spines in the posterior margin of the abdominal terga 1-9 and distinct markings on anterior head, dorsal femora, and abdominal terga. The larvae of A. mnong can be characterized by scattered simple stout setae on the anterior margin and dorsal surface of femora, rowed acute spines on the posterior margin of abdominal terga 1-9, and plain brown body color. Their diagnoses, material data, distributions, and habitat and biological data are provided.