Effects of organic matter and season on leaf litter colonisation and breakdown in cave streams

1. Low organic matter availability is thought to be a primary factor influencing evolutionary and ecological processes in cave ecosystems. We examined links among organic matter abundance, macroinvertebrate community structure and breakdown rates of red maple (Acer rubrum) and corn litter (Zea mays) in coarse‐ (10 × 8 mm) and fine‐mesh (500‐μm) litter bags over two seasonal periods in four cave streams in the south‐eastern U.S.A. 2. Organic matter abundance differed among cave streams, averaging from near zero to 850 g ash‐free dry mass m^?2. Each cave system harboured a different macroinvertebrate community. However, trophic structure was similar among caves, with low shredder biomass (2–17% of total biomass). 3. Corn litter breakdown rates (mean k = 0.005 day^?1) were faster than red maple (mean k = 0.003 day^?1). Breakdown rates in coarse‐mesh bags (k = 0.001–0.012 day^?1) were up to three times faster than in fine‐mesh bags (k = 0.001–0.004 day^?1). Neither invertebrate biomass in litter bags nor breakdown rates were correlated with the ambient abundance of organic matter. Litter breakdown rates showed no significant temporal variation. Epigean (surface‐adapted) invertebrates dominated biomass in litter bags, suggesting that their effects on cave ecosystem processes may be greater than hypogean (cave‐adapted) taxa, the traditional focus of cave studies. 4. The functional diversity of our cave communities and litter breakdown rates are comparable to those found in previous litter breakdown studies in cave streams, suggesting that the factors that control organic matter processing (e.g. trophic structure of communities) may be broadly similar across geographically diverse areas.     

Life histories, production dynamics and resource utilisation of mayflies (Ephemeroptera) in two tropical Asian forest streams

SUMMARY 1. A 2‐year study of the life histories, production dynamics and resource utilisation of five mayfly species was undertaken in two forest streams in Hong Kong [Tai Po Kau Forest Stream (TPKFS) and Shing Mun River (SMR)]. Afronurus sp. and Cinygmina sp. (Heptageniidae), Procloeon sp. and Baetiella pseudofrequenta (Baetidae), and Choroterpes sp. (Leptophlebiidae) were abundant in both streams and contributed more than 50% of the total mayfly populations. 2. All species had asynchronous larval development with recruitment occurring throughout the year. Mean annual production (all mayflies combined) was 3.1 and 2.0 g dry weight m^?2 year^?1 in SMR and TPKFS, respectively – the higher value at SMR reflecting greater mayfly densities – with more than 70% of production occurring during the wet season. Mayfly production varied between years, decreasing by 5% in TPKFS and 43% in SMR during 1996–97, reflecting lower densities of heptageniids relative to 1995–96. Annual biomass turnover rates (P/B) were high in both sites ranging from 27.2 to 94.6 in TPKFS (Cinygmina sp. and Procloeon sp.) and from 31.8 to 109.8 in SMR (Cinygmina sp. and B. pseudofrequenta). 3. Patterns of daily production in both streams showed that Afronurus sp., Cinygmina sp. and Choroterpes sp. were most productive during the wet season, while Procloeon sp. maintained high production levels throughout the year. The highest daily production of B. pseudofrequenta occurred during the wet season in TPKFS, but in the dry season at SMR. Temporal overlap in production and hence resource utilisation in both streams, calculated using the proportional similarity index (PS), ranged from 0.39 to 0.81. It was highest (0.63–0.81) between pairs of species of Heptageniidae and Baetidae, and lowest between Choroterpes sp. and other mayflies (0.39–0.61). No clear temporal segregation was observed among any species. However, when using the fraction of production attributable to each food, lower PS values were obtained for all species in both sites. In SMR, trophic segregation may have occurred between the two species pairs Procloeon sp.–Cinygmina sp. and Procloeon sp.–Choroterpes sp. (PS=0.17 and 0.03, respectively). 4. A combination of production data and information on the stable isotope signature of mayflies revealed that, during both the wet and dry seasons, more than 50% of total mayfly production in TPKFS was derived from autochthonous foods. In SMR, 68% of production was supported by allochthonous foods during the wet season, and 72% by autochthonous sources in the dry season. Considering that more than 70% of the total production occurred in the wet season, the trophic basis of mayfly production in SMR is mostly allochthonous (58%) while in TPKFS it is mainly of autochthonous origin (66%). The year‐round importance of autochthonous foods in shaded streams such as TPKFS is surprising, but the wet season contribution of allochthonous foods (especially in SMR) may have resulted from depletion of algal biomass during spates.     

Life‐history constraints in grassland plant species: a growth‐defence trade‐off is the norm

Plant growth can be limited by resource acquisition and defence against consumers, leading to contrasting trade‐off possibilities. The competition‐defence hypothesis posits a trade‐off between competitive ability and defence against enemies (e.g. herbivores and pathogens). The growth‐defence hypothesis suggests that strong competitors for nutrients are also defended against enemies, at a cost to growth rate. We tested these hypotheses using observations of 706 plant populations of over 500 species before and following identical fertilisation and fencing treatments at 39 grassland sites worldwide. Strong positive covariance in species responses to both treatments provided support for a growth‐defence trade‐off: populations that increased with the removal of nutrient limitation (poor competitors) also increased following removal of consumers. This result held globally across 4 years within plant life‐history groups and within the majority of individual sites. Thus, a growth‐defence trade‐off appears to be the norm, and mechanisms maintaining grassland biodiversity may operate within this constraint.     

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.     

Life history and secondary production of Glossosoma nigrior Banks (Trichoptera: Glossosomatidae) in two Alabama streams with different geology

We studied life history and secondary production of a caddisfly scraper, Glossosoma nigrior, in two Alabama streams. Collier Creek, located within the Appalachian Plateau physiographic province, is underlain by sandstone bedrock, while Hendrick Mill Branch is located in the Valley and Ridge physiographic province with limestone bedrock. G. nigrior populations in both streams exhibited trivoltine life histories, which were attributed to the higher water temperature regimes than those found in more northern streams. Mean larval density (556 m⁻²) and biomass (B) (49.2 mg AFDM m⁻²) were much higher in Hendrick Mill Branch than Collier Creek (78 m⁻² and 6.7 mg AFDM m⁻²). G. nigrior in Hendrick Mill Branch maintained continuous larval growth and higher larval density than Collier Creek throughout the year mainly due to a greater availability of optimal habitat, a more stable hydrology, and warmer winter water temperature. These factors also resulted in the much higher annual secondary production (P) in Hendrick Mill Branch (965 mg AFDM m⁻²; P/B = 18.3) than Collier Creek (115 mg AFDM m⁻²; P/B = 17.9). Gut content analysis revealed that algae (>50%) and detritus (>40%) were the major diet items for G. nigrior, and the majority of secondary production (>80%) was contributed by the consumption of algae. Glossosoma populations play an important role in trophic linkage in these streams with their high production and grazing activities. Handling editor: D. Dudgeon     

Life history and life table analysis of the whitefly predator Delphastus catalinae （Coleoptera： Coccinellidae） on collards

Abstract The ladybeetle, Delphastus catalinae (Horn), is one of the most commonly used predacious natural enemies being commercially reared for controlling whiteflies, including Bemisia tabaci (Gennadius) biotype B (= B. argentifolii Bellows & Perring), on various ornamental and vegetable crops under greenhouse conditions. The development, survivorship, and fecundity of D. catalinae feeding on B. tabaci biotype B on collard plants were determined in the laboratory, and the age-specific life table parameters were analyzed based on the life history data. Developmental time was 4.0, 1.9, 1.1, 1.4, 5.2, and 5.3 days for eggs, first, second, third, fourth instars, and pupae, respectively, with an average of 18.9 days from oviposition to adult emergence for both sexes, 19.0 days for females, and 18.8 days for males. Adult longevities averaged 146.6 days for both sexes, 122.6 days for females, and 170.5 days for males. After an average 4.9 days preoviposition period, females laid a mean of 5.6 eggs per day over a 97.0-day period. Net reproductive rate ( R ₀) and gross reproductive rate (ΣM m_x ) were estimated by life table analysis at 276.8 and 325.1, respectively. Generation time (T) and doubling time (DT) were 35.6 and 4.8 days respectively, and the intrinsic rate of natural population increase ( r_m ) was estimated at 0.158, or l = 1.171 for the finite rate of increase. The r_m value of D. catalinae is similar to or higher than those of the whitefly feeding on most vegetable and ornamental crops, indicating that the ladybeetle is capable of regulating populations of B. tabaci biotype B and other whiteflies under greenhouse conditions.     

Life history differences of Psilotreta locumtenens (Trichoptera: Odontoceridae) in two reaches of a mountain stream in Korea

The life history traits of the caddisfly, Psilotreta locumtenens Botosaneanu (Odontoceridae), were studied in two stream reaches with different thermal ranges (main and branch streams) of the Gapyeong Stream, a typical mountain stream located in the central Korean Peninsula. Psilotreta locumtenens larvae were quantitatively sampled monthly from November 2008 to July 2010, and biweekly during the emergence period (late April to early July), using a Surber sampler (30 × 30 cm). Adults were quantitatively sampled with a sweep net. Larval density in the main stream (324.21 ± 38.59 m^?2) was higher than that in the branch stream (60.48 ± 10.86 m^?2). The larvae hatched in the early summer and overwintered as 5th and 3rd instars in the main and branch streams, respectively. The emergence peak at the main stream was approximately 2 weeks earlier. The sex ratio at both sites was approximately 0.3. The life history in both streams was univoltine. Secondary production in the main stream was much higher than in the branch stream, owing to high larval densities, and the P/B ratios at the two sites were similar. This study demonstrated remarkable differences in larval growth patterns and emergence peaks in P. locumtenens between the two stream reaches due to differences in accumulated degree days (230.30 DD) and other phenological cues such as daily mean threshold water temperature (9°C) during the ascending phase, despite their relatively small mean annual water temperature difference of 0.58°C.     

Control of periphyton standing crop in an Atlantic Forest stream: the relative roles of nutrients,grazers and predators

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Life history and production of the burrowing mayfly Ephoron virgo (Olivier, 1791) (Ephemeroptera: Polymitarcyidae) in the lower Ebro river: a comparison after 18 years

The life history of the burrowing mayfly Ephoron virgo (Olivier, 1791) (Ephemeroptera: Polymitarcyidae) was studied during spring and summer 2005 in the lower Ebro river (Catalonia) and compared to a previous study performed in 1987 (Ibáñez, Escosa, Muñoz and Prat 1991). The results showed an advancement of Ephoron virgo life cycle and an increase of production estimates. In 2005 larval development reached the maximum size one month earlier than in 1987, and adult emergence peak began three weeks earlier. Comparing adult sex ratios (F:M), there was a major presence of females in 2005 (1:4), while the opposite was observed in 1987 (2:1). Secondary production was higher in 2005 than in 1987, obtaining 950 mg dry weight/m²/year with the increment summation method and 1080 mg dry weight/m²/year using the removal summation method. Higher water temperatures were measured for the entire 2005 larval growth period, which were related to higher air temperatures. Therefore, that temperature increment was likely the main cause of changes observed in the Ephoron virgo life cycle.     

Pollination limitation,costs of capsule production and the capsule-to-flower ratio in Dendrobium monophyllum F. Muell. (Orchidaceae)

Abstract In perennial plants, life-history theory suggests that natural selection should result in the optimization of fruit-to-flower ratios within the limits imposed by the trade-offs between resource allocation for present reproduction and future growth and reproduction. The tropical orchid Dendrobium monophyllum F. Muell., an epiphyte or lithophyte, offers no nectar rewards, is self-incompatible and has a capsule-to-flower ratio of about 1:14. The influence of pollination limitation and the costs of capsule production on capsule-to-flower ratios were assessed using experimental and field studies in which individual plants were observed for 3 years. Pollinators visited about 80% of flowers, and capsule production was significantly related to inflorescence size and pollinaria removal. About nine pollinator visits occurred per capsule. Pollinator visitation and capsule production did not vary significantly between years. The inflorescence size classes most successful in capsule production were also the most frequent in natural populations. The experimental supplementation of outcross pollen to flowers increased capsule set over controls by 45% within a year, but was limited to about 53%. A capsule-to-flower ratio of 1:2 in experimental plants significantly decreased the subsequent growth and flowering of individuals relative to controls. A capsule-to-flower ratio above 1:10 in naturally pollinated plants decreased flowering in the subsequent year. Thus, it is suggested than an increase in capsule production above 10% would not necessarily correlate with greater reproductive fitness because of the increased cost of capsule production. The capsule-to-flower ratio recorded in this study could be evolutionarily stable because of trade-offs between selection for pollinator attraction and the cost of capsule production. The production of surplus flowers appears to function in pollinator attraction and increases fitness through male function.     

Effects of nutrient enrichments on primary production in the Ria Formosa coastal lagoon (Southern Portugal)

Small-scale, short-term enrichment experiments were conducted in the Ria Formosa coastal lagoon (southern Portugal), to assess the effects of nutrient availability on primary productivity, biomass (as chlorophyll a), and algal composition. Samples were collected from natural communities at three different sites in the western lagoon: Barra, oceanic inlet; Ramalhete, adjacent to a urban waste water treatment plant; and Ponte, an upstream channel used for recreation and bivalve farming. These samples were enriched separately with nitrogen and phosphorous during the extreme neap tides of the summer solstice at both high (HW) and low water (LW). The experiment was repeated during the autumnal equinox to test for seasonality, and during the following summer solstice to test for replication. The addition of nitrogen consistently stimulated the productivity and biomass during summer experiments at the two sites within the lagoon, identifying N as the most likely primary “potentially limiting nutrient” in the western part of the lagoon for this period. No stimulation of biomass and productivity occurred in September at the same two sites indicating the importance of other factors such as light, sedimentation or grazing pressure, as controlling the pelagic community. However, these outcomes were reversed at the oceanic inlet (Barra-HW) where there was no stimulation by nitrogen during the summer months, but there was in September, suggesting that there is a different nutritional requirement for the coastal community in comparison with the lagoon community. In samples where productivity was stimulated, diatoms were the group most sensitive to enrichment.     

The influence of water level on macrophyte growth and trophic interactions in eutrophic Mediterranean shallow lakes: a mesocosm experiment with and without fish

1. Water‐level fluctuations are typical of lakes located in the semi‐arid Mediterranean region, which is characterised by warm rainy winters and hot dry summers. Ongoing climate change may exacerbate fluctuations and lead to more severe episodes of drought, so information on the effects of water level on the functioning of lake ecosystems in such regions is crucial. 2. In eutrophic Lake Eymir, Turkey, we conducted a 4‐month (summer) field experiment using cylindrical 0.8‐m‐ (low‐water‐level) and 1.6‐m‐deep (high‐water‐level) mesocosms (kept open to the sediment and atmosphere). Fish (tench, Tinca tinca, and bleak, Alburnus escherichii) were added to half of the mesocosms, while the rest were kept fishless. Ten shoots of Potamogeton pectinatus were transplanted to each mesocosm. 3. Sampling for physicochemical variables, chlorophyll a (chl‐a), zooplankton and per cent plant volume inhabited (PVI%) by macrophytes was conducted weekly during the first 5 weeks, and subsequently biweekly. Macrophytes were harvested on the last sampling date. During the course of the experiment, the water level decreased by 0.41 ± 0.06 m. 4. Throughout the experiment, fish affected zooplankton abundance (?), nutrient concentrations (+), chl‐a (+) and water clarity (?) most strongly in the low‐water‐level mesocosms and the zooplankton community shifted towards dominance of small‐sized forms. The fishless mesocosms had a higher zooplankton/phytoplankton ratio, suggesting higher grazing. 5. Greatest macrophyte growth was observed in the low‐water‐level fishless mesocosms. However, despite high nutrient concentrations and low water clarity, macrophytes were also abundant in the fish mesocosms and particularly increased following a water‐level decrease from midsummer onwards. Macrophyte growth was poor in the high‐water‐level mesocosms, even in the fishless ones with high water clarity. This was ascribed to extensive periphyton development reducing light availability for the macrophytes. 6. Our results indicate that a reduction in water level during summer may help maintain the growth of macrophytes in Mediterranean eutrophic shallow lakes, despite a strong negative effect of fish predation on water clarity. It is therefore probable that an expected negative effect of global climate change on water clarity because of eutrophication and enhanced top‐down control of fish may be, at least partly, counteracted by reduced water level, provided that physical disturbance is not severe.     

Effects of gall volume on survival and fecundity of gall-making aphidsAdelges japonicus (Homoptera: Adelgidae)

The number of larvae reaching maturity within the gall of Adelges japonicus was positively related to gall volume, and the relationship between the number of mature larvae and gall volume did not change with different densities of colonized larvae. The population changes in the bud galls of A. japonicus were surveyed by collecting the galls, which did not suffer predation or parasitism within the galls, from young artificial plantations of Picea jezoensis over two years. In the year when the density of colonized larvae was high, they suffered a 42% mortality within the galls, whereas mortality was nearly zero in the low density year. The numbers of larvae per gall were positively correlated with gall volume. The regression lines of the number of colonized larvae on gall volume did not differ significantly in the regression coefficients between the two years, whereas the intercept was significantly higher in the year when the density of colonized larvae was higher. However, different within-gall mortality nullified this difference, and the regression lines of the number of mature larvae on gall volume had no significant difference both in the regression coefficients and the intercepts. This suggests that the number of mature larvae per gall was limited by available resources within the gall which were positively related to gall volume. In 25% of the galls in which mature larvae inhabited, the space within the galls were completely filled by the larvae, indicating that space was one of the limiting resources. Gall volume also affected the number of adults that emerged from the gall and the potential number of their progeny.     

Bacterial abundance and production in river sediments as related to the biochemical composition of particulate organic matter (POM)

The major proportion of heterotrophic activity in running waters islocalized on the solid surfaces of sediments in the benthic and hyporheic zoneand is dominated by microorganisms. However, this assertion is based on thestudies of small streams, and little is known about the microbial metabolism oforganic matter in river ecosystems. We therefore explored the relationshipsbetween bacterial abundance and production and the gradients of organic matterquality and quantity in sediments of a sixth-order lowland river (Spree,Germany). We found vertical gradients of detrital variables (particulateorganicmatter (POM), particulate organic carbon (POC), nitrogen (PN), and protein) andof bacterial variables (abundance, production, turnover time, and proportion ofbacterial carbon in total POC) in two different sediment types. These gradientswere steeper in stratified sediments than in the shifting sediments. Detritalvariables correlated strongly with bacterial abundance and production. The bestcorrelation was found for detrital variables indicating substrate quantity andquality (r_S = 0.90 for PN with abundance). Although bacterialbiomasscomprised only 0.7% of the POC (1.9% of PN, 3.4% of the protein) in sediments,the turnover of sedimentary organic carbon was fast (median = 62d), especially in the shifting sediments. Our findings demonstratethat sediment dynamics significantly foster organic carbon metabolism in riversystems. Thus, these sediments, which are typical for lowland rivers, stronglyinfluence the metabolism of the whole ecosystem.     

Effects of early resource limitation and compensatory growth on lifetime fitness in the ladybird beetle (Harmonia axyridis)

Acceleration of growth following a period of diet restriction may result in either complete or partial catch-up in size. The existence of such compensatory growth indicates that organisms commonly grow at rates below their physiological maxima and this implies a cost for accelerated growth. We examined patterns of accelerated growth in response to temporary resource limitation, and assayed both short and long-term costs of this growth in the ladybird beetle Harmonia axyridis. Subsequent to the period of food restriction, accelerated growth resulted in complete compensation for body sizes, although we observed greater larval mortality during the period of compensation. There were no effects on female fecundity or survivorship within 3 months of maturation. Females did not discriminate against males that had undergone compensatory growth, nor did we observe effects on male mating behaviour. However, individuals that underwent compensatory growth died significantly sooner when deprived of food late in adult life, suggesting that longer-term costs of compensatory growth may be quite mild and detectable only under stressful conditions.     

Life history and production of Agapetus quadratus (Trichoptera: Glossosomatidae) in a temporary, spring-fed stream

1. The life history and trophic basis of production of the caddisfly grazer Agapetus quadratus were studied in the torrent Gorg Blau, a spring‐fed stream on the island of Majorca that dries annually during summer. 2. Quantitative random samples were taken every 2–3 weeks during an annual surficial flow period, from November 2000 to mid‐July 2001. Instars of field‐collected larvae were determined by measurements of head width and pronotum length, and the sex of all pupae was determined to study sexual dimorphism and sex ratio. 3. Stage‐frequency histograms suggested a trivoltine population, with an average cohort time of 4 months. Larval development was asynchronous, with continuous growth and overlapping generations. Recruitment peaks were identified in mid‐November, early March and late June, indicative of winter, spring and summer generations. On average, females were larger than males and the mean sex ratio was 2 : 3 (females : males). Population densities and biomasses derived from the field data were used to calculate production and turnover rate. 4. Annual production of A. quadratus in the torrent Gorg Blau (4.80 g dry mass m^?2 year^?1) was the highest ever reported for the genus, being comparable with that estimated for some insects with rapid development and multiple cohorts. 5. Estimates of production of A. quadratus were combined with foregut content analysis to estimate the fraction of total production derived from the principal food sources: algae and organic detritus. Algae supported a major proportion of the production of this grazer. 6. The low density of predators characteristic of many temporary streams, and the small amplitudes in discharge and temperature during most of the wet period that characterise the spring habitats might allow high levels of grazer production in this particular Mediterranean stream.     

Effects of variation in resource acquisition during different stages of the life cycle on life‐history traits and trade‐offs in a burying beetle

Individual variation in resource acquisition should have consequences for life‐history traits and trade‐offs between them because such variation determines how many resources can be allocated to different life‐history functions, such as growth, survival and reproduction. Since resource acquisition can vary across an individual's life cycle, the consequences for life‐history traits and trade‐offs may depend on when during the life cycle resources are limited. We tested for differential and/or interactive effects of variation in resource acquisition in the burying beetle Nicrophorus vespilloides. We designed an experiment in which individuals acquired high or low amounts of resources across three stages of the life cycle: larval development, prior to breeding and the onset of breeding in a fully crossed design. Resource acquisition during larval development and prior to breeding affected egg size and offspring survival, respectively. Meanwhile, resource acquisition at the onset of breeding affected size and number of both eggs and offspring. In addition, there were interactive effects between resource acquisition at different stages on egg size and offspring survival. However, only when females acquired few resources at the onset of breeding was there evidence for a trade‐off between offspring size and number. Our results demonstrate that individual variation in resource acquisition during different stages of the life cycle has important consequences for life‐history traits but limited effects on trade‐offs. This suggests that in species that acquire a fixed‐sized resource at the onset of breeding, the size of this resource has larger effects on life‐history trade‐offs than resources acquired at earlier stages.     

Effects of the presence of a predatory fish and the phenotype of its prey (a shredding shrimp) on leaf litter decomposition

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