An algae — Desert snail food chain: Energy flow and soil turnover

Summary The feeding ecology of adult desert snails (Sphincterochila zonata), inhabiting a loess plain in the Negev Desert, was studied during a 7-year period. Energy flow and soil turnover determinations were made at the individual and population level on the basis of field observations and data derived from laboratory simulations. Sphincterochila zonata were only active on 8–27 winter days annually, otherwise they were dormant. The snails fed exclusively on algae that grew on the soil surface following rain. The mean annual dry-weight biomass of algae was 197.4(±118.1) gr·m^-2. The dry weight biomass of snails ranged from 0.02–0.14 gr·m^-2. Annual production in the food chain varied substantially from year to year (1–1539 Kj·m^-2), but the net annual energy balance of the snails was always positive. Soil crust turnover, resulting from grazing of snails on algae, was estimated at 142 kg·hectare^-1 during the study period.The annual magnitude of energy flow and soil turnover is determined by the soil surface moisture regime which in turn, is a function of rainfall patterns. Sphincterochila zonata may serve an important role in Negev ecosystems by dispersing soil algae and altering soil structure. The snails are not subject to substanding predation by rodents but may occasionally serve as an important food source for migrating birds.     

The productivity and carbon budget of a natural population of Daphnia lumholtzi Sars

D. lumholtzi in Lake Samsonvale, Queensland, Australia, is a small species (max. size approx. 7 µgC) that occurs in low abundance (max. abundance 6400 m^–3), with an average daily biomass of 3.32 mgC m^–3. Its annual rates of carbon assimilation, production and respiration, are 166, 110, and 56 mgC m^–3 y^–1 respectively. Annual biomass turnover (annual production/average daily biomass) is 33 and production efficiency is 50–66%. The population may consume 1.65–2.20 mgC m^–3 daily, equivalent to about 1% of the average daily standing crop of phytoplankton. Clutch size is small, 2 eggs, but represents 30–80% of a female's weight. A female may only produce 8–10 offspring in a full lifespan, nevertheless egg production may account for 56% of total production. The population shows autumn and spring peaks in abundance, and is believed to oversummer (4 months) as ephippia.     

Distribution and production ofChironomus salinarius (Diptera: Chironomidae) in a shallow coastal lagoon in the Bay of Cádiz

The abundance, generation time and production ofChironomus salinarius larvae in a lagoon fish-pond system in the Bay of Cádiz were studied by taking monthly samples at 3 sites during 1991 and 1992. Numerical abundance and biomass of larvae showed considerable spatial, seasonal and interannual variation (ANCOVAs,P<0.001). The maximum mean annual density was 7048 larvae m^–2, and corresponded to a biomass of 3.08 g dry weight (DW) m^–2. It was recorded at the site with the lowest rate of water renewal. Seasonal patterns were similar at all sites, with main annual peaks of abundance and biomass in autumn-early winter. Chironomid density was positively related to the biomass of benthic macroalgae (P<0.001). The population studied was multivoltine with a probable average of five generations per year, with overlapping cohorts and a predominance of third- and fourth-instar larvae. Estimates of annual production ranged between 72.2 g DW m^–2 yr^–1 at the site with the lowest rate of water renewal in 1991 and 0.1 g DW m^–2 yr^–1 at the site with the highest rate of water renewal in 1992. Mean annual production and the production/biomass ratio for the system was estimated to be 16.8 g DW m^–2 yr^–1 and 12.7, respectively. Possible factors leading to the observed density fluctuations are discussed, as well as possible sources of error in production estimates.     

Earthworm numbers,biomass and respiratory metabolism in a beech woodland-Wytham Woods,Oxford

Summary The mean annual population density of earthworms was found to be 164.6 m^-2 during a period of detailed study between October 1971 and September 1972. In a year of less detailed study between November 1972 and October 1973 the population density was 117.5 m^-2 (139.8 m^-2 when the type of extraction method was allowed for). Mean biomass densities in the two years of investigation were 41.0 g preserved wet wt m^-2 (1971–1972) and 38.6 (possibly 39.2) g preserved wet wt m^-2 (1972–1973).Comparison of the Brogden's Belt population and biomass densities with those reported from other woodlands indicates that soil type is more important than leaf litter type in determining the numerical abundance of earthworms. Population densities are lower in beechwoods on mor soils, mor soils also support significantly fewer species. As with numbers, mean biomass density in beechwoods on mor soils was significantly lower than that occurring in beechwoods on mull soils; the latter, in turn, being lower than those found in mixed deciduous woods on mull soils. Unlike population density biomass density is influenced by both soil and litter type, this is discussed by reference to mean body weights and food quality as reflected by tannin, nitrogen and carbohydrate content.The annual respiratory metabolism of the Brogden's Belt earthworms was calculated to be between 10.7 and 13.41 O₂ m^-2 a^-1, which is equivalent to between 4.1 and 5.1% of the total soil metabolism. A production/biomass ratio of 0.49–0.58 was estimated, as was a net population efficiency of 22%.     

Snail populations,beech litter production,and the role of snails in litter decomposition

Summary The population densities of snails living in beech litter were studied form March 1968 to April 1969. Litter production over one year was measured and the role of snails in litter disappearance assessed.Snails were extracted from litter using a modified Vágvölgyi (1952) flotation method, extraction efficiencies being 84%. The mean annual population density of the twenty-one species of snail recorded on the main sampling site was estimated at 489/m². Carychium tridentatum was the most numerous species, with a mean density of 200/m². Acanthinula aculeata, Punctum pygmaeum and Vitrea contracta also had fairly high mean densities. The mean annual biomass was 699 mg dry wt./m² or 278 mg ash-free dry wt./m². Hygromia striolata and Oxychilus cellarius/alliarius were the most important species in terms of biomass on the main site. Within the limits of accuracy imposed by the sampling regime the population densities of four out of five of the species (C. tridentatum, A. aculeata, V. contracta, Retinella pura) studied remained unchanged throughout the year, whereas P. pygmaeum had a significantly higher autumn population. C. tridentatum populations were highly aggregated at all times of the year, most markedly so in June. Other species were aggregated at certain times of the year only. Samples taken from other sites showed total population densities of snails ranging from 185–1082 snails/m².A total tree litter production of 652 g/m²/annum was recorded of which 584g/m²/annum was of beech material. 72% fell in the October–December period. 58% of the beech litter-fall was leaves, 5.2% bud-scales, 27% fruits and 10% twigs and bark. Summation of appropriate field layer peak standing crops amounted to 23.3 g/m². This was considered as potential litter and was equivalent to 3.4% of the total litter input. The litter standing on the woodland floor in Septermber 1968 was 2,700 g/m², hence, assuming a steady state, litter turnover time was estimated as 4.5 years.It was calculated that the total snail population ingested 0.35–0.43% of the annual litter input, of which 49% was assimilated. The role of the individual species is examined in relation to concepts of key species in ecosystem functioning. The possible role of slugs in decomposition processes is also discussed.     

Life cycle and production of Hydrobia acuta Drap. (Gastropoda: Prosobranchia) in a hypersaline coastal lagoon

The life cycle and annual production of Hydrobia acuta was studied in a hypersaline lagoon (s = 39 in summer), forming a part of solar salt works. Quantitative random samples were taken at regular intervals over a period of 15 months using a corer, and snails collected were counted and measured. Weight and biomass was calculated from a length-weight relationship and from measurements of ash content. H. acuta was a strictly annual species in the study lagoon. Recruitment takes place over a brief period in May and June, after which the breeding population dies. Growth of the new generation was slow during summer, probably due to the unfavourably high salinity. A period of rapid growth took place in autumn coinciding with a drop in salinity caused by rainfall. In winter Hydrobia hibernated by burrowing deeply into the sediment. Growth recommenced in spring when the lagoon was reflooded, but by this time the number of survivors was low.The maximum density of snails was 6 000 m^–2 and maximum biomass 500 mg organic dry wt · m^–2. Annual cohort production was estimated as 786 mg organic dry wt · m^–2 · a^–1. These figures are low compared to other studies on hydrobiid snails, and for production in inland waters, but the value for annual P/B = 4.5 is typical for a univoltine species. The relevance of the results to foraging by wading birds (the main consumers), is discussed.     

Comparative bioenergetics of permanent and temporary pond populations of the freshwater clam,Musculium partumeium (Say)

The population energetics of a temporary and a permanent pond population of Musculium partumeium in Southwest Ohio were studied. In the permanent pond (surface area = 396 m², maximum depth = 0.7 m) the population was bivoltine and iteroparous whereas in the temporary pond (surface area = 1042 m², maximum depth = 0.9 m) the population was usually univoltine and semelparous.Growth and biomass were assessed as total organic carbon and total nitrogen to provide estimates of productivity and seasonal changes in C:N for each generation. Productivity (non-respired assimilation = growth + reproduction; N-R.A. = G + R) was 6939 mgC·m^–2·a^–1 (3858 and 3353 mgC·m^–2·a^–1 for each generation) and 1661 mgC·m^–2·a^–1 for the permanent and temporary pond populations respectively. The average standing crop biomass (B) was 606.8 mgC·m^–2 (357.5 and 249.3 mgC·m^–2 for each generation) and 231.9 mgC·m^–2 with overall productivity: biomass ratios of 11.4 and 7.2 for the permanent pond and temporary pond populations respectively.Respiration rates were converted to carbon equivalents (respired assimilation = R.A.) and used to evaluate the components of total assimilation (T.A. = R.A. + N-R.A.) and the efficiency of partitioning this energy to N-R.A. for G and R. When expressed as a percentage, the production efficiencies (100 × N-R.A.:T.A.) were 50.4 and 62%, and the reproductive efficiencies (100 × R:N-R.A.) were 26.4 and 18% for the permanent and temporary pond populations respectively. The reproductive efficiencies for populations of these viviparous clams are greater than those for most oviparous molluscs.The comparative information on the energetics of these populations does not completely fit any theoretical consideration of reproductive effort or life-history strategy. These data are discussed in relation to selection for population success in temporary ponds.Funded in part by grants to Albert J. Burky from the Ohio Biological Survey and the University of Dayton Research CouncilFunded in part by grants to Albert J. Burky from the Ohio Biological Survey and the University of Dayton Research Council     

Primary productivity and trophic status of Lakes Sorell and Crescent,Tasmania

The primary productivity of two turbid, shallow lakes on the Tasmanian Central Plateau was determined by the C¹⁴ technique from half-light day incubations in situ. Graphical integration of depth-rate curves gave estimates of areal day rates of production and of annual rates.The 2 lakes are closely adjacent and very similar physically and chemically, but have very different phytoplankton populations. Lake Crescent has ten times the standing crop biomass of Lake Sorell but its greater turbidity restricts light penetration, and production per unit of surface per day and per year is only 2.6 times that of Sorell.With day rates of 25-(44)-93 mgCm^–2 and annual production of 16.9 gCm^–2 Lake Sorell could be regarded as oligotrophic. Consideration of standing crop biomass and morphometry however indicates oligo-mesotrophy. Lake Crescent with day rates of 35-(115)-250 mgCm^–2 and annual production of 45 gCm^–2 is moderately eutrophic.Incubations in constant light demonstrated considerable variation in production rates in different parts of Lake Crescent.     

Annual and seasonal changes in fine root biomass of a Quercus ilex L. forest

The biomass, production and mortality of fine roots (roots with diameter <2.5 mm) were studied in a typical Mediterranean holm oak (Quercus ilex L.) forest in NE Spain using the minirhizotron methodology. A total of 1212 roots were monitored between June of 1994 and March of 1997. Mean annual fine root biomass in the holm oak forest of Prades was 71±8 g m^–2 yr^–1. Mean annual production for the period analysed was 260+11 g m^–2 yr^–1. Mortality was similar to production, with a mean value of 253±3 g m^–2 yr^–1. Seasonal fine root biomass presented a cyclic behaviour, with higher values in autumn and winter and lower in spring and summer. Production was highest in winter, and mortality in spring. In summer, production and mortality values were the lowest for the year. Production values in autumn and spring were very similar. The vertical distribution of fine root biomass decreased with increasing depth except for the top 10–20 cm, where values were lower than immediately below. Production and mortality values were similar between 10 and 50 cm depth. In the 0–10 cm and the 50–60 cm depth intervals, both production and mortality were lower.     

Biomass and primary-production of herbaceous plant communities in the Amazon floodplain

The increase in biomass of different aquatic and terrestrial herbaceous plant communities was measured during various growth periods in the Amazon floodplain near Manaus. Maximum biomass varied from 4–11.2 t ha^–1 dry weight in mixed annual terrestrial communities to 6–23 t ha^–1 in aquatic annual species (Paspalum repens, Oryza perennis, Luziola spruceana and Hymenachne amplexicaulis) and 15.6–57.6 t ha^–1 in communities of the perennial species Paspalum fasciculatum. Cumulative biomass of 3 successively growing annual species reached 30 t ha^–1 a^–1. Net primary production is considerably higher than maximum biomass. Paspalum fasciculatum reached 70 t during a growth period of 8 months. If one considers for annual species a monthly loss of 10–25% of the biomass, then net primary production in areas with three successive macrophyte communities and a cumulative maximum biomass of 30 t ha^–1 is estimated to reach up to 50 t ha^–1 a^–1. Annual P/B ratio may reach about 3.     

Enchytraeid numbers,biomass and respiratory metabolism in a beech woodland - Wytham Woods,Oxford

Summary The mean annual population density of enchytraeids in the litter and upper 6 cm of soil was found to be 14,590 m^-2. Mean individual weights approximated 116, 158 and 151 g wet wt. in the litter, 0–3 cm and 3–6 cm strata respectively. The mean biomass was estimated to be 1.908 g wet wt m^-2. Vertical migration was shown to occur, the upward migration in late Autumn was a response to temperatures in the litter being temporarily higher than those of the 0–3 cm stratum. As a result of upward vertical migration and possibly recruitment, numbers reached a maximum in the litter during Winter. On an annual basis the litter, 0–3 and 3–6 cm strata contained 41.43, 46.44 and 12.12% respectively of the extracted enchytraeids. The equivalent biomass values were 33.18, 52.46 and 14.36%. Total numbers and biomass revealed a general picture of high values in late Autumn to Winter which gradually decreased through Spring and Summer except for a minor peak in May–June. The annual oxygen consumption of the enchytraeids approximated 4.285 l O₂ m^-2 yr^-1 (20.461 kcal85.610 kJ) and accounted for 1.63% of the total soil metabolism. A production/biomass (P/B) ratio of 4.93 was estimated as was a net population efficiency of 32%. For this site the contribution by enchytraeids to total soil respiration is about one-third that of the earthworms and an order of magnitude greater than that of the nematodes.     

Production,population biology and diet ofNeomysis integer (Leach) in a shallow Frisian lake (The Netherlands)

TheNeomysis integer population of a shallow Frisian freshwater lake, situated in the north of the Netherlands, was studied from May until October 1980. The results were compared with observations on populations in other Frisian lakes, and with the results reported by other authors on brackish-water and marine populations of this species.Three generations were observed: one overwintering generation and two summer generations. Reproduction stopped completely during winter. During 1980 population densities ofN. integer were relatively low in all Frisian lakes. In Slotermeer the maximum population density was only 6 individuals m^–2, production amounted to 10 mg dry wt m^–2 yr^–1. The annual P/B ratio was 4.0. The diet ofN. integer consisted, in terms of biomass, of more than 95% detritus and animal food, the latter mainly consisting ofBosmina and cyclopoid copepods. Feeding intensity reached a maximum at sunset. It was still high during first part of the night and was low in the morning just after sunrise.     

Production of trout,Salmo trutta,in a Danish stream

Synopsis The numbers of trout,Salmo trutta, in Granslev », Denmark, were estimated by the removal method on 18 dates from March 1974 to March 1976. Populations density varied from 0.39 to 0.74 trout m^–2 in 1974–1975 and from 0.36 to 0.59 m^–2 in 1975–1976 and at all times four or five year classes were present. The age structure of the population was unstable and the variable natural survival, immigration into and emigration from the study site could not be separated. An annual growth cycle with the most rapid growth for all year classes taking place from May to early August was found. Statistically significant differences between different years occurred in the growth of the 0,I and II age groups, but no evidence of density-dependent growth was found. The biomass ranged from 35.4 to 9.5 g m^–2. The total mean annual biomass was 22.8 and 14.7 g m^–2 in the two years and the II group made the greatest contribution, 44 and 48%, respectively. During 1975–1976 the mean annual biomass of each year class only was about two-thirds of that in 1974–1975. Annual production in the two years was 25.7 (range 24.7–28.5) and 12.6 g m^–2 (range 11.7–15.0) and the II group accounted for about 46 and 38% of the production. In addition eel,Anguilla anguilla, produced about 0.5 g m^–2 yr^–2. The unstable age structure of the trout population was compared with trout populations from other streams. The importance of immigration as a recruitment process in middle and lower reaches of streams and of migrations as a mechanism to optimise utilization of the total stream habitat, as well as temperature as a factor controlling the growth rate are discussed.     

Nematode numbers,biomass and respiratory metabolism in a beech woodland—Wytham Woods,Oxford

Summary The mean annual population density of nematodes in the litter and upper 6 cm of soil was found to be 368,000 m^-2. Mean individual live weight biomass approximated 0.2 g and mean biomass was calculated to be 74.6 mg live weight m^-2. No evidence of seasonal vertical migration between the litter, 0–3 cm and 3–6 cm strata was found and on average these strata contained 21.9, 46.2 and 31.9% respectively of the total number of nematodes recovered. The equivalent biomass values were 26.14, 56.57, and 17.29%. Total numbers revealed a general picture of low densities in spring and high ones in early winter, whereas biomass m^-2 was low in late summer — autumn and high in winter. The annual oxygen consumption of the extracted nematodes was calculated to approximate 0.211 m^-2 (4.0 kJ m^-2) but when corrected for the effect of individual biomass (weight specific oxygen uptake) was equivalent to an energy expenditure 6.0 kJ m^-2 which in its turn, because of the efficiencies of extraction, probably accounts for only 87% of the total energy expenditure by the nematode fauna. The nematodes were estimated to be responsible for a minimum of 0.11% to a maximum of 0.13% of the total soil respiration. A production/biomass ratio of 5.16 was estimated as was a net population production efficiency of 36.63%.     

Population studies of three aquatic gastropods in an intermittent backwater

Summary Three snail populations of an intermittent backwater were studied over a 20-month period, and estimates were made of rate of population change, mean biomass, annual and summer net production, and survival under conditions of little standing water. Lymnaea palustris and Physa integra populations were essentially univoltine, while G. parvus appeared to produce several generations during the year. L. palustris formed a protective epiphragm and was apparently better suited to recurrent dry periods than either G. parvus or P. integra, the latter population being maintained through immigration from the stream population. The population density of G. parvus was usually well below, and more aggregated, than the other two species, and there was considerable sampling error associated with its estimated population parameters.The estimated annual production rates for L. palustris and P. integra were 2.18 g/m² and 1.59 g/m², respectively, and these two species accounted for about 98% of the total gastropod production. About 75.9% and 66.5% of the L. palustris and P. integra production, respectively, occurred between June and November. Summer turnover ratios (ratio of production to mean standing crop) in 1969 were 4.69 for L. palustris and 2.94 for P. integra. An extended dry period early in the summer of 1970 reduced total snail production to about 18–24% that of the previous season, although summer turnover ratios were fairly consistent.     

Effect of winter fire on primary productivity and nutrient concentration of a dry tropical savanna

Burning increased the mean annual canopy and belowground biomass of a dry tropical savanna by 40% and 12%, respectively, while littermass was reduced by 85% in comparison to control savanna. Mean annual aboveground and belowground net primary production were 471 and 631 g m^-2 in control, and 584 and 688 g m^-2 in burned savanna, respectively. Fire caused an increase in mean aboveground net production of 24% and in belowground net production of 9%.Concentration of carbon, nitrogen and phosphorus in vegetation of unburned plots ranged between 34.01–38.59%, 0.85–1.53% and 0.04–0.11% and in soil from 0.95–1%, 0.011–0.13% and 0.017–0.02%, respectively. Fire increased the mean concentrations of N and P by 16% and 42% in vegetation and 18.18% and 17.65% in soil, respectively. Thus winter fire can be an important tool for the management of dry tropical savanna with respect to biomass production and nutritive quality.     

Forest floor biomass,litter fall and nutrient return in Central Himalayan oak forests

The seasonal dynamics of forest floor biomass, pattern of litter fall and nutrient return in Central Himalayan oak forests are described. Fresh and partially decomposed litter layers occur throughout the whole year in addition to herbaceous vegetation. The highest leaf litter value is found in April and May and the minimum in September. Partially and largely decomposed litter tended to increase from January to May with a slight decline in June. The wood litter peaked in March and April. The relative contribution of partially decomposed litter to the forest floor remains greatest the year round. The maximum herbaceous vegetation development was found in September with a total annual net production of 104.3 g m^-2yr^-1. The total calculated input of litter was 480.8 g m^-2yr^-1. About 68% of the forest floor was replaced each year with a subsequent turnover time of 1.47 yr. The total annual input of litter ranged from 664 (Quercus floribunda site) –952 g m^-2 (Q. lanuginosa site), of which tree, shrub and herbaceous litter accounted for respectively 72.0–86.3%, 6.4 – 19.4% and 5.2 – 8.6%. The annual nutrient return through litter fall amounted to (kg ha^-1) 178.0 – 291.0 N, 10.0 – 26.9 P, 176.8 – 301.6 Ca, 43.9 – 64.1 K and 3.98 – 6.45 Na. The tree litter showed an annual replacement of 66.0 – 70.0%, for different nutrients the range was 64 and 84%.     

Life history, population dynamics and production of eastern mosquitofish, Gambusia holbrooki (Pisces, Poeciliidae), in rice fields of the lower Mondego River Valley, western Portugal

The introduced population of Gambusia holbrooki from the rice fields of the lower Mondego River Valley, Portugal, was studied for 15 months, relating their life cycle and population dynamics with its production, in order to assess the role of the species in the energy flow and secondary production in this type of agro-ecosystem. Two main annual cohorts (1995 and 1996 cohorts) were identified. The females outnumbered males and the average female/male-ratio was 4. The inspection of ovary developmental stages of this viviparous fish, revealed that the most important reproductive period was between April and August. The first recruits were recorded in June and were present thereafter until October. Males from the parental cohort died before August, whereas parental females could survive until October. Mean adjusted fecundity (number of embryos divided by female standard length) peaked in July 1996 (0.95) and in June 1997 (1.05). Females reached greater sizes, had a higher growth rate and lived longer than males. Annual production was estimated at 3.101 g.m^–2.year^–1 (ash-free dry weight, AFDW), the average biomass at 2.896 g.m^–2 (AFDW), and the P/B ratio was 1.071. A conjugation of life history, population dynamics, production and ecological traits (e.g. fast growth, reduced longevity, viviparity, high productivity, an intermediate position in food chain, and no special habitat requirements for reproduction) clearly show that the populations of G. holbrooki, introduced into rice fields all over the world, may play an important role in the structure and functioning of the biological communities of these important agro-ecosystems.     

Influence of environmental factors on the production of Potamothrix alatus paravanicus Poddubnaya & Pataridze (Tubificidae) in different areas of Lake Sevan

A modification of the Hynes' method was used to estimate production of Potamothrix alatus paravanicus Poddubnaya & Pataridze (Tubificidae), the dominant species of benthic invertebrate in Lake Sevan. Maximum production (5–24 g m^t\-2 wet weight) occurred in the sublittoral, while maximum biomass was found (8–17 g m^–2 wet weight) in the profundal. The turnover ratio of the average cohort varied between 2.4 and 5.0. The annual turnover ratio (P/B) decreased from 2.1–3.1 in the littoral to 0.1–0.4 in the profundal zone.A logarithmic correlation was found between P/B and temperature and oxygen regime, and depth and average size of clitellate specimens.     

Production of Gammarus pulex L. (Amphipoda) in a small Danish stream

Quantitative samples of Gammarus pulex L. taken from a small Danish stream during 1975 showed mean annual population densities varying from 500 m^–2 in early May to 5 500 m^–2 in late September. The mean annual biomass was 1.5 g dry weight m^–2. No discrete cohorts could be distinguished from the size frequency distributions. Annual production, estimated by the size-frequency method, was 3.9 g dry weight M^-2 and P/B ratio was 2.6. The contribution to trout energetics may have been as much as 17%.