The populations of fish in the River Tees system on the Moor House National Nature Reserve, Westmorland

The age, growth, population densities and annual production of an isolated population of brown trout ( Salmo trutta L.) and bullhead ( Coitus gobio L.) were investigated over a 5 year period in Trout Beck in the northern Pennines. Additional data were obtained from two other sites; Great Dodgen Pot Sike and the River Tees.
Trout growth rates were low, particularly in Dodgen Pot Sike, but ages up to VIII were recorded. Bullhead growth rates were similar to those in the Tees at Cow Green (Crisp et al. , 1974), though the Moor House bullheads had a higher survival rate. Population densities of trout ranged from 0.10–0.22/m², with a fry contribution of 0.80%. The bullhead population in Trout Beck was sparse, generally 0.1/m², though an 0 group value of 0.46/m^a was estimated in 1967. At Tees Bridge values ranged from 2.5–7/8/m². Most trout matured by age III, though in Great Dodgen Pot Sike only half of the females were mature at age IV.
Annual production was estimated from Allen graphs. Trout production ranged from 1.02–3.50 g/m²/year, the fry contributing up to 51.5% of the total. A value of 0.48 g/m²/year was obtained for bullheads in the Trout Beck system in a year of good recruitment, whilst bullhead production at Tees Bridge was 7.43 g/m²/year, with fry and I group contributing 48.7 and 37.5% respectively. The high survival rate, irregular recruitment, and poor growth of these high altitude populations is discussed. Growth and reproductive peculiarities of the Great Dodgen Pot Sike trout are also discussed.     

Diel abundance, migration and feeding of fish larvae (Eleotridae) in a floodplain billabong

Eleotrid larvae (2.1–16 mm) were collected from surface waters of a billabong in south-eastern Australia. Estimates of larval density in plankton net samples at night averaged 148.3 larvae per m³ and 16.6 larvae per m³ during the day. In contrast, pump samples provided density estimates of 8.3 larvae per m³ at night and 0.9 larvae per m³ during the day. Larval densities did not differ between open water, snag (fallen tree) and Typha habitats, but Typha habitats yielded larger larvae than other habitats. 32.9% of larvae in pump samples were damaged and unmeasurable, creating a bias favouring larger larvae. The modal length of larvae in net samples at night was 5–6 mm, compared with 3–4 mm during the day, reflecting both greater net avoidance by larger larvae during the daytime and dispersal of smaller larvae from the surface at night. Dispersion patterns of larvae suggest that classes of larvae smaller than, and larger than 5.0 mm exhibit reciprocal diel vertical migration behaviour linked to ontogenetic changes in diet. Larvae less than 5 mm fed only during the day and preyed exclusively on rotifers, whereas larger larvae continued to feed at night and consumed mostly planktonic crustaceans.     

Comparative primary productivity of algal epiphytes on three species of macrophyte in the littoral zone of Lake Ohrid, Yugoslavia

Primary productivity of algal epiphytes on the surfaces of Phragmites, Potamogeton , and Nuphar was measured seasonally from June 1978, through June 1979, in the littoral zone of Lake Ohrid, using ¹⁴C methodology. Surface areas of individual macrophytes were determined throughout the study period through the use of a non-miscible surfactant and a calibration curve of surfactant weight versus known, calculated surface areas.
Mean total surface area available for epiphytic colonization during the study period was 1.032 m² macrophyte surface per m² of littoral zone for Phragmites , 0.810 m² for Potamogeton , and 0.167 m² for Nuphar . Seasonal rates of mean primary productivity of algal epiphytes on Phragmites from the surface to the light-compensation depth ranged from 84–1406 mg C m⁻² littoral zone d⁻¹; ranges for epiphytes on Potamogeton and Nuphar were 77–586 and 69–268 mg C m⁻² littoral zone d⁻¹, respectively. Maximum rates were observed typically during June; minimum rates were observed typically during August to December. Mean daily productivity rates over the 12 month period were for epiphytes on Potamogeton 167.0, on Nuphar 100.4 and on Phragmites 671.2 mg C m⁻² littoral zone d⁻¹. Calculated annual production for epiphytes on Nuphar was 36.65, on Potamogeton 60.95 and on Phragmites 245.0 g C m⁻² littoral zone yr⁻¹. Epiphytic production data were typically considerably higher than production data obtained for littoral and pelagial planktonic algae and compare favorably with published data for epiphytic and periphytic production in Lawrence Lake, Marion Lake, and Borax Lake.     

An estimate of the sustainable rate of shell extraction from the Dutch Wadden Sea

1. Shell production by cockles Cerastoderma edule was studied to examine whether or not the present licensed rate of shell extraction in the Dutch Wadden Sea exceeds the current rate of shell addition to the exploitable stocks.
2. Long-term data on numbers of cockles and weights of their shells were used to estimate their annual production on Balgzand, a 50-km² tidal flat area in the western-most part of the Wadden Sea. During the 1969–97 period, it amounted to an average of 125 g m^–2, including 107 g m^–2 of shells large enough to be exploitable for shell-lime fishery.
3. The very irregular annual recruitment of cockles was the main cause of the wide 95% confidence limits (74 and 140 g m^–2 year^–1) of this 28-year estimate. Moreover, high mortality rates in severe winters substantially reduced production per recruit in some year classes.
4. About one-third of the estimated production does not reach exploitable stocks, because it is fragmented by birds (particularly eider ducks), permanently buried in the sediment, or removed by the fishery for live cockles.
5. During the last few decades, the estimated mean amount added annually to the exploitable stocks was 88 million kg or 132 000 m³ of large cockle shells. This amount compares favourably with the current annual level of removal of 134 000 m³ of shells, three-quarters of which are cockles.
6. Even at temporarily lower production rates, the exploitation of shell stocks at its present rate is not expected to lead to a rapid exhaustion of the existing stocks in the tidal inlets of the Dutch Wadden Sea, as these stocks will be in the order of a few million m³.     

The forest of Vorsø, Denmark: Succession towards a natural, decidous boreal forest influenced by breeding cormorants

The forest on the strictly protected island Vorsø in Horsens Fjord has since 1952 been examined every ten years to discover the number, basal area, wood volume and aboveground biomass of the species. This paper describes the results from 1982 and the development up to that. The dominating species are Fraxinus excelsior, Ulmus glabra and Fagus sylvatica. The two forests, "Vesterskov" (6.09 ha) and "Østerskov" (2.48 ha) respectively, were, in 1982, still immature. Their number of stems per ha were: 756 and 509; basal area: 38 m² and 45 m². volume: 544 m³ and 615 m³; biomass 207 t and 268 t. The annual increase 1972–1982 was respectively 8.8 m³/ha and 0.3 m³/ha. The impact of the cormorant ( Phalacrocorax carbo sinensis ) is especially in "Østerskov" increasing, since the colony here was established in 1976. The succession of the forest is discussed in relation to the disturbance by the cormorants.     

Phytoplankton and pelagic primary productivity in Øvre Heimdalsvatn

A phytoplankton investigation was carried out in the subalpine, low-productive Norwegian lake Øvre Heimdalsvatn in 1969–70 and 1972. This paper describes the temporal and spatial distribution of the standing stock of phytoplankton, and phytoplankton primary productivity. The annual average primary productivity in 1972 was 4.0–4.9 mg C m⁻³ d⁻¹; the annual average standing stock varied from 120 mg m⁻³ (freshweight) in 1969–70, to 250 mg m⁻³ in 1972. Phytoplankton species composition and size distribution is discussed. Throughout the year the phytoplankton is dominated by small (ultraplankton) species; μ-algae (< 5 μm) showed cell concentrations up to 15 mill. cells 1⁻¹. The dominating group was chrysophytes; cryptophytes, dinoflagellates or green algae were at times abundant. A phytoplankton monthly budget and a diagram showing annual average carbon flow through the standing stock of phytoplankton are presented; the phytoplankton dynamics in Øvre Heimdalsvatn is compared to that of other low-productive lakes.     

Effect of air temperature upon mean water temperature in streams in the north Pennines and English Lake District

SUMMARY. Water temperatures from six north Pennine streams (approximate mean annual discharges 0.04–1.5 m³ s^-1) and two Lake District streams (mean annual discharges c. 0.04 m³ s^-1) were used to examine the possibility of predicting 5-day or 7-day mean water temperatures from standard meteorological/hydrological data collected at north Pennine stations. The relationship between mean air temperature (X) and mean water temperature (Y) was approximately linear, except in the Pennine streams for periods when mean air temperature was below 0°C. The calculated regressions of mean water temperature upon mean air temperature accounted for 86–96% of the variance of water temperature even when the air and water temperatures were measured at stations some 50 km apart. A negligible improvement was obtained by the use of multiple regressions incorporating rainfall or stream discharge from a nearby metering weir. Predicted water temperatures were rather lower than observed values during the first half of the calendar year and rather higher during the second half, but these seasonal differences could not be adequately quantified for practical application.     

Effect of fire on the topsoil seed banks of rehabilitated bauxite mine sites in the jarrah forest of Western Australia

Summary Germinable seed stores of 5- and 8-year-old rehabilitated bauxite mine pits in south-west Western Australia were assessed before and after burning. These seed stores were compared to those of adjacent unmined Jarrah ( Eucalyptus marginata ) forest, to identify at what age fire can be reintroduced, in order to measure restoration success and reduce fire hazard. Soils were sampled in early summer (before fire) and late autumn (after fire). Before fire, the mean topsoil seed bank of 5-year-old sites was 2121 seeds per m² while 8-year-old sites had a mean of 1520 seeds per m². Only the 5-year-old sites were significantly different from the forest mean of 1478 seeds per m² for the same season. After summer burns (and possibly due to seasonal effects) topsoil seed banks of rehabilitated areas (sampled in autumn) decreased by an average of 53 per cent. Topsoil seed banks of 5–8-year-old sites were resistant to lower intensity burns, with 362 seeds per m² of native species surviving mild burns and 108 seeds per m² of native species surviving after an intense summer fire. The topsoil seed reserve of 5–8-year-old rehabilitated areas had a high proportion of annual weed species while the forest sites had high levels of subshrubs and native annuals. Low-intensity burns did not alter the composition of life-forms in the soil seed bank, while intense burns favoured annual weed and shrub species. The results indicate that it is not appropriate to introduce fire to rehabilitated areas before 8 years, due to limited fuel reduction benefits and possible adverse effects on obligate seeding species. The large proportion of weed species in the soil seed bank of young rehabilitated areas is a concern, and remains a major consideration for future disturbance of these areas.     

Population dynamics and production of the pelagic amphipod Hyalella montezuma in a thermally constant system

SUMMARY. 1. The population dynamics and annual production of the multivoltine. pelagic amphipod Hyalella montezuma were studied over a 3-year period in the thermally constant environment of Montezuma Well. Arizona.
2. H. montezuma showed two maxima which coincided with spring and autumn phytoplankton maxima. Juveniles comprised over 85% of the population in the pelagic zone compared to 37% in the littoral vegetation and there were significantly more females in the littoral vegetation. It appears that juvenile and adult H. montezuma show different habitat preferences.
3. Total annual mean production for H. montezuma calculated by the size frequency method and adjusted for multiple generations, was 357 kg ha⁻¹ yr⁻¹, which is higher than single-species production estimates reported for most zoobenthic amphipods and multivoltine planktonic crustaceans. Average energy production for H. montezuma was 4640 kJ ha⁻¹ yr⁻¹ in the pelagic zone and 1072 kJ ha⁻¹ yr⁻¹ in the littoral vegetation.
4. Average cohort P / ratios for H. montezuma were higher in the pelagic zone (5.5) than in the litttoral vegetation (3.7). Juveniles had higher cohort P ratios than adults in the pelagic zone, while the reverse relationship was true in the littoral vegetation. We propose that different size-selective predators may contribute to the differences in P ratios for juveniles and adults in these two habitats.     

Influence of water depth on dispersion of juvenile salmonids, Salmo salar L. and S. trutta L., in a Scottish stream

The dispersion of salmon and trout of 0 +, 1 + and 2 + age classes was examined by electrofishing 12 sections of a stream in Perthshire, Scotland at the end of the growing season in September to October in 1972 and 1976. In 1972 sections ranged in surface area from 21 to 122 m² and had volumes of 5.0–17.3 m³ and when surveyed in 1976 surface areas were 15–106 m² and volumes 2.1–6.7 m³. In 1976 widths and areas of sections were 63–87% of the values for 1972, depths were 39–68% and volumes 28–52% of the earlier values. These differences were due to 1976 being drier and warmer. In both years, all sections contained 0 + and 1 + age classes of salmon and trout and some sections contained 2 + age salmon and trout. The total density of fish in a section ranged from 1.9 to 3.9 m⁻² in 1972 and from 3.72 to 6.08 m⁻² in 1976. There was an inverse relationship, significant in 1972 only, between the density of 0 + salmon and that of 1 + trout in the different sections. Densities of both 0 + and 1 + salmon in the sections were inversely correlated, and those of 0 + and 1 + trout were directly correlated, with area of water deeper than 10 cm.     

The cytotoxic and photodynamic inactivation of micro-organisms by Rose Bengal

Rose Bengal was cytotoxic to the following bacteria at the concentrations given in parentheses (highest concentrations of dye in mol/1 at which growth occurred on nutrient medium): Brochothrix thermosphacta and Deinococcus radiodurans (1 times 10^-6 or less); Streptococcus, Micrococcus, Staphylococcus, Bacillus, Arthrobacter and Kurthia spp. (1 times 10^-5–1 x 10^-4), and Pseudomonas spp. and Enterobacteriaceae (5 times 10^-3–1 x 10^-2 or greater). These organisms were killed rapidly when suspended in illuminated (170 μE/m²/s) solutions of Rose Bengal (1 times 10^-4 mol/1) providing oxygen was present. Singlet oxygen was identified as the lethal agent, because the rate of killing was increased by dissolving the dye in deuterium oxide while the organisms were protected against photoinactivation by L-histidine or crocetin. Yeasts from chilled foods were killed in illuminated solutions of Rose Bengal but a light intensity of 315 μE/m²/s was needed for a death rate comparable with that of bacteria. The yeasts present in a range of chilled meat and dairy products failed to form colonies on Rose Bengal (5 times 10^-5 mol/1) media exposed continuously to modest illumination (55–80 μE/m²/s).     

Juvenile salmonid production in a Lake Erie nursery stream

Normandale Creek (2531m²) provides spawning and nursery grounds for lake-run rainbow trout, Salmo gairdneri Richardson, brown trout, Salmo frutta L. , and coho salmon, Oncorhynchus kisutch (Walbaum). Upstream movements are significantly correlated with peak stream discharge (P < 0.05). In 1973–1974, 59 adult salmonids constructed 86 nests of which 60% were disturbed by re-use or sand deposition. From a calculated deposition of 90 400 ova, 483 juveniles, 6 cm in fork length ( f.l. ), were resident in the stream when estimates by electrofishing first became valid. Estimated within-stream (i.e. all sections combined) monthly densities of wild juveniles >6 cm f.l. ranged from six to 22 fish (100 m²)⁻¹, and biomass from 1.3 to 3.1 gm⁻². The highest within-section biomass was 8.3 gm⁻² in November. Additions of hatchery-reared rainbow trout temporarily increased monthly within-stream juvenile density up to 23 fish (100 m²)⁻¹, and biomass to 8.2 g m⁻². Density and biomass were positively correlated (P < 0.05) with both instream and bank cover, and biomass negatively correlated (P < 0.05) with gradient. The importance of substratum and flow characteristics on juvenile density and biomass was strongly indicated although not statistically significant. Annual within-section production of juveniles > 6 cm f.l. ranged from 4.79 to 5.93 g m⁻² year⁻¹ in Sections III and IV, respectively. Inclusion of calculated probable numbers of fish < 6 cm f.l. increased within-section production from 5.93 to 18.65 g m⁻² year^−l in upstream Section IV.     

Adaptations to extreme low light in the fern Trichomanes speciosum

Trichomanes speciosum is a threatened species restricted to sheltered, very humid sites. Uniquely amongst European ferns, differing ecological tolerances of the gametophyte and sporophyte generations are manifested as widely differing distributions. The perennial, vegetatively propagating gametophyte persists in drier, colder, darker habitats than the sporophyte. In sites where the gametophyte grows, light availability was found to be < 1 μmol m² s¹ for approx. 85% of daylight hours, rarely or (in some sites) never rising above 10 μmol m² s¹. Much of the time, light was < 0.01% of full sunlight. Measurements of gas exchange and chlorophyll fluorescence yield show that these plants have optimal photosynthesis at light intensities c . 5–10 μmol m² s¹, the highest light to which they are normally exposed to in their natural environment. The absence of any capacity for reversible nonphotochemical fluorescence quenching means that there is little or no protection of the photosynthetic apparatus from light-induced damage. We conclude that these plants are able to create what are essentially monocultures in their extreme environments only because of a combination of low metabolic rate (at low temperatures) and an ability to make efficient use of what little light is available to them by morphological and physiological means.     

Effects of light and temperature on the cell size and some biochemical components in two freshwater cryptophytes

The effects of light and temperature on cell size and cellular composition (chlorophyll, protein, carbohydrate) of two freshwater cryptophytes were studied with batch cultures. Neither of the species had a constant cell size but the size varied with growth conditions. At each temperature the smallest cells were recorded at the lowest experimental photon flux density. The smallest cells of Cryptomonas 979/67 had an average volume of 232 μm³ and the largest ones 1 020 μm³. In Cryptomonas 979/62 the smallest and largest cells measured 4 306 μm³ and 12 450 μm³. Both species increased their cellular chlorophyll content when PFB dropped below 110–120 μmol m^-2 s^-1. The highest and lowest chlorophyll contents of 979/67 were 7.45 fg μm^-3 and 0.55 fg μm^-2 respectively. For 979/62 the corresponding values were 10.23 fg μm^-3 and 0.93 fg μm^-3. In both species the protein content remained stable at PFDs higher than 110–120 μmol m^-2 S^-1. The highest content of protein measured in 979/67 was 638 fg μm^-3 and the lowest 147 fg μm^-3. For 979/62 these values were 1 036 fg μm^-3 and 148 fg μm^-3 respectively. The carbohydrate results were less clear and no pattern either in response to photon flux density or temperature was obvious. The lowest and highest contents recorded for 979/67 were 62 fg μm^-3 and 409 fg μ^-3 and for 979162, 36 fg μm^-3 and 329 fg μm^-3     

Relative abundance and growth of mussels (Mollusca: Eulamellibranchia) in pools 8,9 and 10 of the Mississippi River

SUMMARY. Twenty-three species of unionid mussels were sampled during the summer of 1975 from a 76-mile reach of the Upper Mississippi River; this represented eight species fewer than were recorded in a 1930–31 survey for the same section of the river. Of the 244 sites sampled in 1975, mussels were found at 116 sites and the ten most productive sites accounted for 40.8% of the 1205 mussels collected. There were as many as thirteen different mussel species taken from one sampling site. The two most abundant species ( Amblema peruviana and Megalonaias gigantea ) constituted 54.5% of the mussels sampled in 1975; these two species made up only 8.3% of the 1930–31 survey. Individuals of the twelve most abundant mussel species were aged, and shell-length growth equations were determined. In addition, shell-free dry weight growth rate and shell weight growth rate was determined for Amblema peruviana . The calculated annual production for A. peruviana (assuming a mean density of 1 per m²) was 482 mg/m² shell-free dry weight, and 14.471 g/m² shell weight.     

Effects of foraging mode and season on the energetics of the Marine Iguana, Amblyrhynchus cristatus

1. Marine Iguanas ( Amblyrhynchus cristatus ) inhabiting the rocky shores of the Galápagos Islands apply two foraging strategies, intertidal and subtidal foraging, in a seasonal climate. Effects of both foraging strategy and seasonality on the daily energy expenditure (DEE) were measured using doubly labelled water.
2. Difference in foraging mode did not result in significant differences in DEE.
3. On Santa Fé the DEE in the warm season was significantly higher than in the cool season (67·8 ± 21·8 kJ kg^–0·8 day^–1 vs 38·0 kJ kg^–0·8 day^–1). This difference can be explained by body temperature. A model estimate of the body temperature was used to predict monthly DEE figures, giving a year round budget. On average a 1-kg iguana would need only 47 kJ day^–1, or 17 mJ year ^–1. This is lower than previous estimates in which body temperatures were not taken into account.
4. The water flux of the Marine Iguana increases with increasing foraging time. The linear rise per minute foraging is roughly two times as high for subtidally foraging animals as for intertidal foragers.     

Influence of habitat characteristics on small mammals in a Mexican high-altitude grassland

Small mammals in a high-altitude grassland area close to Mexico City were studied. Populations of 10 species were censused using live traps in 48 sample quadrats. Within each quadrat, vegetation characterization, including complete floristic listings, cover values for species and layers and values of habitat modification, were assessed. Habitats were described according to plant communities identified using ordination and classification methods. Nine different plant communities were obtained. Densities and abundance of all small mammal species were calculated for each of the habitats classified. Peromyscus alsloni was the most abundant species in all habitats, reaching maximum densities of 55 ha⁻¹ in pine forest with dense ground and herb layer. Peromyscus melanotis also occurred in all habitats but at lower densities (maximum 29 ha⁻¹). Reithrodontomys megalotis was found in all habitats except in tall dense grassland. Densities for this species were generally low (1-9 ha⁻¹) but reached 19 ha⁻¹ in short dense grassland. All other species were largely absent from 4–8 habitats and showed very low densities (0.75–4 ha⁻¹). The densities of the more abundant species were largely correlated with more open habitats and higher indices of habitat modification. Lower altitude grassland habitats have a greater abundance of small mammals and a higher species richness than the medium and higher altitude, physiognomically more complex habitats. Species richness was highest in tall pine-alder forest with a species-rich, dense herb layer and lowest in pine forest with dense ground and herb layers. Species richness was positively correlated with overall small mammal density.     

Floristic composition, structure, and dynamics of an upper montane rain forest in Southern Ecuador

Two 1 ha plots of undisturbed upper montane rain forest in southern Ecuador were sampled for all trees with a dbh ≥ 5 cm. An extraordinarily high α-diversity for a forest near treeline is described. The "non-ridge forest" plot at 2900 m elevation has 75 species and 28 families ha^-1 and is believed to represent an advanced stage of succession whereas, the "ridge forest" plot at 2700 m elevation with 90 species and 38 families ha^-1 has a more rapid turn-over rate probably due to a more unstable environment. Downslope forces (soil creep) are discussed as a possible cause for 32–44% of all trees being inclined more than 30° in the moderately steep terrain (average slope 20° in both study plots). The plot of "non-ridge forest" is characterized by a much greater biomass whether expressed as basal area (44 m² versus 15 m²) or stem volume (214 m³ versus 52 m³), while the density is equally high (2310 versus 2090 trees ha^-1) in both plots. Families with Family Importance Value >25 / ha^-1 are Clusiaceae, Cunoniaceae, Melastomataceae, Myrsinaceae, and Ternstroemiaceae.     

Modelling black fly production dynamics in blackwater streams

SUMMARY. 1. Two predictive models were employed along with intensive field sampling to estimate production of black flies ( Simulium spp.) on snags (submerged wood) in three blackwater streams on the Georgia Coastal Plain of the southeastern U.S.A. One model predicts daily growth rate from temperature and hydrograph pattern; the other predicts habitat abundance (of snags) from river height.
2. In the sixth order Ogeechee River, annual production was twice as high in 1982 (7.1 g dry mass [=DM] m⁻² of snag surface) as in 1983 (3.6 g DM m⁻²). When converted to production per m² of river bottom, values were 35–40% of the snag surface estimates. Annual production was much lower in fourth order Black Creek (1982, 1.3 g DM m⁻² of snag surface) and much higher in the sixth order Satilla River (1975, 15.6–40.0 g DM m⁻²).
3. There was a distinct bimodal pattern of black fly production in the Ogeechee River in both years, with peaks occurring in winter and summer. Similar bimodal patterns of production were found in Black Creek and in the Satilla River. Although there appears to be an intrinsic component to the bimodal pattern, production peaks (growth rate and biomass) appear to be associated with initial stages of flooding.
4. Annual production/biomass ratios (37–85) are the highest reported for black fly populations. The variation of annual P/B ratios among sites was more strongly dependent on the temporal distribution of standing stock biomass than on differences in growth rates. Variation in production among sites appears to be due to differences in current velocity, hydro-graph variability, and abundance of coexisting consumers.     

Bird community energetics in a boreal coniferous forest

The annual minimum energy consumption of the bird community was 2524 × 10³ kcal km⁻², of which 44% was consumed by wintering species and 73% by passerines. The daily energy consumption was in summer 14–16 × 10³ and in winter 1–2 × 10³ kcal km⁻². In spruce forests and in afforested swamps birds required approximately 0.12% of the net primary production. Their total annual energy consumption was covered by invertebrates (59%), vertebrates (2%) and vegetable matter (39%); the food derived from the ground (55%), from trees (44%) and from the air (1%). Arboreal insectivorous passerines, ground passerines and gallinaceous birds were the most important ecological guilds. Among passerines existence metabolism accounted for 73% of the annual energy consumption, extra activity for 17%, breeding activity for 1%, moult for 4% and nestlings for 4%.