Primary production studies in two linked but contrasting Welsh lakes

SUMMARY. Llyn Padarn and Llyn Peris have distinct phytoplankton populations. During 1975–76, the standing crop measured as chlorophyll- a was 5.5 times greater in Padarn than in Peris and the production rate, determined by the ¹⁴CO₂ method, was faster by 3.4 times. These differences were attributed to the higher concentrations of phosphorus in the lower lake caused by treated sewage effluent. Incident light intensity, which was slightly lower in Peris due to mountain shading, and temperature, which was 1–4°C higher in Padarn, made little significant contribution to these differences during the summer. The reduced transparency of Padarn water, compared with that of Peris, resulted from denser phytopiankton crops in Padarn. During the summer, Padarn exhibited carbon dioxide depletion which correlated with the chlorophyll concentration. Light inhibition at the surfaces of both lakes correlated with solar radiation intensity. However, the relationship between pigment content and maximum photosynthetic rate was poor. Extracellular products accounted for about 16% of the total production in the lakes. Uptake of ¹⁴C-labelled acetate was low compared with that of ¹⁴CO₂ uptake.     

Diatom and chrysophycean cyst profiles in sediment cores from two linked but contrasting Welsh lakes

Llyn Padarn and Llyn Peris are linked by a small river, and although originally a single larger lake, the lakes now support contrasting algal communities. A study of the sequences of diatom remains and chrysophycean cysts present in long sediment cores has been carried out to assess the historical extent of the differences in these algae between the two lakes.

Investigations have included statistical analysis to determine the precision of counting sediment samples. The most efficient sampling and counting procedure, resulting in counts of acceptable precision, involved taking a single sample of sediment from a depth horizon, preparing two replicate slides and counting the entire area of each slide. A comparison of the variation in distribution of these siliceous algae in replicate sediment samples taken from the surface sediment with the variation in distribution down the long sediment core from Llyn Padarn showed that the vertical variation greatly exceeded the horizontal variation. Thus variation in diatom numbers down the sediment core reflects changes in the alga reaching the sediments rather than horizontal patchiness of distribution in the top most layers of sediment.

Studies of the sequences of diatoms and chrysophycean cysts were carried out on a 4·75 m core from Llyn Padarn and 2·1 m core from Llyn Peris representing c. 6000 years and c. 900 years of the two lakes respective history. From c. 6000 until c. 2200 years B.P., Llyn Padarn was an acid oligotrophic lake. It appears that about 2200 years B.P., Llyn Padarn became more enriched, Asterionella formosa was the most numerous diatom preserved in the sediment at this time. Reversion to oligotrophic conditions then occurred c. 2000 years B.P. The oldest material from the Peris sediment core was characterized by centric diatoms similar to those found in Padarn sediment dating from 2000 years B.P. to the surface of the Padarn core. Thus at this time, c. 900 years B.P., the algal communities in the two lakes were similar. Both lakes were oligotrophic. However, by 200 years B.P. the chemistry of the sediment core from Peris showed evidence of extensive copper and slate mining in the environment. At this time the species of siliceous algae preserved in the Peris core changed, and the current differences in algal communities in the two lakes appear to originate from this point in history.     

Algal bioassay of the water from two linked but contrasting Welsh lakes

SUMMARY. A comparison of the potential water fertility of two linked lakes, Llyn Peris and Llyn Padarn, was carried out over a year using Asterionella formosa as the test organism in batch cultures. In unenriched lake water, growth was more rapid in Padarn water during autumn-winter-spring, but there was no significant difference between growth in water from the two lakes during the period of thermal stratification. Mean growth rates in treatments with additions of nitrogen and chelated iron were not significantly different from unenriched water samples, both between treatments and lakes during thermal stratification. Growth in water from both lakes was increased greatly by addition of phosphate. Maximum growth rates occurred on addition of nitrogen, phosphate and chelated iron, and there was no significant difference between growth rates in this treatment for the two lakes throughout the year. Comparisons of mean growth rates in bioassays by Tukey's interval estimate showed significant differences between other treatments and the two lakes. The potential fertility of water samples from the two lakes differs more during mixed water conditions than in the period of thermal stratification. Inhibition of A. formosa growth was recorded four times in unenriched filtered water from Peris, once on addition of nitrogen and once on addition of iron. Inhibition was alleviated by the addition of nutrients. The addition of diluted sewage effluent which enters the drainage system between the two lakes, resulted in similar rates of growth of A. formosa in water from both lakes. Results of the bioassays are discussed in relation to differences in phytoplankton between the two lakes. Although A. formosa will grow in samples of unenriched lake water from Peris, other features, including a shorter retention time of water, lower level of incident radiation and, occasionally, inhibitory substances in the lake water, contribute to the rare occurrence of this diatom in Peris. Padarn is enriched by sewage effluent which aids the growth of A. formosa.     

Effect of algal deposition on acetate and methane concentrations in the profundal sediment of a deep lake (Lake Constance)

Abstract In the profundal sediment ot Lake Constance (143 m depth) the temperature is constant at 4 °C. Despite the constant temperature, CH₄ concentrations changed with season between about 120 μM in winter and about 750 μM in summer, measured down to 30 cm depth. The acetate concentration profiles also varied between seasons. In summer, acetate concentration reached a maximum at about 100 μM in 2 or 4 cm depth. In winter, maximal concentrations of about 5 μM were observed over the entire depth. Input of organic material in late spring may be the reason for the seasonal change of both compounds. To simulate such a sedimentation event, intact sediment cores were covered with suspensions of Porphyridium aerugenium or Synechococcus sp. The addition of the phytoplankton material resulted in a drastic increase of acetate concentrations with a maximum at 2 cm depth, similar to in situ acetate concentrations measured in summer. The same applies for CH₄ for which increased concentrations were observed down to 6 cm depth. H₂ concentrations, on the other hand, showed no distinct increase. Treatment of intact sediment cores with ¹⁴C-labeled Synechococcus cells resulted in the formation of ¹⁴C-acetate, ¹⁴CH₄ and ¹⁴CO₂. Maximum concentrations of ¹⁴CH₄ were found at 4 cm depth, i.e. just above the depth to which ¹⁴C-acetate penetrated. The results show that phytoplankton blooms may cause a seasonal variation of acetate and CH₄ in profundal sediments of deep lakes despite the constant low temperature. They also indicate that acetate is the dominant substrate for methanogenic bacteria in the profundal sediments of Lake Constance.     

Genetics of whitefish and vendace in England and Wales

Proportion of loci polymorphic ( P ) and mean heterozygosity per locus ( H _exp) were P = 0.130 and H _exp= 0.096 for C. albula from the English Lake District and P = 0.217 and H _exp= 0.046 for C. lavaretus from the English Lake District and Llyn Tegid, Wales in 1991–2. Significant deviation from the Hardy-Weinberg model was observed at the superoxide dismutase ( sSOD* ) locus in C. albula from Bassenthwaite Lake. This could have been due to chance, to the Wahlund effect or to selection. Genetic distances between the two species were consistent with their taxonomy and genetic differentiation was much greater for the C. lavaretus populations than for the two geographically close C. albula stocks. Llyn Tegid C. lavaretus showed unique alleles at two loci and conservation of fish in this lake is therefore of great importance. Red Tarn C. lavaretus differed most from other Lake District populations probably due to founder effects from early stocking events and/or from selection due to the high altitude of this lake.     

DIATOM PALEOLIMNOLOGY OF TWO FLUVIAL LAKES IN THE ST. LAWRENCE RIVER: A RECONSTRUCTION OF ENVIRONMENTAL CHANGES DURING THE LAST CENTURY

Water quality degradation is a serious concern for the St. Lawrence River. While some environmental data are available for the St. Lawrence ecosystem, long-term monitoring data are generally lacking. To infer past environmental changes, we undertook a paleolimnological assessment of diatom assemblages preserved in four ²¹⁰Pb- and ¹³⁷Cs-dated sediment cores from two fluvial lakes in the river, and used diatom transfer functions to infer past shoreline habitat characteristics. At sites in Lake Saint-François, a fluvial lake downstream from Cornwall, water quality decreased this century in response to human impacts (e.g. macrophyte density and nutrient levels increased). These trends were apparent from an increase in epiphytic diatom taxa, followed by an increase in eutrophic planktonic taxa. Water quality, however, appears to have improved somewhat in response to rehabilitation measures during the last two decades. From a sediment core near Montréal (Lake Saint-Louis), we also noted a large proportion of eutrophic and epiphytic taxa, but less evidence was recorded of a recent improvement in water quality. The diatom-based inference model for habitat characteristics appeared to reconstruct environmental conditions in the St. Lawrence River during the last century. The most notable shift has been an increase in diatom taxa commonly associated with macrophyte substrates. Trends in some of the planktonic diatoms were similar to those recorded in paleolimnological investigations from Lake Ontario, but cores from the river also may be reflecting local environments. This study shows that diatom-based paleolimnological studies are possible in large river systems, if coring sites (e.g. fluvial lakes) are carefully selected.     

Chernobyl-derived radiocaesium in fish as dependent on water quality and lake morphometry

¹³⁷Cs concentrations in perch Perca fluviatilis , pike Esox lucius and roach Rutilus rutilus obtained from lakes of different size and water quality in an area which received about 10–67 kBq m⁻²¹³⁷Cs, were compared with environmental data. Radiocaesium concentrations were highest in pike, and were about two to three times higher in the pike and perch than in the roach. The largest perch had about four times more ¹³⁷Cs than the smallest ones, but the activities in the pike and roach were independent of fish size. All of the water quality parameters examined correlated with ¹³⁷Cs concentrations in the fish, but the concentrations of ¹³⁷Cs in the bottom sediment did not. ^l37Cs fallout and water retention time in lakes showed a positive correlation with fish ¹³⁷Cs, while the size of the catchment area, phosphorus content of the water, water colour, pH and electrical conductivity showed a negative one. Thus there was more ¹³⁷Cs in the fish from the oligotrophic lakes than from the eutrophic ones, from the less humic lakes than from the more humic ones, from the more acid than from the less acid ones and from the lakes containing less potassium than from those containing more potassium. The depth, area and volume of the lake did not affect radiocaesium in the fish. Regression equations are presented for predicting ¹³⁷Cs in fish on the basis of environmental data. Fallout, total phosphorus and pH together had the highest influence in multiple regression models but were less significant in the roach than in the perch and pike.     

Holocene carbon burial by lakes in SW Greenland

The role of the Arctic in future global change processes is predicted to be important because of the large carbon (C) stocks contained in frozen soils and peatlands. Lakes are an important component of arctic landscapes although their role in storing C is not well prescribed. The area around Kangerlussuaq, SW Greenland (66–68°N, 49–54°W) has extremely high lake density, with ∼20 000 lakes that cover about 14% of the land area. C accumulation rates and standing stock (kg C m⁻²), representing late- to mid-Holocene C burial, were calculated from AMS ¹⁴C-dated sediment cores from 11 lakes. Lake ages range from ∼10 000 cal yr  bp to ∼5400 cal yr  bp , and reflect the withdrawal of the ice sheet from west to east. Total standing stock of C accumulated in the studied lakes for the last ∼8000 years ranged from 28 to 71 kg C m⁻², (mean: ∼42 kg C m⁻²). These standing stock determinations yield organic C accumulation rates of 3.5–11.5 g C m⁻² yr⁻¹ (mean: ∼6 g C m⁻² yr⁻¹) for the last 4500 years. Mean C accumulation rates are not different for the periods 8–4.5 and 4.5–0 ka, despite cooling trends associated with the neoglacial period after 4.5 ka. We used the mean C standing stock to estimate the total C pool in small lakes (<100 ha) of the Kangerlussuaq region to be ∼4.9 × 10¹³ g C. This C stock is about half of that estimated for the soil pool in this region (but in 5% of the land area) and indicates the importance of incorporating lakes into models of regional C balance at high latitudes.     

Microbial sulfate reduction in acidic (pH 3) strip-mine lakes

Abstract ³⁵SO₄ reduction was detected in slurries of sediments obtained from Reservoir 29 (pH 3.8) and Lake B (pH 6.2), two acid strip-mine lakes in Indiana. The rates varied seasonally and were higher in summer and fall than in the spring. The optimal pH for sulfate reduction in Reservoir 29 sediments was 5, but samples had increased activity at pH 7 within 24 h after adjusting the pH to this value. In Lake B, the optimal pH for sulfate reduction was the in situ pH (6.2). Sulfate reduction in both lakes was stimulated 2–3-fold by increasing p _H2. High concentrations (5 mM) of organic acids inhibited sulfate reduction at pH 3.8, but stimulation was observed at concentrations of 0.1 mM. Acid-volatile sulfides accounted for about 70% of the products of ³⁵SO₄ reduction.     

Chironomid stratigraphy in the shallow and eutrophic Lake Søbygaard, Denmark: chironomid–macrophyte co-occurrence

1. A sediment core from the shallow, hypertrophic Lake Søbygaard (mean depth ∼1 m; [TP] 310 μg P L⁻¹) was analysed for subfossil remains to reconstruct chironomid community changes in relation to the succession and disappearance of aquatic macrophytes. 2. Species composition in the 1.10 m core indicates a succession from a 'naturally' eutrophic state to a hypertrophic state during recent centuries. Radiometric dating (²¹⁰Pb) of the uppermost 20 cm of the sediment core (∼1932–93) indicates that sediment accumulation rate had doubled in recent decades. 3. Changes in chironomid assemblages were in close agreement with changes in both diatoms and macrophyte remains in the same core. Distinct changes in chironomid communities reflect the eutrophication process and macrophyte succession through Chara , Ceratophyllum and Potamogeton dominance to the present state, with complete loss of submerged vegetation and dominance by phytoplankton. 4. The co-occurrence and relationship between aquatic macrophyte diversity and recent subfossil chironomid assemblages were assessed from an additional 25 Danish lakes. There was good agreement between the macrophyte and chironomid-based lake groupings. Overall, a significant difference ( P <0.001) was found in chironomid assemblages among lakes in different macrophyte classes. In a pair-wise comparison, the poorly buffered mesotrophic lakes and the alkaline eutrophic lakes had significantly different chironomid assemblages. 5. Chironomid taxa commonly reported to be associated with macrophytes ( Cricotopus , Endochironomus and Glyptotendipes ) were shown also to be indicators of highly productive lakes lacking abundant submerged vegetation.     

Sediment processing by gizzard shad, Dorosoma cepedianum (Lesueur), in Acton Lake, Ohio, U.S.A.

Gizzard shad are primarily detritivorous in Acton Lake, a 253-ha impoundment in southwestern Ohio, U.S.A. To determine the magnitude of sediment utilization by the gizzard shad population in Acton Lake. I used data on population density and age structure, daily ration, and feeding selectivity in estimating the mass of sediments processed by shad daily from April through November. At densities of 4595–10 645 fish ha⁻¹(wet weight biomass = 90–121 kgha ¹), gizzard shad could process 3.8–23.0 kg of dry sediments ha⁻¹ day ¹. On average throughout the growing season, gizzard shad could process a dry mass of sediments each day equivalent to 13% of shad wet weight biomass. Because of the high rate of sedimentation (> 700 kg dry sediment ha⁻¹ day⁻¹) in Acton Lake, gizzard shad can process < 4% of the freshly deposited sediments each day, and therefore are likely to have little effect on benthic community dynamics in the system.     

Sulphur stable isotopes can distinguish trophic dependence on sediments and plankton in boreal lakes

1. Stable isotopes of carbon are useful for differentiating between freshwater food chains based on planktonic algae or benthic algae, but are reported to be of limited use for identifying food chains based on sedimentary detritus. Because data from marine systems suggest that stable isotopes of sulphur (δ³⁴S values) have potential in this regard, we tested their utility in freshwater lakes.
2. We found that sulphate in the water column of four boreal lakes was enriched in ³⁴S compared to the sulphur in bulk sediments from these lakes. Furthermore, within a given lake, insects known to feed on sediment (directly or via predation) had δ³⁴S values similar to those of sediment, whereas planktonic and benthic invertebrates known to feed on suspended particles had δ³⁴S values similar to those of sulphate in the water column.
3. Using the stable S isotope values of invertebrates that obtain their S from either the sediment or the water column as end members in a two-source mixing model, we show that two fish species obtain their food from both planktonic and sedimentary sources. Furthermore, model results suggest that, as expected, the more benthic-feeding fish species obtains more of its S from the sediment compartment than does the species that feeds in the water-column.
4. Our results suggest that measurements of stable sulphur isotopes provide a means of distinguishing between members of food chains that are based in the water column from those based on sedimentary detritus. As such, they would be a useful complement to stable C isotopes that are used to distinguish between food chains based on planktonic or benthic algae.     

Relationships between DOC concentration and epilithon stable isotopes in boreal lakes

1. Littoral biota in boreal lakes are known to assimilate epilithon. Being able to predict the stable isotopic composition of these alga will help to identify those systems in which δ¹³C and δ¹⁵N analysis can be used in foodweb investigations of allochthony and biomagnification.
2. In a survey of 15 boreal lakes, the concentration of dissolved organic carbon (DOC) explained 76% of the variation in epilithon δ¹³C, and 86% of the variation in epilithon δ¹⁵N.
3. Because both δ¹³C and δ¹⁵N values were depressed and similar to terrestrial values in humic (high DOC) lakes, it will be more difficult to successfully employ stable isotopic techniques for estimating allochthony in such systems. Lower δ¹⁵N values in humic lakes also indicate that trophic positions estimated by stable isotopes are not directly comparable to those of similar biota inhabiting clearwater lakes, unless autochthonous baseline corrections are made.     

Ammonium and phosphate regenerationby the zooplankton of an Amazon floodplain lake

SUMMARY. 1. Regeneration of ammonium and phosphate by macro-zooplankton (Cladocera. adult copepods. and copepodites) was measured in Lake Calado. an Amazon floodplain lake, Macrozooplanktonabundances ranged between 1×10⁴ and 3×10⁵ individuals m⁻².
2. Phosphate regeneration ranged from 0.2 to 1.3 μ mol PO₄ m⁻² b⁻¹at station 1. located 2 km from the Solimoes River, and from 1.6 to8.3 μ mol PO₄ m⁻² h ⁻¹ at station 3, located 7 km from the SolimoesRiver. Ammonium regeneration at stations 1 and 3 ranged from 1.7 to11.9 and from 13.4 to 77.2 μ mol NH₄ m⁻² h⁻¹. respectively.
3. Zooplankton regenerated ammonium and phosphate at similarrates during rising and falling waier. Regeneration by macrozooplankton was low compared to other tropical lakes and compared to microbesand microzooplankton in Lake Calado.     

Inorganic carbon limitation of photosynthesis in lake phytoplankton

1. Inorganic carbon availability influences species composition of phytoplankton in acidic and highly alkaline lakes, whereas the overall influence on community photosynthesis and growth is subject to debate.
2. The influence of total dissolved inorganic carbon (DIC) and free CO₂ on community photosynthesis was studied in six Danish lakes during the summer of 1995. The lakes were selected to ensure a wide range of chlorophyll a concentrations (1–120 μg l^–1), pH (5.6–9.6) and DIC concentration (0.02–2.5 m m ). Photosynthesis experiments were performed using the ¹⁴C technique in CO₂-manipulated water samples, either by changing the pH or by adding/removing CO₂.
3. Lake waters were naturally CO₂ supersaturated during most of the experimental period and inorganic carbon limitation of photosynthetic rates did not occur under ambient conditions. However, photosynthesis by phytoplankton in lakes with low and intermediate DIC concentrations was seriously restricted when CO₂ concentrations declined. Similarly, photosynthesis was limited by low CO₂ concentrations during phytoplankton blooms in the hardwater alkaline lakes.     

The population biology of pike, Esox lucius L., in two gravel pit lakes, with special reference to early life history

The population biology of pike in two gravel pit lakes of contrasting habitat type were studied. The density of pike (1 + and over) in November 1987 in the Main Lake was 7.7 fish ha⁻¹ (19 kg ha⁻¹) and in St Peter's was 71 fish ha'(53 kg ha⁻¹). The age distribution showed that recruitment was consistent in St Peter's but variable in the Main Lake. The main factor causing this difference in recruitment is abundance of aquatic vegetation. Survival up to November 1987 of 0-group pike in the Main Lake was 0.00175% from potential egg deposition, producing 6.6 fish ha⁻¹ (0.83 kg ha⁻¹). In St Peter's survival was 0.00315%, producing 23 fish ha⁻¹ (3.04 kg ha ¹).     

Stable isotope variability of meso-zooplankton along a gradient of dissolved organic carbon

1.  The δ¹³C and δ¹⁵N signatures of zooplankton vary with dissolved organic carbon (DOC), but inconsistent and limited taxonomic resolution of previous studies have masked differences that may exist among orders, genera or species and are attributable to dietary and/or habitat differences. Here we investigate differences among the isotopic signatures of five zooplankton taxa ( Daphnia , Holopedium , large Calanoida, small Calanoida and Cyclopoida) in Precambrian shield lakes with a sixfold range of DOC concentration.
2.  δ¹³C signatures of Daphnia , small calanoids and large calanoids became more depleted with increasing lake DOC, whereas Holopedium and cyclopoid δ¹³C became enriched with increasing DOC concentration.
3.  The variability of δ¹³C and δ¹⁵N isotopic signatures among zooplankton groups was reduced in high-DOC, compared to low-DOC lakes, especially for δ¹³C. Differences in δ¹³C and POM-corrected δ¹⁵N accounted for up to 33.7% and 19.5% of the variance, respectively, among lakes of varying DOC concentration.
4.  The narrow range of signatures found in higher DOC lakes suggests that different taxa have similar food sources and/or habitats. In contrast, the wide range of signatures in low-DOC lakes suggests that different taxa are exploiting different food sources and/or habitats. Together with the variable trends in zooplankton isotopic signatures along our DOC gradient, these results suggest that food web dynamics within the zooplankton community of temperate lakes will change as climate and lake DOC concentrations change.     

Temporal and geographical variation in lake trophic status in the English Lake District: evidence from (sub)fossil diatoms and aquatic macrophytes

1. A sediment core (representing 250–300 years) was taken from each of three lakes of conservation interest and contrasting trophic status in the English Lake District: Wastwater, Bassenthwaite Lake and Esthwaite Water. Lithostratigraphic analyses, radiometric dating and analysis of fossil diatoms were carried out.
2. Transfer functions, based on the diatoms, were used to reconstruct total phosphorus (TP) and, thus, eutrophication at the study lakes. In Wastwater, changes in lake pH were also reconstructed.
3. The lakes were also classified according to their present macrophyte flora, the latter being compared with previous records.
4. The fossil diatoms of Wastwater were continuously dominated by taxa typical of oligotrophic, circumneutral waters, indicating that the lake has not been enriched or acidified in the last 250 years. The aquatic macrophyte flora has probably remained unchanged since before the Industrial Revolution.
5. The diatom assemblages of both Bassenthwaite Lake and Esthwaite Water began to change in the mid-1800s. Further change occurred from the 1960s, at the onset of a recent period of eutrophication. These two lakes have experienced continued nutrient enrichment throughout the 1970s, 80s and 90s, largely associated with increasing phosphorus inputs from sewage effluent. There is no evidence of any recovery in response to recent reductions in external nutrient loads.
6. Only in Esthwaite Water has the change in aquatic macrophytes been pronounced.
7. Palaeolimnological reconstruction is useful in determining background conditions and natural variation in lake ecosystems.     

Abundance and distribution of anaerobic protozoa and their contribution to methane production in Lake Cadagno (Switzerland)

Abstract In the uupermost layers of the anoxic sediment in Lake Cadagno, 9 different species of anaerobic protozoa were identified. The total number of these organisms was about 580 cells·ml⁻¹ sediment. Most pf these protozoa contained endosymbiotic methanogenic bacteria which in total amounted to 10⁶ methanogens·ml⁻¹ sediment. In addition to the methanogenic endosymbionts, cells of Metopus setosus and Caenomorpha lata also contained a non-fluorescent bacterial rod inside the cytoplasm. In some individual cells of C. lata this second type of endosymbiotic bacterium was sometimes the only endosymbiont observed. Contrary to earlier suggestions, anaerobic protozoa do not seem to play a major role in methane production at least in Lake Cadagno. No significant methane production due to the anaerobic protozoa and their methanogenic endosymbionts was found in situ. Isolated ciliates and amoebae produced methane at 12°C, but not at 6°C, probably as a result of temperature limitation. In the sediment of Lake Cadagno sulfate reduction seemed to be the dominant terminal degradation process.     

Stable isotope ecology of fossil hippopotamids from the Lake Turkana Basin of East Africa

The diet of African hippopotamids can be documented through δ ¹³C analyses of enamel and other tissues. Analysis of a 10-million-year sequence of hippopotamids in and near the Lake Turkana Basin of northern Kenya shows that hippos have included a substantial fraction of C₃ vegetation in their diets since the late Miocene when C₄ vegetation first appears in hippo diet as a measurable fraction. The C₄ component of vegetation becomes dominant (>50%) by Upper Burgi time ( c . 2 million years ago) but does not reach 100% for all individuals. It is therefore not unexpected that the δ ¹³C values of modern hippopotamids show a higher fraction of dietary C₃ biomass than has been estimated from traditional observations. Analysis of δ ¹⁸O of hippos from different stratigraphic levels shows no systematic trend over time; the average value for fossil hippos over the last 10 million years is similar to that of modern hippos from the Omo River system.