Massive fish kills within the Nyanza Gulf of Lake Victoria,Kenya

The morphometry and hydrology of the Nyanza Gulf of Lake Victoria can be greatly affected by violent storms. This can result in a condition in which nutrient-rich bottom mud is mixed with the sediment-laden runoff water from neighbouring marshes and rivers. This situation can lead to massive fish kills. It is such an event in 1984 which will be described that caused the sudden death of over 400 000 fish weighing over 2400 tonnes.The unusually low precipitation in the catchment area during that year resulted in a drop in the lake level from 12.4 to 11.8 m. This low water level, combined with a severe storm created the conditions which caused this large fish kill. The species affected were Lates niloticus and Oreochromis niloticus. Mortality was attributed to several factors which included high levels of suspended material in the water column (detritus and algae) which clogged the gills of the fish, low dissolved oxygen, low pH, and high concentrations of algae.     

The Shrimp Caridina nilotica in Lake Victoria (East Africa), Before and After the Nile Perch Increase

The shrimp Caridina nilotica is a major prey of the introduced Nile perch in Lake Victoria. In spite of heavy predation, the density of shrimps increased after the Nile perch boom and the concomitant disappearance of the haplochromine cichlids. In the same period, the mean size of gravid shrimps and the size at first maturity declined. This seems to indicate an increased predation pressure on adult shrimps. Before the Nile perch upsurge, specialised shrimp eaters and piscivores, among the haplochromine cichlids, only took adult shrimps, whereas we assume that most haplochromines used to include juvenile shrimps into their diet. Another important predator on adult shrimps was Bagrus docmak. The combined density of predators on adult shrimps in the pre-Nile perch era was estimated at 10 kg ha⁻¹ and the potential predators on juveniles were estimated at 170 kg ha⁻¹. After the Nile perch upsurge, only Nile perch up to 10 cm TL and Rastrineobola argentea fed on juvenile shrimps (ca. 36 kg ha⁻¹) and Nile perch from 10 to 50 cm TL (ca. 13 kg ha⁻¹) fed on adults. These rough estimates of the biomass of predators on shrimps before and after the Nile perch upsurge indicate a reduced predation pressure on juvenile shrimps. The disappearance of the haplochromines may have released competition with small Nile perch for juvenile shrimps, thus enhancing the recruitment of Nile perch.     

Potential fishery of Nile perch Lates niloticus Linne (Pisces: Centropomidae) in Nyanza Gulf of Lake Victoria,East Africa

From bottom trawls, Lates niloticus (Lin.) in Nyanza Gulf was observed to have a gulf-wide distribution. Lates dominated the catch at Homa point. Catch rates in kg h^-1 varied from one location to another and from month to month.Its estimated production is 11 000 metric tons, netting about 22 000 000 Kenya shillings (US $2 000 000).Abundance of food items such as freshwater shrimps of the genus Caridina, and Engraulicypris argenteus, have been used as basis for estimating the growth and production of Lates.     

Reproductive potential of the Nile perch,Lates niloticus L. and the establishment of the species in Lakes Kyoga and Victoria (East Africa)

The size at first maturity, sex ratio and fecundity of L. niloticus in Lake Kyoga have been examined, and compared with the situation in other aquatic systems. The species has the ability to reproduce enormously. It produces up to 16 million eggs. In Lake Kyoga, fecundity (F) increases with length (L) in cm according to the equation: % MathType!MTEF!2!1!+-% feaafiart1ev1aaatCvAUfeBSjuyZL2yd9gzLbvyNv2CaerbuLwBLn% hiov2DGi1BTfMBaeXatLxBI9gBaerbd9wDYLwzYbItLDharqqtubsr% 4rNCHbGeaGqiVu0Je9sqqrpepC0xbbL8F4rqqrFfpeea0xe9Lq-Jc9% vqaqpepm0xbba9pwe9Q8fs0-yqaqpepae9pg0FirpepeKkFr0xfr-x% fr-xb9adbaqaaeGaciGaaiaabeqaamaabaabaaGcbaGaciOraiaac2% dacaGGGaGaaiinaiaac6cacaGG0aGaai4maiaacAdacaGGGaGaaiiE% aiaacccacaGGXaGaaiimamaaCaaaleqabaGaaiylaiaacAdaaaGcca% GGmbWaaWbaaSqabeaacaGGYaGaaiOlaiaacMdacaGGYaaaaaaa!44D8!\[\operatorname{F} = 4.436 x 10^{ - 6} L^{2.92} \]Information from different habitats shows that females grow to a larger size than males but the growth rate is the same in both sexes. Males mature earlier than females at 50–65 cm total length with females maturing between 60–95 cm. There are about twice as many males as females. The rapid establishment of L. niloticus in Lake Kyoga and the Nyanza Gulf of Lake Victoria following its introduction is attributed to the high reproductive potential of the species under favourable environmental conditions.     

Male-male competition and speciation: aggression bias towards differently coloured rivals varies between stages of speciation in a Lake Victoria cichlid species complex

Sympatric speciation driven by sexual selection by female mate choice on a male trait is a much debated topic. The process is problematic because of the lack of negative frequency-dependent selection that can facilitate the invasion of a novel colour phenotype and stabilize trait polymorphism. It has recently been proposed that male-male competition for mating territories can generate frequency-dependent selection on male colouration. Rare male cichlid fish would enjoy a fitness advantage if territorial defenders bias aggression towards male cichlid fish of their own colour. We used blue (ancestral type) and red phenotypes of the Lake Victoria cichlid species complex Pundamilia. We tested the aggression bias of wild-caught territorial blue male cichlid fish from five separate populations for blue vs. red rival male cichlid fish using simulated intruder choice tests. The different populations vary in the frequency of red male cichlid fish, and in the degree of reproductive isolation between red and blue, reflecting different stages of speciation. Blue male cichlid fish from a population that lack red phenotypes biased aggression towards blue stimulus male cichlid fish. The same was found in two populations where blue and red are reproductively isolated sister species. This aggression bias may facilitate the invasion of a novel colour phenotype and species coexistence. Blue male cichlid fish from two populations where red and blue are hybridizing incipient species biased aggression towards red stimulus male cichlid fish. Thus, after a successful invasion of red, aggression bias alone is not likely to generate frequency dependence required to stabilize the coexistence of phenotypes. The findings show that aggression bias varies between stages of speciation, but is not enough to stabilize the process of speciation.     

Effects of a Whole-lake,Experimental Fertilization on Lake Trout in a Small Oligotrophic Arctic Lake

We tested whether increased phosphorus and nitrogen concentrations would affect a lake trout (Salvelinus namaycush) population in a small oligotrophic lake with a benthically dominated food web. From 1990 to 1994, nitrogen and phosphorus were added to Lake N1 (4.4 ha) at the arctic Long-Term Ecological Research site in Alaska. We used mark/recapture methods to determine the lake trout population size, size structure, recruitment, and individual growth from 1987 to 1999. Data were also collected on water chemistry and food availability. Fertilization resulted in increased pelagic primary productivity, chlorophyll a, turbidity, snail density, and hypoxia in summer and winter. Lake trout density was not affected by the manipulation however growth and average size increased. Recruitment was high initially, but declined throughout the fertilization. These results suggest that lake trout were affected through increased food availability and changes to the physical characteristics of the lake. During fertilization, hypoxia near the sediments may have killed over-wintering embryos and decreased habitat availability. Although lake trout responded strongly to increased nutrients, loss of recruitment might jeopardize lake trout persistence if arctic lakes undergo eutrophication.     

Population dynamics of two small cichlid fish species in a tropical man-made lake (Lake Kariba)

The population dynamics of two small cichlid fishes (Pharyngochromis darlingi andPseudocrenilabrus philander) were studied in Lake Kariba, a very large African man-made lake. They are of no economic importance but make up about 14% and 7% respectively of the inshore fish population and are the major components of the diet of fish-eating birds on the lake.P. darlingi isthe larger species (L = 156.5 mm) and is found on both shelving and steep, eroding shores. Its mortality rate differs in each habitat (Z = 0.44 and 0.72 month^–1 respectively), only 0.79% survive for 12 months and its % MathType!MTEF!2!1!+-% feaafiart1ev1aaatCvAUfeBSjuyZL2yd9gzLbvyNv2CaerbuLwBLn% hiov2DGi1BTfMBaeXatLxBI9gBaerbd9wDYLwzYbItLDharqqtubsr% 4rNCHbGeaGqiVu0Je9sqqrpepC0xbbL8F4rqqrFfpeea0xe9Lq-Jc9% vqaqpepm0xbba9pwe9Q8fs0-yqaqpepae9pg0FirpepeKkFr0xfr-x% fr-xb9adbaqaaeGaciGaaiaabeqaamaabaabaaGcbaGaamiuaiaac+% cadaqdaaqaaiaadkeaaaaaaa!384D!\[P/\overline B \] ratio is 5.45 (on shelving shores).Ps. philander is smaller (L = 83.9 mm) and is restricted to shelving areas with abundant vegetation. Its monthly mortality rate was high (Z = 7.69), only 0.05% survive to 12 months whilst its % MathType!MTEF!2!1!+-% feaafiart1ev1aaatCvAUfeBSjuyZL2yd9gzLbvyNv2CaerbuLwBLn% hiov2DGi1BTfMBaeXatLxBI9gBaerbd9wDYLwzYbItLDharqqtubsr% 4rNCHbGeaGqiVu0Je9sqqrpepC0xbbL8F4rqqrFfpeea0xe9Lq-Jc9% vqaqpepm0xbba9pwe9Q8fs0-yqaqpepae9pg0FirpepeKkFr0xfr-x% fr-xb9adbaqaaeGaciGaaiaabeqaamaabaabaaGcbaGaamiuaiaac+% cadaqdaaqaaiaadkeaaaaaaa!384D!\[P/\overline B \] ratio was very high (7.69). The estimates of growth obtained forP. darlingi differ considerably from those given in an earlier study in Lake Kariba and some possible reasons for this are discussed. In suitable habitats, the combined production of both species could be 40 kg ha^–1 yr^–1 which indicates their potential importance to the ecology of the lake.     

Functioning and dynamics of wetland vegetation of Lake Victoria: an overview

The aquatic macrophytic vegetation constituting the wetlands situated along the coast of Lake Victoria provides valuable services to both local and regional communities as well as an important ecological function through the transition between terrestrial and aquatic ecosystems. The wetland vegetation is typically rooted in the substrate on the landward side of the lake, but forms a floating mat towards the middle of the wetland and at the wetland/lake interface. Cyperus papyrus and Miscanthidium violaceum vegetation typically dominate the permanently inundated wetland areas along most of the shores of Lake Victoria. Due to the prevailing climatic and hydrological catchment conditions, these macrophytic plants (papyrus in particular) tend to exhibit high net productivity and nutrient uptake which strongly influences both wetland status and lake water quality. In addition, these wetlands provide important economic livelihoods for the local populations. The integrity and physical structure of these wetlands strongly influences their associated mass transport mechanisms (water, nutrients and carbon) and ecosystem processes. Wetland degradation in Africa is an increasing problem, as these ecosystems are relied upon to attenuate industrial, urban and agricultural pollution and supply numerous services and resources. In an integrated project focused on the wetlands of Lake Victoria, the ecological and economic aspects of littoral wetlands were examined and new instruments developed for their sustainable management.     

The invasion of an introduced predator,Nile perch (<Emphasis Type="Italic">Lates niloticus</Emphasis>, L.) in Lake Victoria (East Africa): chronology and causes

Nile perch, a large predatory fish, was introduced into Lake Victoria in 1954. The upsurge of Nile perch in Lake Victoria was first observed in the Nyanza Gulf, Kenya, in 1979. In Ugandan waters this occurred 2–3 years later and in the Tanzanian Mwanza Gulf 4–5 years later. At the beginning of the upsurge in the Mwanza Gulf in 1983/1984 only sub-adult and adult fishes were found. The first juveniles appeared in 1985, suggesting that the initial increase of Nile perch was mainly caused by migration of sub-adults and adults. Shortly after the onset of trawl fishery in the area in 1973, haplochromines in the Mwanza Gulf started to decline. The final disappearance of the haplochromines, in 1987, only occurred after the Nile perch boom, and despite the abandoning of the haplochromine fishery in 1986. We hypothesize that the decline of haplochromines decreased predation on and competition with juvenile Nile perch and then facilitated survival of these juveniles. Consequently the immigration of sub-adult and adult Nile perch in an area may have paved the way for successful recruitment. Over-exploitation of haplochromine cichlids in the 1970s in the Nyanza Gulf, where the Nile perch upsurge was first observed, may have played a similar role.     

Ladybird population dynamics in potato: comparison of native species with an invasive species, Harmonia axyridis

Following the detection of the harlequin ladybird, Harmonia axyridis, in 2003 in potato crops in Belgium, a study was carried out between 2004 and 2006 on the phenology of this species compared to native species in potato. The results confirmed the success of H. axyridis, with high population levels in 2004 and 2005. In 2006, aphid populations were very low and no H. axyridis larvae were sampled in potato, but the indigenous species Coccinella septempunctata and Propylea quatuordecimpunctata were detected. A species by species comparison of the date of first larvae detection, the larvae population peak, and the difference between this peak and the aphid population peak was performed. Results showed a clear correlation between C. septempunctata and P. quatuordecimpunctata and potato aphids, with a delay of 3.5 and 6.5 days between the aphid and ladybird population peaks for the two native species. H. axyridis arrived 7–8 days after the two indigenous species and the larval peak population occurred 15.8 days after the aphid population peak. This meant that H. axyridis had to complete its larval development with very low aphid populations or even with no aphids at all. The reason for its late arrival and the possible food resources used by H. axyridis larvae are discussed.     

Growth of dominant zooplankton species feeding on plankton microflora in Lake Shira

Batch cultures and continuous flow cultures were used to study the growth rates of zooplankton species from Shira lake, the rotifer Brachionus plicatilis Muller and calanoid copepod Arctodiaptomus salinus Daday, which were fed on phytoplankton and bacterioplankton from the lake. Analyses of the birth and survival rates were used to demonstrate that the lake phytoplankton, consisting mostly of cyanobacteria and diatomaceous algae, is inadequotes for optimal realisation of the reproductive potential of B. plicatilis when compared with the bacterial diet. The study revealed that the kinetic growth characteristics of the two zooplankters were similar: B. plicatilis r _max, 0.120 d^–1; S ₀, 0.253; and K _s, 0.114 mg dry mass l^–1; and for A. salinus r _max, 0.129 d^–1; S ₀, 0.240; and K _s, 0.171 mg dry mass l^–1. Fluctuations in natural food concentration reduced the growth rate of both species. Even though the threshold concentration of food for B. plicatilis and A. salinus were quite similar, the copepods were less sensitive to food limitation.     

Environmental effects of introducing whitefish,Coregonus lavaretus (L.), in Lake Ring

The impact of whitefish (Coregonus lavaretus (L.)) on the trophic structure of eutrophic lakes was studied in Lake Ring, a small eutrophic Danish lake (22.5 ha, mean depth 2.9 m) in which the natural fish fauna is dominated by pike (Esox lucius L.), perch (Perca fluviatilis L.), and eel (Anguilla anguilla (L.)), roach (Rutilus rutilus (L.)) and burbot (Lota lota (L.)) being the only other fish species present. A total of 10993 0+ whitefish were stocked in the lake from October 1989 to July 1990 and the structure of the fish, zooplankton and benthic invertebrate communities studied during the period 1989–1991. Stomach contents analysis revealed that the whitefish mainly ate Daphnia and copepods in 1990–1991, the proportion of copepods decreasing with increasing size of the fish and Daphnia being the overall most important food source. The density of Daphnia in the lake decreased from 72 ind. 1^-1 in 1989 to 9 ind. 1^-1 in 1991; concomitantly the large species Daphnia magna and D. pulex almost disappeared and the density of cyclopoid copepods increased from 72 to 101 ind. 1^-1, presumably because of improved food conditions, while that of calanoid copepods remained virtually unchanged. As a result chl-a increased from 19 to 47 µg 1^-1 and Secchi depth decreased from 2.4 m to 1.7 m, despite there being no change in total P and total N (0.6 mg P 1^-1 and 1.3 mg N 1^-1, respectively). Changes were also observed in the benthic invertebrates; Chaoborus, oligochaetes, and chironomids all decreased, whereas Pisidium increased. It is concluded that the stocking of whitefish in eutrophic lakes for commercial purposes may delay their recovery, or even lead to enhanced eutrophication.     

Female-to-male shift of mouthbrooding in a cichlid fish,Tanganicodus irsacae,with notes on breeding habits of two related species in Lake Tanganyika

Synopsis Parental-care patterns and mating systems of three goby-like cichlids in Lake Tanganyika were investigated. In Tanganicodus irsacae females mouthbrooded eggs and small young for about two weeks and then males took over the role for about one week. Field observations of tagged fish suggest that this species is monogamous: a male's home range largely overlapped with that of its mate, while their home ranges were segregated from those of similar-sized consexual adults. Eretmodus cyanostictus also performed female-to-male shift of mouthbrooding and appeared to be monogamous. The third species, Spathodus marlieri, however, exhibited exclusively maternal mouthbrooding. The differences in parental care and mating system among the three species are discussed in relation to their feeding habits, and the pattern of monogamy in the goby-like cichlids is compared with those of other fishes.     

Food web manipulation in a small,eutrophic Lake Wirbel,Poland: effect of stocking with juvenile pike on planktivorous fish

In a four year experiment (1988–1991) carried out in a shallow, eutrophic lake, an increase in young-of-the-year pike (Esox lucius) density was used to control planktivorous fish assemblages consisting mainly of small-sized cyprinids: roach (Rutilus rutilus), Leucaspius delineatus, and white bream (Blicca bjoerkna). Stocking with pike fry of ca 30 mm in three successive springs resulted in large-scale mortality among prey of vulnerable sizes. Rotenone treatment at the end of the fourth year showed that roach and white bream populations contained few 0+ to 2+ fish, but very high numbers of 3+ to 6+ fish. By this time, short-lived, slow-growing L. delineatus, with no size refuge from predators, had been driven almost to extinction.     

Morphological and Morphometrical Variations of Selected Rotifer Species in Response to Predation: a Seasonal Study of Selected Brachionid Species from Lake Xochimilco (Mexico)

Allelopathy of filamentous green algae (FGA) has been less studied than that of macrophytes. Little Budworth Pool, Cheshire, UK is a small, shallow, clear-water lake with high TP concentrations, very high NO_3-N concentrations, only moderate phytoplankton density, high FGA growth (mainly Spirogyra sp.) and no submerged plants. Experiments were carried out to test the possible allelopathic effects of Spirogyra on the phytoplankton of this lake and on a submerged plant Elodea nuttallii. Changes in phytoplankton growth, phytoplankton species dynamics and species composition were apparently not influenced by allelopathy of live or decaying Spirogyra. A shift from diatom (Cyclotella sp) – cryptomonad (Chroomonas acuta and Cryptomonas erosa) dominance to Chlorococcales (Micractinium pusillum, Monoraphidium contortum and Scenedesmus opoliensis) – Volvocales (Chlorogonium elongatum and Pandorina morum) dominance was recorded in both control and FGA treatments, suggesting an effect of nutrient enrichment. Nutrient concentrations and differences in competitiveness among phytoplankton species can also explain differences in their growth rates in Spirogyra filtrate. Spirogyra also did not influence apex number per plant, shoot length or growth rate of E. nuttallii. This FGA species probably cannot control phytoplankton or E. nuttallii growth in nutrient rich conditions through allelopathy.     

Resilience of Alternative Stable States during the Recovery of Shallow Lakes from Eutrophication: Lake Veluwe as a Case Study

In this paper we analyze a long-term dataset on the recovery from eutrophication of Lake Veluwe (The Netherlands). Clear hysteresis was observed in a number of ecosystem variables: the route to recovery differed significantly from the route that led to loss of clear water. The macrophyte dominated state disappeared in the late 1960s at TP above 0.20 mg l⁻¹, whereas its return occurred at less than 0.10 mg TP l⁻¹. Several regime shifts resulting in the occurrence of three alternative stable states were observed over a period of 30 years. The turbid state showed resistance to change, despite a strong and prompt reduction in Chl-a following reduction of external P-loading. The most important component that determined hysteresis in the return to clear water was not internal P-loading, but a high level of nonalgal light attenuation (through sediment resuspension) maintained by the interaction between wind and benthivorous fish. Although Chara was able to re-colonize the most shallow parts of the lake, recovery stalled and for a number of years clear (above charophyte beds) and turbid (deeper parts of the lake) water co-existed, as a separate alternative state on route to full recovery. Lake-wide clear water was re-established after bream density had been reduced substantially. This allowed a return of zebra mussels to the lake, whose high filtration capacity helped in maintaining clear water. In this study, we were able to identify the main drivers of hysteresis and regime shifts, although formal demonstration of cause and effect was not possible on the basis of field data alone. We argue that resilience of the present clear water state of Lake Veluwe very much depends on sizable populations of a few keystone species, especially Chara (stoneworts) and Dreissena (zebra mussels), and that careful management of these species is equally important as control of nutrients. Lake management should strive to maintain and strengthen resilience of the ecosystem, and this should offer protection against a renewed collapse of the clear state.     

Recovery of species diversity and composition of braconid parasitic wasps after reforestation of degraded grasslands in lowland East Kalimantan

Human-induced disturbances and wildfires can transform areas of tropical rainforest into Imperata-dominated grassland, but it may be possible that recovery of biodiversity is facilitated by reforestation with fast-growing trees. We compared the assemblages of braconid wasps as parasitoids of taxonomically diverse groups of insects among Imperata grasslands, young and mature plantations of Acacia mangium, young secondary forests after wildfires, and old secondary forests in the lowland of East Kalimantan. The abundance and species richness of braconids, which had declined in Imperata grasslands, somewhat increased in Acacia plantations, and also the species composition of braconids in Acacia plantations was transitional between Imperata grasslands and old secondary forests. Parasitoids of detritivores and wood borers increased markedly after plantation, while those of herbivores showed a distinct turnover of species all over the range from grasslands to old secondary forests. The plantation of A. mangium had most likely facilitated the recovery of the diversity of host forest and their parasitoids, but the recovery was just at the rudimentary stage even in mature plantations. Monitoring of parasitic wasps would be useful to test the continuous recovery of forest biodiversity in plantation stands.     

The food and feeding habits of Eutropius niloticus (Ruppell), family Schilbeidae (Osteichthyes: Siluriformes) in Lake Kainji,Nigeria

Studies on the food and feeding habits of Eutropius niloticus in Lake Kainji, Nigeria revealed that the fish is omnivorous but fed more on insects than on any other food. The most important insects were Povilla adusta Navas nymphs and imagos and adults of the common grasshoppers. Fish, particularly the clupeids Sierrathrissa leonensis Thys Van den Audernaerde and Pellonula afzeliusi Johnels, also formed an important part of the diet. There was seasonal variation of organisms selected for food and there were variations in the dietary items selected by different age groups. Accumulation of fat was found to correlate with seasonal feeding intensity. E. niloticus serves as food for other fishes.     

The elicitor-induced oxidative processes in leaves of Solanum species with differential polygenic resistance to Phytophthora infestans

Previous studies have shown that suspension-cultured cells of Solanum genotypes with various polygenic resistances to Phytophthora infestans differed in activities of early oxidative processes in response to culture filtrate (CF) from this pathogen. These studies have now been extended by analysing production of reactive oxygen species (ROS), lipid peroxidation and Lipoxygenase (LOX, E.C.1.13.11.12) activity induced by CF in detached leaves of S. tuberosum cv Bzura and clone H-8105, polygenically resistant and susceptible, respectively, as well as S. nigrum, nonhost, completely resistant. The relative increase in the ROS production was higher in the susceptible clone H-8105 than in both resistant genotypes. Lipid peroxidation increased only in the nonhost S. nigrum. An increase in lipid peroxidation in S. nigrum leaves coincided with enhanced LOX activity. In both S. tuberosum genotypes, significant increases in LOX activity were delayed and unaccompanied by changes in the level of lipid peroxidation. LOX activity attained a higher level in both of the resistant genotypes than in the susceptible one. The present results suggest that the involvement of both ROS production and LOX activity in the defense strategy in Solanum species/P. infestans interactions.     

Cyclomorphosis in Daphnia: an adaptation to avoid invertebrate predation

Seasonal morphological changes in three Daphnia species were followed over a two-year period in two lakes that differ in invertebrate and fish pressure. Whereas the morphology of D. hyalina, the biggest of the three species, varied little from season to season, D. cucullata, the smallest, exhibited the most pronounced seasonal changes in head height/carapace length ratio. The pattern of seasonal changes of body proportions was similar in all size classes and isometric growth of the head was reported for D. cucullata. Unlike the head, tail spine length/carapace length ratio almost did not vary seasonally. Strong negative allometry of tail spine growth was observed. These results are consistent with the hypothesis that helmets and tail spines provide protection against invertebrates in the two smallest, thus most endangered species.