Do Chaoborus larvae migrate in temporary pools?

A diurnal vertical migration of larvae of Chaoborus punctipennis (Say) was observed in shallow temporary woodland pools in East Texas. In the laboratory, in 153 cm tall columns, the larvae underwent a much greater migration than possible in the shallow pools. We hypothesize that the migratory behavior and transparent body of Chaoborus larvae may have originally evolved in such shallow water habitats. These traits enabled Chaoborus to successfully invade the quite different adaptive zone of predation in the plankton of deep lakes.     

Biology of the African clupeid Limnothrissa miodon with reference to its small size in artificial lakes

Summary Limnothrissa miodon is a clupeid from Lake Tanganyika which has been introduced to Lakes Kivu and Kariba and which invaded Lake Cahora Bassa. These lakes differ considerably from one another but the biology of Limnothrissa is similar in many respects in all of them. Similarities include its feeding and breeding biology, whilst its populations fluctuate, on both an annual and a seasonal basis, in relation to environmental changes. The major differences between the lakes concern the size to which Limnothrissa grows. Their average length in the two natural lakes, Lakes Tanganyika and Kivu, is about twice the average length in the two artificial lakes, Lakes Kariba and Cahora Bassa. This in turn affects their age of maturity and they breed during their first year in the reservoirs but during their second in the natural lakes. Their growth rates and performance are comparable up to about 6 months of age and the difference in their size seems to be due to high mortality in the artificial lakes where few Limnothrissa survive for more than a year. Possible explanations for their small size include the genetic composition of the original introduction and the unpredictability of the environment. Neither of these explanations is supported by evidence at present. Deficiencies in the available food and the effects of intense predation are probably the most likely causes. In both natural lakes, Limnothrissa are predatory or cannibalistic from 100 mm in length and 12 months in age, and fish in their diet may enable them to survive into their second year. The fish in Lakes Kariba and Cahora Bassa, in contrast, are neither predatory nor cannibalistic to any degree. There are no predatory fish species in Lake Kivu but there is a diverse community of pelagic piscivores in Lake Tanganyika which tends to contradict the predation hypothesis. However, the position of Limnothrissa in the reservoirs probably resembles that occupied in Lake Tanganyika by Stolothrissa which is also a small, annual species. The significance of this phenomenon is that Limnothrissa can maintain a high biomass and productivity in the face of intense predation. This trait may be of importance to fisheries management because it means that their yield can be greatly increased.     

Impact of Holoshestes heterodon Eigenmann (Pisces: Characidae) on the plankton community of a subtropical reservoir: the importance of predation by Chaoborus larvae

We analyzed the effects of planktivorous Holeshestes heterodon Eigenmann (Characidae) predation on the plankton community of a small subtropical reservoir, using four enclosures (volume about 17.5 m³), open to the sediment, established in the littoral zone. Two enclosures were stocked with fish (mean TL 5.7 cm), at a density of about 4–5 fish m^–3 (approx. 8 g m^–3), whereas two remained fishless. The experiment lasted a little longer than one month. In the fish enclosures, most Crustacea and Chaoborus larvae remained scarce, probably as a result of visually selective fish predation. In both fishless enclosures, Chaoborus larvae became abundant. However, in only one of these did large individuals become relatively numerous; this discrepancy in the demographic structure of the Chaoborus populations between the two fishless enclosures is unexplained. Only in the fishless enclosure without appreciable numbers of large Chaoborus did densities of Crustacea increase greatly. It is suggested that in the enclosure containing large Chaoborus individuals, crustacean populations were prevented from developing due to predation pressure, while the small Chaoborus larvae of the other enclosure could not readily consume these prey. Rotifers were low in abundance in the absence of fish, probably as a consequence of Chaoborus predation. Phytoplankton density increased in all four enclosures, due probably to the lack of water flow. Only in the fishless enclosure with high densities of crustaceans did phytoplankton abundance decrease markedly at the end of the experiment, perhaps because of grazing losses.     

Chaoborus predation and zooplankton community structure in a rotifer-dominated lake

During summer, Chaoborus punctipennis larval densities in the water column of fishless, eutrophic Triangle Lake become very high, and coincidently, the spined loricate rotifer Kelfcottia bostoniensis becomes the dominant zooplankter. Research was done to test the hypothesis that selective predation by Chaoborus on soft-bodied rotifers controls species dominance in the mid-summer zooplankton of this lake. In situ predation experiments showed positive selection by Chaoborus for the soft-bodied Synchaeta oblonga, negative selection for K. bostoniensis, and intermediate selection for Polyarthra vulgaris, a species with rapid escape tactics. However, during a 21 day in situ mesocosm experiment, zooplankton dominance and succession in Chaoborus-free enclosures was identical to that in enclosures with Chaoborus at lake density. Despite the selective predation, Chaoborus larvae may not exert significant top-down control on rotifers, whose intense reproductive output during mid-summer in temperate eutrophic lakes results in new individuals at rates that exceed predatory losses.     

Chaoborus and fish-mediated influences on Daphnia longispina population structure,dynamics and life history strategies

Summary This study examined the long term effects of predation by larvae of the midge Chaoborus and simulated fish predation on experimental Daphnia longispina populations. Chaoborus predation, relative to fish predation, led to populations composed of larger individuals as a whole, larger egg-bearing individuals, and a larger primiparous instar. Daphnia retained helmets beyond the first instar in response to the presence of Chaoborus. Both types of predation, relative to predator-free controls, reduced prey population size and rates of increase, but increased population death rates. The reduction in population size due to predation led to increased resource availability for individuals remaining in the populations and increased individual fecundity in the predation treatments. The differences noted between the Chaoborus, fish, and control treatments increased with predation intensity.     

Influence of benthic and interstitial processes on nutrient changes along a regulated reach of a large river (Rhône River,France)

An enclosure study was conducted in Ranger Lake in south-central Ontario, Canada from 4 July to 5 August 1997 to determine predation effects of the larvae of the phantom midge fly Chaoboruson the zooplankton community. Zooplankton assemblages were established in 12 enclosures (2 m in diameter, 7.5 m deep). Three densities of fourth-instar Chaoborus trivittatus (0 l^–1, 0.1 l^–1 and 0.5 l^–1) were introduced as predator treatments to the enclosures. Temperature, dissolved oxygen and zooplankton community composition were monitored for six weeks. To determine if the zooplankton community composition changed, a repeated measures multivariate analysis was performed on percent biomass of Bosmina and calanoid copepods. There were no significant differences in mean taxon percent biomass among predator treatments. There were significant differences in mean taxon percent biomass between water layers (epilimnion and metalimnion). There were also significant differences in lengths of Bosmina and calanoid copepods among predator treatments at the end of the experiment. Crop content analysis of C. trivittatusshowed that Bosmina constituted 88–98% of the prey items found in the crops. These results demonstrate that the use of deep enclosures, a Chaoborus species which vertically migrates, and lower natural densities of Chaoborus may provide prey with an important natural refuge from predation and so allow a more accurate determination of the predation impact of Chaoborus trivittatusin temperate lakes where fish control Chaoborus densities.     

Morphological and genetic differentiation of the African clupeid Limnothrissa miodon 34 years after its introduction to Lake Kivu

The evolutionary consequences of an artificial introduction of the clupeid Limnothrissa miodon from Lake Tanganyika into Lake Kivu, East Africa were examined. In 1959, 57 400 fry (mixture of Limnothrissa and the related clupeid, Stolothrissa tanganicae ), were released into Lake Kivu to boost fisheries production. Comparisons were made between respective source and transplant populations 34 years later (1993) using morphometrics ('truss' method), allozymes (29 enzyme-coding loci) and mitochondrial (mt) DNA variation (RFLP analysis of PCR-amplified ND5/6 genes). Significant morphological and genetic differentiation between source and transplant samples was detected, with a distinct clustering of Kivu Limnothrissa on respective dendrograms, especially at the morphometric and mtDNA levels. Differentiation within Lake Tanganyika was, however, consistently higher than that between lakes. Allozymic diversity was similar in samples from both lakes (Lake Tanganyika: heterozygosity = 0.0658, mean number of alleles=1.44; Lake Kivu: heterozygosity = 0.0655; mean number of alleles = 1.48), however, a significantly lower mtDNA haplotype diversity was detected in Lake Kivu (Lake Tanganyika: 0.905; Lake Kivu: 0.755). Data suggest that high post-introduction mortality and various demographic factors reduced the effective population size of the introduced population to tens rather than thousands of individuals, resulting in a reduction in genetic diversity and founder effect.     

Molecular phylogenetic investigations of the Viviparidae (Gastropoda: Caenogastropoda) in the lakes of the Rift Valley area of Africa

The freshwater gastropod family Viviparidae is nearly cosmopolitan, but absent from South America. On the African continent, two genera are recognized; the widespread Bellamya and the monotypic Neothauma, which is confined to Lake Tanganyika. Most of the African Bellamya species are confined to the major lakes of the Rift Valley area in Africa, i.e. Lake Albert, Lake Malawi, Lake Mweru, and Lake Victoria. The phylogenetic analyses of mitochondrial (COI and 16S) and nuclear (H3, 18S and 28S) DNA inferred three major lake-clades; i.e. Lake Victoria/Kyoga/Albert, Lake Malawi and Lake Mweru/Bangweulu. The endemic B. rubicunda from Lake Albert and B. unicolor from Lake Kyoga were inferred to be part of the Lake Victoria clade. Bellamya capillata as identified by shell characters was polyphyletic in gene trees. The monophyletic Bellamya species radiation in Lake Malawi was most nearly related to the Lake Victoria/Kyoga/Albert-clade. Taxa from the Zambian lakes, Mweru and Bangweulu, were inferred together and placed ancestral to the other lakes. Neothauma tanganyicense was inferred as the sister-group to the Zambian Bellamya. Within the lake-clades the endemic radiations show very low genetic diversities (0–4.1% in COI), suggesting much faster morphological divergence than molecular divergence. Alternatively, Bellamya in Africa constitutes only a few species with several sub-species or eco-phenotypic morphs. The African viviparids were inferred to be the sister-group to a clade comprising Asian species, and the relatively low genetic diversity between the clades (12.6–15.5% in COI) makes a recent Miocene dispersal event from Asia to Africa much more likely than an ancient Gondwana vicarience distribution.     

Zooplankton Migration in Three Lakes of Western New York

We examined the diel vertical migration (DVM) of zooplankton in three lakes of western New York; Lakes Conesus, Lime, and McCargo. In all three lakes, the dipteran predator Chaoborus was a more dramatic migrator than any of the cladocerans or copepods. In contrast, another invertebrate predator, Leptodora, seemed to have the least vertical dispersal. In Conesus Lake, following the evening ascent and upper-water night-positioning of Chaoborus flavicans, Daphnia pulicaria expanded its own vertical range of dispersal thereby decreasing its degree of overlap with Chaoborus. The vertical distribution of Diaptomus sicilis was mostly below that of Daphnia pulicaria, thus reducing possible competition for food resources. Poor oxygen conditions in the lower waters of Lakes Lime and McCargo restricted all zooplankton (except Chaoborus) to an epilimnetic zone where the regions of niche overlap and predation were narrowed. Such annually-induced restrictions on vertical dispersal are probably common features of numerous stratified lakes with hypolimnetic reducing conditions. No fixed pattern of vertical dispersal or migration is likely in lakes where stratification and non-stratification follow each other seasonally. Vertical migration does provide some flexibility in niche separation, but the potential for chaotic behavior in interacting predators and prey may confound simple explanations of DVM.     

Larval chaoborid mandibles in surface sediments of small shallow lakes in Finland: implications for palaeolimnology

The aim of this study was to assess the value of phantom midge Chaoborus (Diptera: Chaoboridae) larvae as a palaeoenvironmental proxy in northern Europe. The presence of Chaoborus was examined in 80 small shallow lakes across Finland (60–70°N), based on their subfossil mandibles preserved in lake sediments. Chaoborus flavicans was present in 33 lakes, but was the most abundant midge taxon in only one of the lakes. Chaoborus crystallinus/obscurripes mandibles were present in two lakes. Statistical analyses showed that the distribution of Chaoborus was significantly correlated with lake depth and mean July air temperature. Chaoborus was absent in the northern study lakes, whereas the results indicated that C. flavicans was frequently found in small shallow lakes in southern Finland. Chaoborus flavicans was particularly abundant in fish-free, macrophyte-rich, dystrophic lakes, but absent in oligotrophic clear-water lakes having fish. The results of this study indicate that the subfossil mandibles of Chaoborus can provide valuable information in multiproxy palaeolimnological studies, especially when investigating past changes in water level and temperature in small shallow lakes in northern Europe. Guest editors: K. Buczkó, J. Korponai, J. Padisák & S. W. Starratt Palaeolimnological Proxies as Tools of Environmental Reconstruction in Fresh Water     

Zooplankton community structure driven by vertebrate and invertebrate predators

Summary A zooplankton community was established in outdoor experimental ponds, into which a vertebrate predator (topmouth gudgeon: Pseudorasbora parva) and/or an invertebrate predator (phantom midge larva: Chaoborus flavicans) were introduced and their predation effects on the zooplankton community structure were evaluated. In the ponds which had Chaoborus but not fish, small- and medium-sized cladocerans and calanoid copepods were eliminated while rotifers became abundant. A large-sized cladoceran Daphnia longispina, whose juveniles had high helmets and long tailspines as anti-predator devices, escaped from Chaoborus predation and increased. In the ponds which had fish but not Chaoborus, the large-sized Daphnia was selectively predated by the fish while small-and medium-sized cladocerans and calanoid copepods predominated. In the ponds containing both Chaoborus and fish, the fish reduced the late instar larvae (III and IV) of Chaoborus but increased the early instar larvae (I and II). Small- and large-sized cladocerans were scarcely found. The former might have been eliminated by predation of the early instar larvae of Chaoborus, while the latter was probably predated by fish. Consequently, the medium-sized cladocerans, which may have succeeded in escaping from both types of predator, appeared abundantly. The results suggest that various combinations of vertebrate and invertebrate predators are able to drive various kinds of zooplankton community structure.     

Contrasting patterns of body size for Daphnia species that segregate by habitat

Summary We investigate how body size of two coexisting Daphnia species varies among 7 lakes that represent a gradient of predation risk. The two species segregate vertically in stratified lakes; D. galeata mendotae is typically smaller and more eplimnetic than D. pulicaria. The extent of vertical habitat partitioning, however, varies seasonally within and among lakes in apparent response to predation intensity by epilimnetic planktivorous fishes. Daphnia pulicaria uses the epilimnion at low levels of fish predation but is restricted to the hypolimnion under high fish predation, whereas D. galaeta mendotae always utilizes the epilimnion. The species display contrasting patterns of genetic variation in neonate size and size at maturity. D. pulicaria is larger in lakes with higher fish and Chaoborus densities whereas D. galeata mendotae is smaller. This contrast in body size in lakes with high predation is associated with greater habitat segregation in those lakes. In lakes with low predation risk, the two species are similar in body size at birth and maturity.Authorship order alphabetical     

The extant Chaoborus assemblage can be assessed using subfossil mandibles

1. Assemblages of Chaoborus were examined in 80 thermally stratified southern central Canadian Shield lakes to explore whether subfossil mandibles could be useful in assemblage‐level studies of Chaoborus. 2. Chaoborus (Sayomyia) (probably Chaoborus punctipennis in this study region) and Chaoborus flavicans were the most common taxa recorded, while Chaoborus trivittatus was rarer. Chaoborus americanus was not recorded in subfossil assemblages, because no fishless lakes were included in this study. Chaoborus flavicans had higher relative abundances (%) in lakes with higher dissolved organic carbon (DOC), probably because of reduced fish predation in less transparent water. 3. Results from logistic regression indicate that patterns of presence/absence for Chaoborus mandibles in the study lakes were influenced primarily by hypolimnetic oxygen concentration, probably because of the presence or extent of a hypolimnetic refugium from fish predation. 4. Chaoborus species richness in lakes, derived from subfossil assemblages, did not differ significantly from species richness estimates derived from plankton sampling with a net. Patterns of dominance and coexistence [e.g. the widespread co‐occurrence of C. flavicans and C. (Sayomyia)] determined from subfossil assemblages agreed with previous studies of the contemporary living assemblage. 5. These results suggest that subfossil assemblages may be used as an alternative to nocturnal plankton sampling to carry out research on the community ecology of Chaoborus. 6. We propose a hierarchical conceptual model of assemblage‐level patterns of Chaoborus in temperate lakes. Chaoborus americanus dominates in fishless lakes, whereas in lakes with fish Chaoborus is typically absent where there is no anoxic hyplimnion. In lakes with anoxic strata, C. trivittatus tends to dominate in lakes with few fish; in the remaining lakes, C. flavicans and C. (Sayomyia) dominate, although C. flavicans is more relatively abundant in lakes with lower water clarity (higher DOC).     

Unique qualities and special problems of the African Great Lakes

Synopsis The African Great Lakes consist of large, deep rift valley lakes (e.g. Malawi & Tanganyika) and shallower lakes between the Eastern and Western Rifts (e.g. Victoria). They are a group comparable in size to the North American Great Lakes, but are old. Most are seasonally thermally stratified, and wind is the decisive factor that determines the annual cycle of cooling and mixing. Lakes Tanganyika, Malawi and Kivu are meromictic, with deep relict hypolimnia. Large magnitudes and time scales of periodic internal motion, where these have been measured, appear unique among lakes. These lakes harbour the world's richest lacustrine fish faunas, and the family Cichlidae provides the supreme example of geographically circumscribed vertebrate evolution. The lakes provide a unique comparative series of natural laboratories for evolutionary studies. Primary production is generally high, but in the deeper lakes standing stocks of plankton and of small fish species are low. These pelagic populations are characterised by very high P:B ratios. The fisheries are productive and of socio-economic importance. Large-scale mechanised fishing is not compatible with the survival of the diverse fish communities. Cichlids appear especially vulnerable to unselective fishing. Aquatic reserves might offer a means of survival for at least some communities. Various pollution threats exist. Because water retention times are long, extremely long for some deep lakes, and flushing rates are low, the lakes are vulnerable to pollution which would be long-lasting. Introductions of alien fishes have mostly had undesirable or disastrous results. While the faunas are one of the significant natural heritages of mankind, their conservation must realistically be linked to the legitimate development of the lakes for the well-being of the people who live there. Scientific value alone will not protect the lakes. Just as survival of African terrestrial wildlife in extensive reserves depends heavily upon tourism, so also might the cichlid flocks in underwater reserves. Greater interest from the international scientific community is needed to further rational development and conservation of these great lakes.Invited Editorial     

Creating conditions for changes in prey community structure by Chaoborus spp. in a lake in Sweden

In Lake G»rdsjön (Southwest Sweden), liming as an experimental improvement of living conditions for pelagic algae, resulted in a significant increase of algal biomass and a reduction of mean cell size. The algal development was beneficial for small sized filter feeding zooplankton, particularly rotifers, which showed a significant increase. The increase in abundance of small sized zooplankton created better food conditions for the smaller instars, and thus a much better overall survival of Chaoborus larvae. The resulting, 6–7 times larger population of Chaoborus larvae significantly changed the structure of the crustacean zooplankton community. Bosmina coregoni, the fastest swimmer of the crustacean species suffered most and was strongly reduced by the increased predation from Chaoborus. The share of cladocerans decreased, while copepods increased in importance.     

Influence of Chaoborus density on the effects of an insecticide on zooplankton communities in ponds

Zooplankton communities with a high or low density of Chaoborus larvae were established in outdoor concrete ponds, to which a carbamate insecticide, carbaryl, was applied at 0.1 or 0.5 mg l⁻¹. The lower concentration of the chemical was harmful only to Cladocera. The higher concentration damaged Chaoborus, Copepoda, and some rotifer species, as well as Cladocera. In the ponds with a low density of Chaoborus, chemical application altered the cladoceran community from dominance by Daphnia to that by Bosmina and Moina. In the ponds with a high density of Chaoborus, Chaoborus excluded cladocerans from the zooplankton community presumably by predation, and supported the dominance of rotifers. Cladocera did not recover after application of the chemical, even when Chaoborus was eliminated by the higher concentration of chemical. The relatively rapid recovery of Chaoborus seemed to interrupt the recovery of Cladocera.     

Abundance and distribution of picoplankton in tropical, oligotrophic Lake Kivu, eastern Africa

1. We used flow cytometry to characterize freshwater photosynthetic picoplankton (PPP) and heterotrophic bacteria (HB) in Lake Kivu, one of the East‐African great lakes. Throughout three cruises run in different seasons, covering the four major basins, phycoerythrin‐rich cells dominated the PPP. Heterotrophic bacteria and PPP cell numbers were always high and spatial variations were modest. This represents an important difference from temperate and high latitude lakes that show high fluctuations in cell abundance over an annual cycle. 2. Three populations of picocyanobacteria were identified: one corresponded to single‐cells (identified as Synechococcus by epifluorescence microscopy, molecular methods and pigment content), and the two other that most probably correspond to two and four celled colonies of the same taxon. The proportion of these two subpopulations was greater under stratified conditions, with stronger nutrient limitation. 3. High PPP concentrations (c. 10⁵ cell mL^?1) relative to HB (c. 10⁶ cell mL^?1) were always found. Lake Kivu supports relatively less bacteria than phytoplankton biomass than temperate systems, probably as a consequence of factors such as temperature, oligotrophy, nutrient limitation and trophic structure. 4. A review of PPP concentration across aquatic systems suggests that the abundance of Synechococcus‐like cyanobacteria in large, oligotrophic, tropical lakes is very high. 5. Photosynthetic picoplankton cell abundances in the oligotrophic tropical lakes Kivu and Tanganyika are comparable to those of eutrophic temperate lakes. This apparently contradicts the view that PPP abundance increases with increasing eutrophy. More data on PPP in tropical lakes are needed to explore further this particular pattern.     

A comparison between predation effects on zooplankton communities by Neomysis and Chaoborus

Community level effects of predation by two invertebrate predators, the opossum shrimp (Neomysis intermedia), and the larva of the phantom midge (Chaoborus flavicans) were studied and compared. N. intermedia appeared abundantly in the shallow eutrophic Lake Kasumigaura and had a significant impact on the zooplankton community. The predation pressure by Neomysis was highest on cladocerans, followed by rotifers, and finally copepods. At high densities (maximum nearly 20 000 individuals m^–2), Neomysis excluded almost all cladocerans, rotifers and copepods from the lake.Zooplankton communities were established in experimental ponds, into which C. flavicans was introduced. The predator's density was around 1 individual l^–1, and was probably controled by cannibalism. Although Chaoborus larvae feed on various zooplankton species, their predation impact on zooplankton populations was markedly selective. They eliminated medium- and small-sized cladocerans and calanoid copepods from the ponds, but rotifers increased.Although the feeding selectivities of Neomysis and Chaoborus individuals were similar, the predation effects on zooplankton communities by the two predators were different.     

A metalimnetic oxygen minimum indirectly contributing to the low biomass of cladocerans in Lake Hiidenvesi – a diurnal study on the refuge effect

The diurnal vertical migrations of smelt (Osmerus eperlanus), larvae of phantom midge (Chaoborus flavicans) and cladoceran zooplankton in eutrophic Lake Hiidenvesi were studied in order to clarify the factors behind the low zooplankton biomass. In the study area, an oxygen minimum occurred in the metalimnion in the 10–15 m depth. No diurnal fluctuations in the position of the minimum were observed. Cladocerans inhabited the epilimnion throughout the study period and their vertical movements were restricted to above the thermocline and above the oxygen minimum. C. flavicansconducted a diurnal migration. During the day, the majority of the population inhabited the 12 – 15 m depth just in the oxygen minimum, while during darkness they were found in the uppermost 8 m. Smelts started ascending towards the water surface before sunset and reached the uppermost 3 m around 23:00. During daytime, the majority of smelts inhabited the depth of 7–9 m, where the water temperature was unfavourably high for them (18 °C). Smelts thus probably avoided the steep oxygen gradient in the metalimnion, whereas Chaoborusused the oxygen minimum as a refuge against predation. Those smelts that were found in the same water layers as Chaoborusused the larvae as their main prey. The metalimnetic oxygen minimum thus seemed to favour the coexistence of vertebrate and invertebrate predators, leading to a depression of cladoceran zooplankton.     

Chaoborus predation and the function of phenotypic variation in Daphnia

To investigate the role of helmet formation in defense against predation, laboratory experiments were used to analyze the effects of morphological changes in Daphnia on susceptibility to Chaoborus predation. Behavioral observations of Chaoborus preying on helmeted and non-helmeted Daphnia suggest pre-contact advantages for helmeted prey but post-contact advantages for non-helmeted prey. Helmeted Daphnia are better at evading capture by Chaoborus but may also be more easily handled by the predator. Swimming behavior of the prey, which is influenced by the presence of a tailspine, may affect Chaoborus strike distance. These results re-emphasize the potential hydromechanical importance of body shape changes in defense against predation.