The Ecology of Baetis muticus and Baetis rhodani (Insecta,Ephemeroptera), with Special Emphasis on Acid Water Backgrounds

The two mayflies (Insecta, Ephemeroptera) Baetis muticus and Baetis rhodani are absent from the acid water (pH 4.8–5.2) streams of the Upper Duddon drainage basin. Field observations and laboratory experiments were used in an attempt to explain this absence. Baetis muticus could not tolerate acid water; in addition it was deduced that available foods in the Upper Duddon streams would not sustain it there, even if the water could be tolerated. Baetis muticus feeds by browsing on decaying, allochthonous leaves, implying an input of micro-organisms, especially fungi, to the diet. Well-decayed, allochthonous leaves do not occur in the acid water, Upper Duddon streams. Baetis rhodani proved to be acid water tolerant, but only if the background waters had high ionic loadings. Since Upper Duddon waters have only low ionic loadings, Baetis rhodani is apparently excluded from them on this account. The evidence was that if the Upper Duddon waters were chemically suitable, then Baetis rhodani could survive in them, because a suitable food, the alga Hormidium subtile is present.     

Stream drift as a consequence of disturbance by invertebrate predators

Summary We carried out an experimental field study in a Swedish stream in order to determine whether mobile predators enhance the drift of stream insects. We increased the density of nymphs of the predaceous perlid stonefly, Dinocras cephalotes, in an experimental section of a stream up to densities in another more densely populated part of the same stream. The drift of several benthic species increased significantly compared to a control section where D. cephalotes were rare. Experiments carried out in September showed a strongly elevated drift response in nymphs of the mayfly Baetis rhodani only, whereas May experiments resulted in increased drift in B. rhodani as well as the amphipod Gammarus pulex, the stonefly Leuctra fusca, chironomids, and the total number of drifting animals. In September, we found that the drift response of Baetis rhodani to predator disturbance was dependent on the size of mayfly nymphs; small nymphs appeared in greater numbers in the drift nets than did large nymphs. A subsequent laboratory analysis of drift lengths of B. rhodani nymphs supported the hypothesis that small nymphs travel in the drift for longer than do large nymphs, particularly in darkness. We suggest that morphological constraints in vision or swimming performance, or both, cause small nymphs to drift longer. In May, size-dependent drift was less obvious, probably because the size of the nymphs was considerably greater than in September.     

The status of the Mayfly species described by O. A. Tshernova from the Amur basin in the genus Baetis leach (Ephemeroptera, Baetidae)

Type specimens of Baetis obtusiceps, B. obscuriventris, B. diversicolor and B. mongolicus have been reexamined; lectotypes of B. obtusiceps, B. obscuriventris and B. mongolicus are designated. The following new synonymy is suggested: Baetis vernus Curtis, 1834 = B. obtusiceps Tshernova, 1952, syn. n.; Baetis fuscatus (Linnaeus, 1761) = B. diversicolor Tshernova, 1952, syn. n.; Labiobaetis tricolor (Tshernova, 1928) = Baetis obscuriventris Tshernova, 1952; = B. mongolicus Tshernova, 1952, syn. n.     

Intensive sampling and transplantation experiments reveal continued effects of episodic acidification on sensitive stream invertebrates

1. Although mean pH is increasing in acidified streams throughout Europe, benthic invertebrates are recovering only slowly. One developing hypothesis is that acid episodes continue to restrict recolonisation and recovery. Here, we used intra‐annual sampling in replicate circumneutral, acid and episodic streams at the Llyn Brianne experimental catchments to assess possible mayfly exposure to episodic effects quantified from a risk index based on long‐term discharge. Episodic effects were then tested using short‐term transplantations of animals in enclosures. 2. No mayflies occurred in acid streams while Baetis rhodani occurred in episodic streams but declined to zero density following low pH in autumn. Ephemerella ignita, Baetis vernus, B. muticus, Rhithrogena semicolorata, Ecdyonurus spp. and Heptagenia lateralis occurred only in the circumneutral streams. The first two species were present as nymphs only during July and August so that episodic exposure risk was minimal, but the remaining species occurred as nymphs in all months. 3. Baetis rhodani transplanted during base‐flow (September 2003) and high‐flow (April 2004) were exposed to either; (i) continual (chronic) exposure in the acid streams over 16 days or (ii) repeated short‐term (episodic) exposure to acid conditions for 2 × 4 day periods interspersed with 4‐day recovery periods in the circumneutral streams. Baetis survival in the circumneutral streams (always pH > 5.7) remained high during both low‐ and high‐flow. By comparison, mortality increased (P < 0.01) during chronic and episodic exposure, but only during high‐flow (mean pH 3.8–3.9, cf. 5.5–5.8 at low flow) when mortality varied significantly in the order chronic (>80%) > episodic (>40%) >circumneutral (<10%). 4. We conclude that, despite Europe‐wide trends towards chemical recovery from acidification, even short exposures to high‐flow events at Llyn Brianne are still sufficiently acid to reduce the survival of B. rhodani. Most mayfly species absent from acid and episodic streams have life cycles that would increase acid exposure risks during autumn and winter, and this may be sufficient to explain their current distribution.     

Predator effects on prey population dynamics in open systems

Animal population dynamics in open systems are affected not only by agents of mortality and the influence of species interactions on behavior and life histories, but also by dispersal and recruitment. We used an extensive data set to compare natural loss rates of two mayfly species that co-occur in high-elevation streams varying in predation risk, and experience different abiotic conditions during larval development. Our goals were to generate hypotheses relating predation to variation in prey population dynamics and to evaluate alternative mechanisms to explain such variation. While neither loss rates nor abundance of the species that develops during snowmelt (Baetis bicaudatus) varied systematically with fish, loss rates of the species that develops during baseflow (Baetis B) were higher in streams containing brook trout than streams without fish; and surprisingly, larvae of this species were most abundant in trout streams. This counter-intuitive pattern could not be explained by a trophic cascade, because densities of intermediate predators (stoneflies) did not differ between fish and fishless streams and predation by trout on stoneflies was negligible. A statistical model estimated that higher recruitment and accelerated development enables Baetis B to maintain larger populations in trout streams despite higher mortality from predation. Experimental estimates suggested that predation by trout potentially accounts for natural losses of Baetis B, but not Baetis bicaudatus. Predation by stoneflies on Baetis is negligible in fish streams, but could make an important contribution to observed losses of both species in fishless streams. Non-predatory sources of loss were higher for B. bicaudatus in trout streams, and for Baetis B in fishless streams. We conclude that predation alone cannot explain variation in population dynamics of either species; and the relative importance of predation is species- and environment-specific compared to non-predatory losses, such as other agents of mortality and non-consumptive effects of predators. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Mortality of the terrestrial adult and aquatic nymphal life stages of Baetis vernus and Baetis rhodani in the Breitenbach, Germany (Insecta: Ephemeroptera)

1. The mortality of Baetis vernus Curtis and Baetis rhodani Pictet during the terrestrial-aerial and aquatic life stages was studied at the Breitenbach near Schlitz, Hesse, Germany. The number of females emerging from the stream was recorded with emergence traps. To estimate mortality of females of both species during terrestrial life, numbers of emerging females were compared with numbers of females returning to the stream for oviposition, as shown by numbers of egg masses found in the stream. 2. Mortality of female B. vernus during their terrestrial life stage was 98.8%. It was 91.2 and 96.6%, respectively, during the first and second generations of B. rhodani. 3. To estimate the mortality of both sexes of B. rhodani during the aquatic life stage, the number of eggs laid by the first generation was compared with the number of adults emerging during the second generation. 4. Mortality during the aquatic phase (both sexes combined) of B. rhodani was 91.1%. 5. Mortality during the terrestrial life stages was at least as high, if not higher than during the aquatic stages. Evidently, there is a considerable export of organic material into the terrestrial environment around the stream. Mortality during terrestrial life may be an important regulator of population size.     

Egg envelopes of Baetis rhodani and Cloeon dipterum (Ephemeroptera, Baetidae): a comparative analysis between an oviparous and an ovoviviparous species

A comparative ultrastructural investigation on the eggshell (vitelline and chorionic envelopes) has been carried out in the nymphs of two mayfly species encompassed into the Baetidae, namely Baetis rhodani and Cloeon dipterum. During oocyte differentiation in the meroistic telotrophic ovarioles, gametes are connected to discrete nurse cells by trophic cords. In B. rhodani, each ovariole contains several eggs arranged in sequence, whereas in C. dipterum each contains a single egg. Follicle cells are competent for vitelline and chorionic envelope synthesis. Baetis rhodani is an oviparous species and the chorion is fairly thick, formed by an alveolate endochorion and a fibrillar exochorion delimited by a honey‐comb roof. Cloeon dipterum stands out among Ephemeroptera for being ovoviviparous. In B. rhodani, ovulation starts in the older nymphs with dark wing‐pads, whereas in C. dipterum choriogenesis ends in the imaginal stage. Here the chorion is very thin and laid eggs hatch almost immediately, allowing the larvulae to move out. The maturation of a single egg per ovariole is synchronized with the achievement of the adult stage. The acquisition of ovoviviparity has led to remarkable changes in the ovariole organization along with a simplification of the eggshell structure.     

Escape from Gregarine Parasites Affects the Competitive Interactions of an Invasive Mosquito

When a species is introduced into a new location, it may escape, at least temporarily, from its natural enemies. In field surveys, we found that when the exotic, invasive mosquito, Aedes albopictus, invades new sites, it initially experiences reduced infection by its gut parasite, Ascogregarina taiwanensis. To determine the effect of this escape from parasitism on the competitive ability of A. albopictus, we performed a laboratory competition experiment in which infected and uninfected A. albopictus larvae were reared in microcosms alone and in competition with larvae of the native mosquito, Ochlerotatus triseriatus. We analyzed the effect of parasitism by A. taiwanensis on A. albopictus performance when subjected to intra- and interspecific competition across a range of larval densities, as well as the effect of A. albopictus parasitism by A. taiwanensis on the competitive impact of A. albopictus on O. triseriatus. At a density of 30 O. triseriatus larvae, O. triseriatus survivorship was significantly reduced by the addition of 30 unifected A. albopictus, but not by addition of 30 infected A. albopictus, and not by addition of 15 A. albopictus whether infected or uninfected. Although estimated finite rate of population increase (') showed similar trends, and was significantly affected by treatments, no pairwise differences in rate of increase were significant. Infection by A. taiwanensis also significantly prolonged A. albopictus female development time and reduced the intraspecific competitive effect of increased density of A. albopictus, but did not affect A. albopictus survivorship, mass, or estimated finite rate of population increase. Thus, when A. albopictus escapes from this parasite as it colonizes new sites, this escape may give it a small, but significant, added competitive advantage over O. triseriatus, which may facilitate range expansion of A. albopictus and enhance A. albopictus's initial impact on resident species.     

Systematic revision of the genus Reinboldiella (Ceramiaceae,Rhodophyta) from Taiwan based on comparative morphology and rbcL sequence analyses,including two new species of Reinboldiella

The genus Reinboldiella essentially consists of tiny, membranous blades, epiphytic on benthic red algae, with a wide distribution in the north-western Pacific Ocean (Taiwan, Korea and Japan). In this study, we re-examined the generitype R. schmitziana from Taiwan and neighbouring islands. Two new species of Reinboldiella, R. orientalis sp. nov. and R. taiwanensis sp. nov. are recognized from Taiwan based on comparative morphological studies and molecular analyses of RuBisCO large subunit (rbcL) gene sequences. Reinboldiella orientalis is characterized by its rosette-like appearance, with both erect and prostrate, membranous flat blades whereas R. taiwanensis is distinguishable by mainly consisting of erect, membranous blades with few, short prostrate blades in basal parts of the thalli. Thalli of R. schmitziana also have a rosette-like appearance, but their erect flat blades have cylindrical bases or stipes. The rbcL phylogenetic analyses also support the separation of R.‘schmitziana’ from Taiwan and R. schmitziana from Japan and Korea.     

Population structure and dynamics of Pinus taiwanensis Hayata at Songyang county,Zhejiang Province,China

Pinus taiwanensis is a widely distributed species in the southeastern China (Zhejiang, Anhui, Hubei, Human, Jiangxi, Fujian, Taiwan provinces) at an elevation of 700–2000 m. This pine is a pioneer in forest succession and is often used as a species for afforestation in this region at an elevation above 700 m. This study was carried out at Guanshanyuan, Zhejiang province, at a latitude of N 28° 18, a longitude of E 119° 16 and an altitude of 800–1502 m above sea level. On the basis of a census of all individuals of Pinus taiwanensis at different successional stages and various habitats, age structure, spatial pattern, density, biomass of population and their dynamics were described. Considering the population dynamics throughout the successional process, three phases could be recognized. Until about 9–10 years after Pinus taiwanensis invaded the stands, the density of population was increased by the recruitments along with increase of the mean tree weight and population biomass (phase I). Thereafter, the population was in full density state, the biomass of population and the mean tree weight increased exponentially, while the density was decreased drastically by the self-thinning and the invasion of other broad-leaved trees (phase II). The –3/2 power law of natural thinning was applicable to the populations in this phase. When the broad-leaved trees reached the canopy, although the mean tree weight increased slowly, the density and biomass of Pinus taiwanensis population decreased gradually (phase III) until the population senesced and retreated from the successional series completely. The population dynamics of Pinus taiwanensis during the successional process was in common with pioneer species in forest succession. At some special habitats such as rocky steep slopes and ridges, however, Pinus taiwanensis population could form such an edaphic climax community that the population density, biomass and the mean tree weight in phase III could be in a stable state for very long period.     

木芙蓉三个品种及近缘种的叶绿体基因组比较分析

木芙蓉(Hibiscus mutabilis)栽培历史悠久,是原产中国的古老园林树种和药用植物。为了探讨木芙蓉品种及近缘种的进化特征,厘清木芙蓉品种间及其与近缘种间的亲缘关系,以及探究木芙蓉叶绿体基因组(chloroplast DNA, cpDNA)的遗传方式,该文选择了一个杂交组合中的3个木芙蓉栽培品种(‘单瓣白’‘金秋颂’‘牡丹粉’),用高通量测序平台Illumina NovaSeq对其cpDNA进行首次测序。经组装注释后得到3条完整的cpDNA序列,结合该团队已经完成的近缘种台湾芙蓉(H.taiwanensis)和来自基因库的木槿、朱槿的cpDNA,对木槿属4种及木芙蓉种下的3个品种进行了cpDNA组成和结构特征的比较分析,并完成了其系统发育树重建。结果表明：(1)‘单瓣白’‘金秋颂’‘牡丹粉’的cpDNA序列长度分别为160 880、160 879、160 920 bp,基因数目均为130个,其中蛋白编码基因85个、核糖体RNA 8个和转运RNA 37个。(2)比较分析结果显示,木芙蓉的种下3个品种及其近缘种台湾芙蓉在cpDNA上高度保守,反向重复区(IR)均为26 300 b...     

Nested species subsets, gaps, and discrepancy

Chemical cues from fish can alter the behaviour of stream invertebrates in experimental tanks but their effect in natural streams has received little attention. By adding brook trout (Salvelinus fontinalis) odour to a trout stream in the Rocky Mountains of Colorado, USA, we tested whether changes in the concentration of chemical cues from visually feeding predatory fish would alter the drift of mayfly nymphs (Ephemeroptera). Stream water was piped from stream-side tanks with (odour) and without (control) three brook trout to two locations in the stream 3.5 m upstream of drift nets at six replicate sites. Five-minute drift samples were collected downstream from odour and control pipes before, during and after the release of water from the tanks into the stream during both the day and night. Almost all drift occurred at night and consisted predominantly of Baetis bicaudatus nymphs. The odour manipulation had no measurable effect on Baetis drift during the day but statistical power was low. During the night, however, the drift of large (>0.65 mm head capsule width, HCW) Baetis nymphs decreased significantly during the odour addition compared to control drift. In contrast, the drift of small nymphs (≤0.65 mm HCW) increased both during and after the odour addition in comparison to control drift. Since the stream contains brook trout (0.04–0.18 m⁻²), and water from the stream (presumably containing fish odour) altered the behaviour of fishless-stream Baetis nymphs in another experiment, we conclude that the changes in Baetis drift density were a response to an increase in the concentration of fish odour in the stream. Furthermore, we were able to detect the effect within 5 min. of odour addition, indicating that mayfly behavioural response to trout odour was rapid. These results suggest that mayflies can distinguish different concentrations of trout odour in natural streams and that the response is size-specific, according to the relative risk of predation of large and small Baetis. Received: 12 May 1998 / Accepted: 23 October 1998     

戴云山黄山松群落生态（火用）空间异质性

热力学指标生态（火用）作为传统群落结构指数的补充,通过衡量物种包含的遗传信息量化群落生物热力学空间结构,评估其群落稳定性和健康程度。基于生态（火用）理论研究了戴云山黄山松（Pinus taiwanensis）群落乔木层、灌木层生态（火用）和结构（火用）的空间异质性及分布规律。主要结果如下：（1）乔木层和灌木层分别是群落生态（火用）和结构（火用）的主要贡献者。（2）黄山松种群在群落中具有更高的竞争能力和生存概率,群落生态（火用）贡献率为98.39%,对维持群落结构稳定性具有关键作用。（3）乔灌层的生态（火用）和结构（火用）在研究尺度上均具有强烈空间自相关,其空间异质性主要由结构性因素引起,所占比例大于75%。（4）乔灌层的生态（火用）和结构（火用）均呈明显条带状及斑块状分布,有多个明显高值区,等值线较弯曲且密集,空间异质性程度较高。研究确定了戴云山黄山松种群的热力学地位,黄山松在维系戴云山群落结构稳定和群落健康中发挥着重要作用,结果可为黄山松群落演替动态和可持续管理提供科学依据。     

Species composition and distribution patterns of baetid nymphs (Baetidae: Ephemeroptera) in a Japanese stream

Species composition and distributional patterns among nymphs of five baetid genera (Ephemeroptera), Baetis, Tenuibaetis, Labiobaetis, Nigrobaetis and Alainites were investigated in Yura Stream, Kyoto Prefecture. I collected 13 species: B. sahoensis, B. thermicus, B. sp. F, B. sp. J, B. sp. M1, B. sp. S1, T. sp. E, T. sp. H, L. sp. G, N. chocoratus, N. sp. D, N. sp. I and A. yoshinensis, among which B. thermicus, B. sp. S1 and T. sp. E were dominant, whereas B. sahoensis, B. sp. F, B. sp. M1 and N.sp. I were scarce. Based on their longitudinal distribution patterns, the 13 species were classified into upper species, upper-middle species, middle species, middle-lower species and lower species. Baetis thermicusand A. yoshinensis showed long downstream tails. Baetis sp. J and N. sp. D extended their longitudinal distribution upstream in summer. With regard to habitat preference, Alainites and Labiobaetis were restricted to riffle and vegetated zones, respectively. Tenuibaetis consisted of riffle-vegetated zone species, whereas Baetis and Nigrobaetiscontained both riffle species and ubiquitous species. Habitat partitioning (`sumiwake') along the watercourse (macro-sumiwake) was evident in Tenuibaetis, and that between habitat types (micro-sumiwake) in Labiobaetis vs. Baetis (rhodanigroup species) and Labiobaetis vs. Alainites.     

Constitutively solvent-tolerant Pseudomonas taiwanensis VLB120∆ C∆ ttgV supports particularly high-styrene epoxidation activities when grown under glucose excess conditions

Solvent-tolerant bacteria represent an interesting option to deal with the substrate and product toxicity in bioprocesses. Recently, constitutive solvent tolerance was achieved for Pseudomonas taiwanensis VLB120 via knockout of the regulator TtgV, making tedious adaptation unnecessary. Remarkably, ttgV knockout increased styrene epoxidation activities of P. taiwanensis VLB120Δ C. With the aim to characterize and exploit the biocatalytic potential of P. taiwanensis VLB120Δ C and VLB120Δ CΔ ttgV, we investigated and correlated growth physiology, native styrene monooxygenase (StyAB) gene expression, whole-cell bioconversion kinetics, and epoxidation performance. Substrate inhibition kinetics was identified but was attenuated in two-liquid phase bioreactor setups. StyA fusion to the enhanced green fluorescent protein enabled precise enzyme level monitoring without affecting epoxidation activity. Glucose limitation compromised styAB expression and specific activities (30–40 U/g _CDW for both strains), whereas unlimited batch cultivation enabled specific activities up to 180 U/g _CDW for VLB120Δ CΔ ttgV strains, which is unrivaled for bioreactor-based whole-cell oxygenase biocatalysis. These extraordinarily high specific activities of constitutively solvent-tolerant P. taiwanensis VLB120∆ C∆ ttgV could be attributed to its high metabolic capacity, which also enabled high expression levels. This, together with the high product yields on glucose and biomass obtained qualifies the VLB120∆ ttgV strain as a highly attractive tool for the development of ecoefficient oxyfunctionalization processes and redox biocatalysis in general.     

Historical biogeography and phylogenetic relationships of the genus Chamaecyparis (Cupressaceae) inferred from chloroplast DNA polymorphism

The genus Chamaecyparis comprises five species and one variety native to Taiwan, Japan, Canada, and USA, which demonstrates a classical eastern Asian, western North American, and eastern North American disjunct distributional pattern. The phylogenetic relationships of the species of Chamaecyparis were inferred by comparing 1130 bp of the combined data set of chloroplast trnV intron and petG-trnP intergenic spacer. The phylogenetic tree shows that Chamaecyparis nootkatensis (Cupressus nootkatensis or Xanthocyparis nootkatensis) is clearly diverged from other Chamaecyparis species. For Chamaecyparis species, C. thyoides is sister to C. formosensis and C. pisifera and these together form a monophyletic group. C. lawsoniana is sister to C. obtusa and C. taiwanensis; and these form another monophyletic group. Homogeneity in evolutionary rates was found among species in these two monophyletic groups. Results indicate the divergent evolution of C. taiwanensis and C. formosensis and molecular evidence in this investigation supports C. taiwanensis as a variety of C. obtusa. Utility of cpDNA intergenic spacer petG-trnP in Chamaecyparis is also discussed. Several biogeographical implications were inferred: (1) at least two divergence events have produced the eastern Asian, and both western and eastern North American disjunct distribution in Chamaecyparis; (2) intercontinental sister species pairs are found in Chamaecyparis; (3) cpDNA divergence between two intercontinental sister pairs of C. thyoides and C. pisifera, and C. lawsoniana and C. obtusa is 2.8% and 1.1%, which suggest an estimated divergence time of 14 and 5.5 million years ago during middle and late Miocene, respectively; (4) cpDNA divergence of two Asian Chamaecyparis groups between C. obtusa and C. taiwanensis, and between C. pisifera and C. formosensis is 0.25% and 0.57%, which suggest an estimated divergence time of 1.3 and 2.9 million years ago during Pleistocene and late Pliocene, respectively; these estimated divergence times suggest a relatively recent migration of Chamaecyparis to Taiwan from the Japanese Archipelago; (5) that climatic deterioration caused the disappearance of Chamaecyparis in continental Asia is probable.     

Life history and larval feeding of some species of Ephemeroptera and Plecoptera (Insecta) in the Sierra Nevada (Southern Iberian Peninsula)

We monitored two high mountain streams from the Sierra Nevada (Southern Spain) monthly throughout an annual cycle, measuring and analyzing physical and chemical parameters. Temperature was hourly registered. We studied life cycles and larval feeding of four mayfly species [Serratella ignita (Poda, 1761), Serratella spinosa nevadensis (Alba-Tercedor, 1981), Alainites muticus (Linnaeus, 1758), and Baetis alpinus Pictet, 1843] and two stonefly species [Amphinemura triangularis (Ris, 1902) and Capnioneura mitis Despax, 1932]. Some species changed their strategies (growth rate, life cycle duration, and/or voltinism) in accordance with different conditions (depending on the altitude and temperature). The gut content data showed that species fed mainly on detritus (as collector-gatherers or as shredders), except B. alpinus that behaved as a scraper in one of the streams. Differences in the larval diet of the same species in different streams, and even through its life, support the idea of not using general models to extrapolate the function of a species from a higher taxonomical level. Handling editor: D. Dudgeon     

Diel vertical movements by lotic mayfly nymphs under variable predation risk

1. The diel foraging periodicities of two grazing mayfly (Ephemeroptera) nymphs, Heptagenia dalecarlica and Baetis rhodani, under variable fish (European minnow) predation risk were examined in a series of laboratory experiments. 2. Heptagenia dalecarlica were almost exclusively nocturnal in their use of feeding areas on stone tops. There was a sharp increase in the proportion of nymphs out of refuge at nightfall, both in the control and fish-odour treatments. In a treatment with freely moving fish, H. dalecarlica decreased their activity during both the day and night. In feeding trials with three freely foraging minnows, nymphs were completely safe when they had access to refuges beneath stones, whereas almost half the nymphs were consumed when no refuges were available. 3. Baetis rhodani nymphs reduced their use of stone tops when exposed to four caged minnows or a freely moving fish, but this occurred both day and night. In feeding trials, B. rhodani were captured only while in the water column, and their mortality risk was independent of refuge availability. 4. It is suggested that because H. dalecarlica lack efficient post-encounter defences, they must rely on pre-encounter mechanisms to reduce the threat of fish predation. It appears that in order to dwell sympatrically with fish, H. dalecarlica have evolved a coexistence by hiding strategy. In contrast, Baetis are vulnerable to fish attacks only if they enter drift in daylight, and are thus able to remain on stone tops both during the day and at night.