Water intake by the adult mayflyEpeorus ikanosis (Ephemeroptera: Heptageniidae) and its effect on their longevity

Water drinking habits during flight in adult mayflies ofEpeorus ikanonis Takahashi were observed and the effect of water intake on their longevity was examined. The study was carried out in a mountain stream in western Japan. Adult males collected a water droplet under the head capsule while alighting on the water surface and consumed it after moving to nearby riparian vegetation. The amount of water intake was experimentally estimated to be 9.7% of the bodyweight for males. Although females did not show the water drinking behavior in the field, they imbibed as much as 5.5% of the bodyweight of water in an experimental situation. Morphological observation of the mouthparts revealed that the water was drawn in through the pits at the base of the labrum. In the field caging experiment, males with a water supply had a substantially longer life span than those without. Increase in adult life span by drinking water was less marked in females. Multiple copulation was observed in both sexes during the experiment. The water drinking behavior of the males may closely relate to mating success through increased longevity. Variation in the longevity of adult mayflies was discussed with regard to the possibility of water intake. This paper forms part of a PhD thesis submitted to the Department of Zoology in Kyoto University (1990).     

Production of Hexagenia limbata nymphs in contaminated sediments in the Upper Great Lakes Connecting Channels

In April through October 1986, we sampled sediments and populations of nymphs of the burrowing mayfly, Hexagenia limbata (Serville), at 11 locations throughout the connecting channels of the upper Great Lakes, to determine if sediment contaminants adversely affected nymph production. Production over this period was high (980 to 9231 mg dry wt m^-2) at the five locations where measured sediment levels of oil, cyanide, and six metals were below the threshold criteria of the U.S. Environmental Protection Agency and the Ontario Ministry of Environment for contaminated or polluted sediments, and also where the criterion for visible oil given in the Water Quality Agreement between the U.S.A. and Canada for connecting waters of the Great Lakes was not exceeded. At the other six locations where sediments were polluted, production was markedly lower (359 to 872 mg dry wt m^-2). This finding is significant because it indicates that existing sediment quality criteria can be applied to protect H. limbata from oil, cyanide, and metals in the Great Lakes and connecting channels where the species fulfills a major role in secondary production and trophic transfer of energy.Contribution 733, of the National Fisheries Research Center-Great Lakes, U.S. Fish and Wildlife Service, 1451 Green Road, Ann Arbor, MI 48105.     

Paravelia anta sp. n. and P. hungerfordi (Drake and Harris) from South America (Heteroptera: Veliidae)

Paravelia anta sp.n. from Argentina and Paraguay is described and compared with its close relative P. hungerfordi (Drake & Harris) from Brasil, which is redescribed. In addition, characteristics to separate these two from the remaining species of Paravelia are given.     

Predator density and dissolved oxygen affect body condition of <Emphasis Type="Italic">Stenonema tripunctatum</Emphasis> (Ephemeroptera,Heptageniidae) from intermittent streams

The effects of population density, fish density, and dissolved oxygen on body condition of late-instar nymphs of Stenonema tripunctatum (Ephemeroptera, Heptageniidae) were investigated using nymphs sampled from isolated, upland stream pools over summer in central Arkansas, USA. All three factors exhibited high variation among pools. Body condition was negatively related to fish density, and positively related to dissolved oxygen (when included in the model). High fish densities may be related to low body condition because they cause reduced foraging or force earlier emergence at small body sizes. These results emphasize the combined effects of biotic and abiotic factors on body condition in mayflies, and support earlier findings that population density is a less-important factor.     

胜利河连续系统中蜉蝣优势种的生产量动态和营养基础

大型底栖动物在河流生态系统中发挥着重要作用, 2009 年3月至2010 年3月间对长江中游支流巴河流域的胜利河大型底栖动物群落优势种类的生产力进行为期1 周年的调查研究, 结果表明, 主要蜉蝣优势种扁蜉、等蜉和红斑蜉的生活史为3代/年、3代/年和 2代/年。现存量呈现出1-3级河流增加,而4级又较3级有所下降的趋势。采用龄期频率法( instar-frequency method) 测算的平均周年生产量分别为扁蜉, 200.13 g/(m².a) , P/B 为23.69; 等蜉, 82.06 g/(m².a), P/B 为18.12;红斑蜉, 12.30 g/(m².a), P/B 为8.78。三种蜉蝣的生产量动态在时间上与现存量动态较一致,但彼此各不相同。扁蜉的日均产量于2009年3月在二级河流中达到最大(363.56 mg/m².d),等蜉于2010年3月在三级河流中到达最大(282.76 mg/m².d),而红斑蜉于2009年3月在一级河流中到达最大(33.36 mg/m².d)。生产量的营养基础分析结果表明,扁蜉前肠内含物中无形态碎屑、动物组织、植物纤维、丝状藻类、硅藻所占平均比例为74.37%,4.19%,17.11%,4.29%,0.04%,对生产量的贡献率分别为77.15%,11.27%,6.57%,4.95% ,0.04%;等蜉前肠内含物中无形态碎屑、动物组织、植物纤维、真菌、丝状藻类和硅藻所占平均比例为65.64%,6.17%,23.04%,0.54%,4.53%,对生产量的贡献率分别为: 68.16%,16.61%,8.86%,1.03%,5.23%,0.09%,0.10%;红斑蜉前肠内含物中,无形态碎屑、动物组织、植物纤维、真菌、丝状藻类和硅藻所占平均比例为41.14%,5.96%,38.04%,1.34%,11.21%,2.31%,对生产量的贡献率分别为46.67%,17.52%,15.98%,2.81%,14.13%,2.91%。这与我们在黑竹冲和叹气沟的研究结果存在一定差异,可能与这些溪流自身环境和地区分布有关。     

Growth responses of littoral mayflies to the phosphorus content of their food

We examined how mayfly growth rates and body stoichiometry respond to changing phosphorus (P) content in food. In two experiments, mayfly nymphs were given high or low quantities of food at different carbon:phosphorus (C:P) ratios and their growth was measured. Low food quantity resulted in negative growth rates in both experiments, regardless of food P content. However, under high food availability, mayfly growth was affected by the type of food eaten, with low C:P ratio food producing more rapid growth. In addition, mayfly growth increased somewhat when P-poor food was artificially enriched with inorganic P although this effect was not statistically significant. Mayfly body P content was inversely related to body size but increased in animals fed artificially P-enriched food. A model was constructed to simulate mass balance constraints on mayfly growth imposed by the relative supply of two elements (C and P) in food. The model shows that mayfly growth should be limited by food P content at moderately low C:P ratios ( c . 120, by mass). Given high C:P ratios (mean c.  270, by mass) in periphyton from oligotrophic boreal lakes, our experimental and theoretical results indicate that stoichiometric constraints are important factors affecting benthic food webs in lakes from the Canadian Shield and perhaps in other systems with similarly high C:P ratios in periphyton.     

Predation on mayfly nymph, Baetis rhodani, by native and introduced Gammarus: direct effects and the facilitation of predation by salmonids

1. In a series of laboratory experiments, we assessed the predatory nature of the native Irish amphipod, Gammarus duebeni celticus , and the introduced G. pulex , towards the mayfly nymph Baetis rhodani . We also investigated alterations in microhabitat use and drift behaviour of B . rhodani in the presence of Gammarus , and indirect predatory interactions with juvenile Atlantic salmon, Salmo salar .
2. In trials with single predators and prey, B. rhodani survival was significantly lower when Gammarus were free to interact with nymphs as than when Gammarus were isolated from them. The invader G. pulex reduced the survival of B. rhodani more rapidly than did the native G. d. celticus . Both Gammarus spp. were active predators.
3. In `patch' experiments, B. rhodani survival was significantly lower both when G. pulex and G. d. celticus were present, although the effect of the two Gammarus species did not differ. Again, active predation of nymphs by Gammarus was observed. Significantly more nymphs occurred on the top and sides of a tile, and per capita drifts were significantly higher, when Gammarus were present. Baetis rhodani per capita drift was also significantly higher in the presence of the introduced G. pulex than with the native G. d. celticus .
4.  Gammarus facilitated predation by salmon parr of B. rhodani by significantly increasing fish–nymph encounters on exposed gravel and in the drift. There were no differential effects of the two Gammarus spp. on fish – B . rhodani encounters or consumption.
5. We conclude that Gammarus as a predator can have lethal, nonlethal, direct and indirect effects in freshwaters. We stress the need for recognition of this predatory role when assigning Gammarus spp. to a `Functional Feeding Group'.     

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.     

The role of altitude and associated habitat stability in determining patterns of population genetic structure in two species of Atalophlebia (Ephemeroptera: Leptophlebiidae)

1. Previous studies have identified lowland areas as barriers to gene flow (dispersal) between distinct mountain ranges in montane species of aquatic insects. In this study, we investigated the population genetic structure of two closely related Atalophlebia (mayfly) species inhabiting lowland areas of south‐east Queensland, Australia, with the expectation of widespread gene flow throughout the low‐altitude environment and associated homogeneous genetic structure. 2. In particular, we asked whether species with lower‐altitude distributions demonstrate greater spatial distribution of mtDNA (COI) alleles than the upland species studied previously. This pattern would be expected if good dispersal ability is associated with population persistence in these drought‐prone habitats. 3. The two species demonstrated contrasting genetic population structure. Atalophlebia sp. AV13 D revealed strong population structure, with populations on each side of the low‐altitude area isolated from each other for a long time (c.350 kya), and the presence of an isolation‐by‐distance pattern over relatively small geographical distances (<40 km). In contrast, Atalophlebia sp. AV13 A was panmictic at the scale investigated (≤160 km), with no history of past population fragmentation. 4. Examination of sample distribution along the altitudinal gradient reveals that Atalophlebia sp. AV13 D may have a more upland distribution (associated with greater habitat stability) than previously supposed, while Atalophlebia sp. AV13 A inhabits more xeric lowland areas, where freshwater habitats are less stable. We consequently hypothesise that these contrasting genetic population structures result from differences in habitat stability along the altitudinal gradient, only species with good dispersal ability being able to persist in unstable habitats. These findings may be applicable to other regions of the globe where habitat instability is associated with altitudinal gradients.     

Nocturnal drift of mayfly nymphs as a post-contact antipredator mechanism

1. The predominantly nocturnal constrained drift of stream invertebrates is commonly regarded as a behaviour that avoids encounters with visually foraging fish in the water column. The alternative explanation, that drift peaks are caused by bottom-feeding, nocturnal predators, has rarely been tested.
2. We examined these hypotheses by collecting invertebrate drift in five streams in northern Finland: one with brown trout ( Salmo trutta , a drift-feeding fish), one with alpine bullhead ( Cottus poecilopus , a benthic fish), one with both species, and two fishless streams.
3. Drift by Baetis mayflies was aperiodic or slightly diurnal in both fishless streams on all sampling occasions. In contrast, drift was nocturnal in streams with trout and, to a lesser extent, in the stream with bullhead. Non-dipteran prey drifted mainly nocturnally in all streams with fish, whereas Diptera larvae were less responsive to the presence of fish.
4. In laboratory experiments, bullheads were night-active, causing a much higher frequency of drift by touching Baetis at night than during the day. Thus, increased nocturnal drift may serve to avoid both visual predators (a pre-contact response) and benthic fish (a post-contact response). In streams with bottom-feeding fish, nocturnal drift should be caused by increased drift by night rather than by reduced drift by day.