Inland dispersal of adult aquatic insects

1. Adult caddisflies (Trichoptera) and mayflies (Ephemeroptera) were light-trapped on summer evenings along the Detroit River and Lake St. Clair, near Windsor, Ontario, Canada. Light traps were located at the shore and at increasing distances inland up to 5 km, and were operated simultaneously for 2 h following sunset. Catches of five species of caddisflies of the family Hydropsychidae ( Cheumatopsyche campyla , Cheumatopsyche speciosa , Hydropsyche hageni , Hydropsyche phalerata , Macrostemum zebratum ) and the mayfly Hexagenia (Ephemeridae) were used to examine inland distribution.
2. Inland dispersal was limited: catches of caddisflies declined at a greater than exponential rate with increasing distance from shore. Mean dispersal distance from the shoreline ranged from 650 to 1845 m. Smaller caddisfly species dispersed shorter distances than larger caddisflies and Hexagenia .
3. Inland distribution of adult caddisflies exhibited considerable interspecific variation: distribution was inconsistent among trials for Hexagenia , possibly owing to timing of collections in relation to periods of peak emergence.
4. Sex ratios of caddisflies were female biased at most sites. No consistent bias was observed for Hexagenia . Different inland distribution patterns were observed for males and females. The differences appeared to reflect species-specific reproductive strategies.     

Inland dispersal of adult aquatic insects

1. Adult caddisflies (Trichoptera) and mayflies (Ephemeroptera) were light-trapped on summer evenings along the Detroit River and Lake St. Clair, near Windsor, Ontario, Canada. Light traps were located at the shore and at increasing distances inland up to 5 km, and were operated simultaneously for 2 h following sunset. Catches of five species of caddisflies of the family Hydropsychidae ( Cheumatopsyche campyla , Cheumatopsyche speciosa , Hydropsyche hageni , Hydropsyche phalerata , Macrostemum zebratum ) and the mayfly Hexagenia (Ephemeridae) were used to examine inland distribution.
2. Inland dispersal was limited: catches of caddisflies declined at a greater than exponential rate with increasing distance from shore. Mean dispersal distance from the shoreline ranged from 650 to 1845 m. Smaller caddisfly species dispersed shorter distances than larger caddisflies and Hexagenia .
3. Inland distribution of adult caddisflies exhibited considerable interspecific variation: distribution was inconsistent among trials for Hexagenia , possibly owing to timing of collections in relation to periods of peak emergence.
4. Sex ratios of caddisflies were female biased at most sites. No consistent bias was observed for Hexagenia . Different inland distribution patterns were observed for males and females. The differences appeared to reflect species-specific reproductive strategies.     

Distribution and abundance of blackflies (Diptera: Simuliidae) in relation to impoundments in the Buffalo River, eastern Cape, South Africa

1. The abundance of blackfly species along the length (140 km) of the multiply impounded Buffalo River, eastern Cape, South Africa, was studied between 1986 and 1988. 2. Twenty-three species of blackflies were recorded, most of which (95%) were usually found 5–10 km from the source. 3. Impoundments favoured five lower-reach (pest) blackfly species at the expense of two upper-reach (non-pest) species. Impoundments have therefore shifted the distribution of blackfly species ‘upstream’, and have increased the distribution of pest blackflies. 4. The smallest impoundment, situated in the Upper Foothill Stony Run Zone, had the greatest effect on blackfly distribution. 5. There was no typical filter-feeding guild below impoundments. Possible reasons for this included the discharge of toxic algae from impoundments, the lack of compensatory releases, the sudden drying of impoundment tailwaters and the paucity of natural lakes in southern Africa which precludes a pre-adapted ‘outlet’ fauna.     

Omnivory by Planktivores Stabilizes Plankton Dynamics, but May Either Promote or Reduce Algal Biomass

Classical models of phytoplankton–zooplankton interaction show that with nutrient enrichment such systems may abruptly shift from limit cycles to stable phytoplankton domination due to zooplankton predation by planktivorous fish. Such models assume that planktivorous fish eat only zooplankton, but there are various species of filter-feeding fish that may also feed on phytoplankton. Here, we extend these classical models to systematically explore the effects of omnivory by planktivorous fish. Our analysis indicates that if fish forage on phytoplankton in addition to zooplankton, the alternative attractors predicted by the classical models disappear for all realistic parameter settings, even if omnivorous fish have a strong preference for zooplankton. Our model also shows that the level of fish biomass above which zooplankton collapse should be higher when fish are omnivorous than when fish are zooplanktivorous. We also used the model to explore the potential effects of the now increasingly common practice of stocking lakes with filter-feeding fish to control cyanobacteria. Because omnivorous filter-feeding fish forage on phytoplankton as well as on the main grazers of phytoplankton, the net effect of such fish on the phytoplankton biomass is not obvious. Our model suggests that there may be a unimodal relationship between the biomass of omnivorous filter-feeding fish and the biomass of phytoplankton. This implies that to manage for reductions in phytoplankton biomass, heavy stocking or strong reduction of such fish is best.     

Barriers to the recovery of aquatic insect communities in urban streams

1. Worldwide, increasing numbers of stream restoration projects are being initiated to rehabilitate waterways modified by urbanisation. However, many of these projects have limited success in restoring stream communities. Prompted by this, we investigated previously unrecognised barriers to aquatic insect colonisation in urban streams. 2. To investigate whether the availability of suitable substrata for oviposition limited the longitudinal distribution of caddisflies, large boulders were added to the upstream reaches of one stream. Prior to the addition, more egg masses were observed downstream and this longitudinal pattern persisted subsequently. 3. Malaise trapping revealed that adult caddisfly diversity and abundance was greater downstream than upstream. Furthermore, in a previous study the authors found larval caddisflies reflected the longitudinal distribution of adults. 4. The only obvious potential obstructions between reaches were roads beneath which the stream flowed through culverted crossings. Malaise trapping was used to examine the effect of road culverts and bridges on caddisfly dispersal. Numbers of caddisflies caught declined upstream and about 2.5 × more individuals were taken in traps immediately below than above five culverts. 4. Bridges, which had a more open structure than culverts, had no significant effect on the size of catches made above and below them. 5. Road culverts could act as partial barriers to upstream flight, with consequences for larval recruitment in urban streams. We recommend that urban planners and designers of restoration projects consider possible synergistic effects of poor oviposition habitat and barriers to aquatic insect dispersal, which may be critical for the colonisation of urban headwater streams and for the maintenance of stream insect populations.     

背角无齿蚌(Anodonta woodiana)对浅水系统底栖和浮游藻类竞争关系的影响

底栖藻类和浮游藻类之间的竞争关系对浅水生态系统的结构、功能具有重要的影响,双壳类可通过滤食控制浮游藻类,从而改变底栖藻类与浮游藻类之间的竞争结果。论文通过比较放养背角无齿蚌(Anodonta woodiana)(蚌处理组)与不放养背角无齿蚌(对照组)系统中底栖藻类、浮游藻类的生物量和优势种等的变化,研究了滤食性双壳类对底栖藻类和浮游藻类间竞争的影响。结果表明,背角无齿蚌可显著降低浮游藻类生物量,提高水体透明度和沉积物表面光照条件,从而显著提高底栖藻类的生物量;背角无齿蚌也改变了浮游藻类的优势种,使优势种由蓝藻转变成硅藻。因此,滤食性双壳类有利于促进浅水生态系统从混水态向清水态转变,本研究结果对富营养化浅水湖泊修复与管理具有一定的参考意义。     

Long-term changes in the benthic communities of the Pomeranian Bay (Southern Baltic Sea)

Long-term changes in the macrofauna of the Pomeranian Bay were studied by comparing survey data from the 1950s, 1980s, and 1990s. The study area has undergone significant eutrophication during the period of investigation. Biomass of filter-feeding bivalves increased significantly. Spatial distribution patterns of several species have changed. Strong decreases in species richness were caused by oxygen depletion at stations deeper than 15 m.Saduria entomon, Monoporeia affinis, andPontoporeia femorata vanished entirely between 1981 and 1993. Although a causal relationship between simultaneous increases of nutrient levels and macrobenthic biomass cannot be verified, eutrophication is proposed to be the major process affecing changes in macrofauna assemblages. In addition, changes in hydrography and climate increased frequency and severity of oxygen depletion events in the Pomeranian Bay since the mid 1980s.     

Influence of a collector-filterer caddisfly (Trichoptera :Hydropsyche betteni Ross) on leaf processing rates in an artificial stream

A 2 × 6 unbalanced fixed effects factorial treatment design was used to examine the effects of a collector-filterer on leaf processing rates of American Cottonwood (Populus deltoides Bartr.) in an artificial stream over a six week period. Pre-leached leaves were placed into 30 small enclosures; collector-filterer caddisflies (Hydropsyche betteni Ross) were added to 18 enclosures resulting in two treatment factors (leaves with and without caddisflies). Changes in leaf dry-weight and caddisfly biomass were determined at weekly sampling intervals. Leaf processing rates were significantly different between treatments over time, with enclosures without caddisflies exhibiting a greater weight loss (higher k value) than enclosures with caddisflies (p < 0.01). The reduction in processing rates of the leaves due to the presence of caddisflies was attributed to the construction of retreats by these collector-filterers. Under the influence of the caddisfly, leaf material may be able to remain as a viable food resource for a longer period in the stream.     

Amphibiotic insects of mountain lakes and small watercourses in the Urals

A total of 127 species of stoneflies, dayflies, caddisflies, and chironomids were found in the lakes and small watercourses of the Northern and Pre-Polar Urals. The highest species diversity (78 species) and number were the characteristics of chironomids. Along with widely distributed species, Siberian and Arctic species were revealed in the fauna. The Arctic stonefly Capnia zaicevi Klap and 24 chironomid species were found for the first time on the territory of Komi Republic. Lists of registered insect species and data on their occurrence, abundance, and biomass are given.     

Experimental study of trophic cascade effect of silver carp (Hypophthalmichthys molitrixon) in a subtropical lake, Lake Donghu: on plankton community and underlying mechanisms of changes of crustacean community

An enclosure experiment was carried out to test trophic cascade effect of filter-feeding fish on the ecosystem: growth of crustacean zooplankton, and possible mechanism of changes of crustacean community structure. Four fish biomass levels were set as follows: 0, 116, 176 and 316 g m^-2, and lake water (containing ca. 190 g m^-2 of filter-feeding fishes) was comparatively monitored. Nutrient levels were high in all treatments during the experiment. Lowest algal biomass were measured in fishless treatment. Algal biomass decreased during days 21–56 as a function of fish biomass in treatments of low (LF), medium (MF) and high (HF) fish biomass. Crustaceans biomass decreased with increasing fish biomass. Small-bodied cladocerans, Moina micrura, Diaphanosoma brachyurum and Scapholeberis kingii survived when fish biomass was high whilst, large-bodied cladocerans Daphnia spp. and the cyclopoids Theromcyclops taihokuensis, T. brevifuratus, Mescyclops notius and Cyclops vicinus were abundant only in NF enclosures. Evasive calanoid Sinodiaptomus sarsi was significantly enhanced in LF, but decreased significantly with further increase of fish biomass. Demographic data indicated that M. micrura was well developed in all treatments. Our study indicates that algal biomass might be controlled by silver carp biomass in eutrophic environment. Changes of crustacean community are probably affected by the age of the first generation of species. Species with short generation time were dominant and species with long generation time survived less with high fish biomass. Evasive calanoids hardly developed in treatments with high fish biomass because of the (bottle neck) effect of nauplii. Species abundance were positively related to fish predation avoidance. Other than direct predation, zooplankton might also be suppressed by filter-feeding fish via competition.     

三峡水库初次蓄水后干流库区枝角类的空间分布与季节变化

为了阐明三峡水库初次蓄水后干流库区枝角类的时空分布规律, 2004年4月至2005年1月, 在600余公里干流库区设置10个样点, 定性和定量采集枝角类标本。本次调查共采到9种枝角类, 其中春季8种, 夏季5种, 秋季2种, 冬季3种。枝角类种类组成的空间分布差异明显, 河流状态的江段有4种, 水库状态的江段有7种, 水库状态的江段距大坝越近, 枝角类种类越丰富。枝角类的密度和生物量的空间分布和季节变化规律一致, 季节间差异极显著(P<0.01), 春季最高, 夏季次之, 秋季和冬季较低; 空间分布上表现为在干流库区的纵向分布上差异极显著(P<0.01), 河流状态样点远低于水库状态的样点, 水库上游江段又低于下游坝前段。结果显示三峡水库的枝角类具有明显的纵向分布和季节变化特征。     

Coexistence in a guild of filter feeding caddis larvae: do different instars act as different species?

Summary Guild structure and coexistence of filter feeding caddis larvae was studied in three lake-outlet streams in northern Finland. Natural stones were used as sampling units, and for each stone measurements of six microhabitat variables were taken. Species and species instar relationships in the multivariate microhabitat space were analyzed by a method based on Principal Components Analysis. Seven species of filtering caddis larvae were found at the study sites, but detailed analysis was restricted to the four most abundant ones (Polycentropus flavomaculatus, Hydropsyche angustipennis, H. pellucidula, H. saxonica). P. flavomaculatus was the microhabitat generalist of the guild, whereas all the hydropsychids were narrowly specialized in at least one larval instar. In all species, different instars utilized the microhabitat somewhat differentially. Moss content and Froude number were significantly different between instar microhabitats in all comparisons. Ontogenetic niche changes not only separated species instars from each other, but also profoundly changed the biotic as well as abiotic environment of the species at different phases of larval development. This may have important effects on the dynamics of interacting species and, consequently, on the guild structure. While some of the more classical models of coexistence may also be relevant for understanding coexistence in this guild of sessile stream insects, ontogenetic niche changes clearly need to be taken into consideration in future studies of guild structure and species interactions of filter feeding caddisfly larvae.     

Feeding habits and food resources of filter-feeding Trichoptera in a regulated mountain stream

A year-round study was conducted to examine feeding habits and food resources of the filter-feeding Trichoptera Arctopsyche grandis and Brachycentrus occidentalis along a regulated mountain stream gradient. There was a well defined longitudinal species replacement with A. grandis reaching maximum densities 2.3 kilometers below the impoundment, and concomitant with its decline downstream was an increase in B. occidentalis. At all sampling sites the < 75 µm organic seston fraction usually consisted primarily of diatoms (>70%, by areal estimate on microscope slides), whereas the 75–250 µm and > 250 µm seston fractions were predominantly composed of detritus (> 80 %). B. occidentalis larvae consumed primarily detritus and diatoms (> 70 % of the diet), while A. grandis ingested a variety of materials with animals, detritus and/or filamentous algae often constituting > 80% of the diet. Animal material was over-represented in the diets of both species when compared with amounts in the seston. Feeding habits provided partial explanations for the distinct longitudinal distribution patterns of filter-feeding Trichoptera observed in the regulated river.     

Terrestrial–aquatic transitions: Local abundances and movements of mature female caddisflies are related to oviposition habits but not flight capability

1. Movement behaviours of adult aquatic insects can produce distinct spatial distribution patterns. Studies of adult abundance with distance away from water bodies are common and may invoke flight capability to explain species differences. In contrast, distribution patterns along river channels are poorly described, but are no less important for understanding population dynamics. Longitudinal patterns in adult abundance along short river lengths may differ between sexes and at different life stage transitions between aquatic and terrestrial environments, i.e. at emergence and oviposition. Flight capability is unlikely to influence longitudinal patterns created at emergence, but may influence local abundances of mature females seeking to lay eggs. We tested hypotheses about how local abundances of mature females might differ according to oviposition habits and flight capability.
2. We surveyed abundances of mature female caddisflies at adjacent riffle–pool pairs along short river lengths with homogeneous riparian cover. Our survey included nine species in three families (Hydrobiosidae, Leptoceridae, Hydropsychidae), which encompassed multiple different oviposition habits and a range of wing sizes and shapes. Several of the species oviposit preferentially in riffles. Accordingly, we tested for differences in female abundance between channel units (adjacent riffle–pool pairs). We also tested whether females attained higher abundances in some places along channels than others (i.e. over larger spatial scales and regardless of channel unit) which imply movements along the channel and aggregation in some locations. Wing morphology was used as a proxy measure of flight capability and included measures of wing span, area, aspect ratio and the second moment of wing area.
3. Three distinctly different distribution patterns of mature female caddisflies were identified. The abundance of three species varied over larger scales only (multiple channel units). Six species that oviposit preferentially in riffles had higher female abundances at riffles than pools, but for only one did abundances also vary over larger scales. There was no association between these different patterns and measures of wing morphology, after removing metrics that were correlated and that differed systematically between taxonomic families. However, we could not reject the hypothesis that some aspect of flight behaviour may have contributed to observed patterns.
4. The diverse but distinct distributions of mature female caddisflies we observed along short channel lengths are novel and suggest that species differ in their propensity for movement along streams, which could have consequences for local densities of eggs and juveniles in the aquatic environment. The degree to which population sizes are coupled across the terrestrial-to-aquatic transition is rarely investigated in aquatic insects and may provide fresh insight into sources of spatial variation within populations. Similarly, a more nuanced approach to research on the flight of aquatic insects, including age- and sex-specific phenomena, may provide greater insight into the diverse ecological functions and consequences of movement.

     

Longitudinal river zonation in the tropics: examples of fish and caddisflies from the endorheic Awash River,Ethiopia

Specific concepts of fluvial ecology are well studied in riverine ecosystems of the temperate zone but poorly investigated in the Afrotropical region. Hence, we examined the longitudinal zonation of fish and adult caddisfly (Trichoptera) assemblages in the endorheic Awash River (1,250 km in length), Ethiopia. We expected that species assemblages are structured along environmental gradients, reflecting the pattern of large-scale freshwater ecoregions. We applied multivariate statistical methods to test for differences in spatial species assemblage structure and identified characteristic taxa of the observed biocoenoses by indicator species analyses. Fish and caddisfly assemblages were clustered into highland and lowland communities, following the freshwater ecoregions, but separated by an ecotone with highest biodiversity. Moreover, the caddisfly results suggest separating the heterogeneous highlands into a forested and a deforested zone. Surprisingly, the Awash drainage is rather species-poor: only 11 fish (1 endemic, 2 introduced) and 28 caddisfly species (8 new records for Ethiopia) were recorded from the mainstem and its major tributaries. Nevertheless, specialized species characterize the highland forests, whereas the lowlands primarily host geographically widely distributed species. This study showed that a combined approach of fish and caddisflies is a suitable method for assessing regional characteristics of fluvial ecosystems in the tropics.

     

Nutrient Subsidies from Iteroparous Fish Migrations Can Enhance Stream Productivity

Migratory animals often transfer nutrients between ecosystems, enhancing productivity in the subsidized system. Most research on nutrient subsidies by migratory fishes has focused on Pacific salmon, whose semelparous life history is unusual among migratory fishes. To test whether iteroparous species can provide ecologically important nutrient inputs to stream ecosystems, we experimentally blocked the migration of suckers (Catostomidae) midway up an oligotrophic tributary of Lake Michigan. Comparing reaches upstream of the barrier to downstream reaches containing thousands of breeding fish, we found that suckers elevated phosphorus and nitrogen concentrations three- to five-fold. Algal accrual was doubled and caddisflies grew 12% larger in subsidized reaches relative to reference reaches. An enclosure experiment demonstrated that caddisflies with access to a fish carcass rapidly became enriched in ¹⁵N and ¹³C, and experimental carcass additions were rapidly colonized by high densities of caddisflies. However, under natural conditions below the experimental barrier, caddisflies became enriched in ¹⁵N but not ¹³C, indicating that fish-derived nutrients entered the stream food web primarily through indirect pathways rather than direct consumption of carcasses or gametes. At pupation, an average of 18% of caddisfly tissue N below the barrier was sucker-derived. In comparison to our focal stream, a reference stream with few suckers showed no seasonal or longitudinal patterns in nutrients and stable isotopes. These results demonstrate that iteroparous fish migrations can spur productivity via nutrient subsidies, despite low mortality rates. Thus, concerns about negative ecosystem-level consequences of blocking migrations of semelparous fishes are also applicable to iteroparous species when migrations are large.     

Tufa Barriers from a Caddisfly's Point of View: Streams or Lake Outlets?

The caddisfly community composition in different microhabitats at tufa barriers was studied in the Plitvice Lakes NP, Croatia. Three tufa barriers were investigated and six emergence traps were installed at each site covering various microhabitats. Sampling was conducted monthly during the year 2008. Tufa barriers are lake outlet habitats, but they feature a variety of microhabitats similar to streams, which is reflected in the composition of caddisfly communities regarding longitudinal distribution preferences. The caddisfly communities at all three sites were dominated by species typical for the rhithral zone, but there was a shift in species composition along a longitudinal gradient, from the epirhithral to the metarhithral zone. Analysis of functional feeding guilds showed considerable differences between the caddisfly community at the Labudovac barrier and both downstream barriers, shifting from one with a quite diverse composition, to one completely dominated by passive filter‐feeders. Passive filter feeders were not represented by the same taxa at up‐ and downstream barriers (i.e., by Hydropsyche species and Wormaldia species, at the Labudovac barrier and at both downstream barriers, respectively). Due to high complexity and habitat diversity, the highest diversity and equitability of caddisfly communities were recorded at microhabitats with particulate tufa and medium current velocity (10–20 cm/s). The lowest diversity and species richness were recorded for silt with low current velocity (0–10 cm/s). Abundance of caddisflies was positively correlated with current velocity due to a very high proportion of rheophilic passive filter feeders in the communities. However, community composition and structure is only to some degree influenced by microhabitat characteristics, but rather by their general position within the barrage‐lake system, i.e., longitudinal distribution and thereby availability of different food resources. (© 2012 WILEY‐VCH Verlag GmbH & Co. KGaA, Weinheim)     

Zooplankton biomass in tropical reservoirs in southern Brazil

In order to test the hypothesis that zooplankton biomass distribution (total and taxonomic groups) was influenced by the nutrient concentration and primary productivity distribution in three tropical reservoirs, subsurface samples were taken in the fluvial, transitional and lacustrine regions of three reservoirs (oligotrophic, mesotrophic and eutrophic) in southern Brazil (Paraná State) in March and September 2002. Zooplankton biomass ranged from 0.04 to 264.47 mg DW m⁻³. Higher biomass values were observed for cladocerans (73.60%; 0.01–259.86 mg DW m⁻³), followed by copepods (22.05%; 0.01–69.69 mg DW m⁻³) and rotifers (4.35%; 0.01–11.52 mg DW m⁻³). In general, the total zooplankton, rotifer, cladoceran and copepod biomass, and chlorophyll-a and total nutrient concentrations showed a similar longitudinal distribution within the reservoirs. Total zooplankton, rotifer and cladoceran biomass were related to the chlorophyll-a concentration, and zooplankton biomass was related to the total phosphorus distribution. This may have been due to the significant multicolinearity between the chlorophyll-a and total phosphorus concentrations. Cyanobacteria influenced the taxonomic group biomass results by interfering with the filter feeding in larger zooplankton species, which favoured the dominance of smaller species. As regards the longitudinal distribution of copepod biomass, cyanobacteria biomass determined the displacement of the microcrustaceans to the fluvial region of Iraí Reservoir. Our results supported the hypothesis formulated and the primary productivity was the major predictor of the zooplankton biomass distribution in the reservoirs. Handling editor: S. Dodson     

Vegetation Cover and Substrate Type as Factors Influencing the Spatial Distribution of Trichopterans along a Karstic River

In the longitudinal continuum of the Kupa River the vegetation cover and substrate type were the important environmental factors influencing the spatial differences in the biomass and community composition. Of total macroinvertebrate biomass, a significantly greater percentage of trichopterans was found on boulder and cobble substrata covered with moss (54.3% on boulders, 55.8% on cobbles) than on substrata covered with periphyton (9.9% on boulders, 14.8% on cobbles). In the potamal, trichopterans were markedly reduced (<2.5% of total macroinvertebrate biomass) on gravel substrata. A comparison of the Shannon diversity index values suggested that for trichopteran species diversity the substrate type was a more influential factor than vegetation cover. On the other hand, multidimensional scaling analysis showed that trichopteran community composition was related more significantly to vegetation cover and river area than to substrate type. In the rhithral the vegetation cover was an important factor influencing the functional feeding group composition of trichopterans. The spatial distribution of scrapers and filtering collectors depended significantly on the vegetation cover associated with substrate type, and shredder trichopterans were related to vegetation cover only. Predatory trichopterans made up 17–65% of total predator biomass, and in the rhithron area they were correlated significantly only with vegetation cover. On gravel substrata in the potamal, vegetation cover did not affect the spatial distribution of shredder and collector‐filterer trichopterans significantly.     

Spatial Distribution of the Phytoplankton in a Tropical Lake (Lake Lanao,Philippines)

The distribution of phytoplankton was studied in Lake Lanao, Philippines. In calm weather, the species segregate vertically with motile species and buoyancy-regulating species above other species in the water column. On a distance scale of 12.5 km, horizontal variability yields a coefficient of variation of 23.6% for total biomass, 30.2% for net primary production, and an average of 30.3% for individual species biomass after removal of error variances. Variabilities on a scale of 1 km are about half as large. Horizontal patchiness of individual species is not related to their abundance, surface to volume ratio, morphotype, or taxon, but is negatively related to their size.