Scale-dependent habitat use by a large free-ranging predator,the Mediterranean fin whale

Since the heterogeneity of oceanographic conditions drives abundance, distribution, and availability of prey, it is essential to understand how foraging predators interact with their dynamic environment at various spatial and temporal scales. We examined the spatio-temporal relationships between oceanographic features and abundance of fin whales (Balaenoptera physalus), the largest free-ranging predator in the Western Mediterranean Sea (WM), through two independent approaches. First, spatial modeling was used to estimate whale density, using waiting distance (the distance between detections) for fin whales along ferry routes across the WM, in relation to remotely sensed oceanographic parameters. At a large scale (basin and year), fin whales exhibited fidelity to the northern WM with a summer-aggregated and winter-dispersed pattern. At mesoscale (20–100 km), whales were found in colder, saltier (from an on-board system) and dynamic areas defined by steep altimetric and temperature gradients. Second, using an independent fin whale satellite tracking dataset, we showed that tracked whales were effectively preferentially located in favorable habitats, i.e. in areas of high predicted densities as identified by our previous model using oceanographic data contemporaneous to the tracking period. We suggest that the large-scale fidelity corresponds to temporally and spatially predictable habitat of whale favorite prey, the northern krill (Meganyctiphanes norvegica), while mesoscale relationships are likely to identify areas of high prey concentration and availability.     

Does small-scale vertical distribution of juvenile schooling fish affect prey availability to surface-feeding seabirds in the Wadden Sea?

Food availability is a key variable influencing breeding performance and demography of marine top predators. Due to methodological problems, proportionality between fish abundance and availability is often assumed without being explicitly tested. More specifically, better breeding performance of surface-feeding seabirds at times of large prey stocks suggests that prey availability is also a function of prey abundance. Using vertically resolved stow net sampling we tested whether local abundance and length composition of pelagic fish are reliable predictors of the availability of these fish to surface-feeding Common Terns (Sterna hirundo) breeding in the German Wadden Sea. Prey fish were found to concentrate below the maximum diving depth of the terns. Individuals caught close to the surface were in most cases smaller than conspecifics caught at greater depth. Correlations between fish abundance within and out of reach of the terns appeared to be both species- and site-specific rather than driven by overall fish abundance. Vertical distribution patterns of the terns' main prey fish could be explained as anti-predator behavior, reducing prey availability to the terns. In 2007, when breeding performance was much better than in 2006, herring and whiting were much more abundant, suggesting that overall prey abundance may also increase prey availability in habitats other than those represented by the stow net sampling.     

Comparative dietary analysis of a recently described fish species complex (Galaxiidae) in a New Zealand river

The diets of several populations of four recently described species of galaxiid with allopatric distributions were examined in the Taieri River, New Zealand. Comparisons of available prey items and prey consumed were carried out to examine whether the distributions of the galaxiid species might be influenced by food availability and preferences. Seventy‐three prey categories were identified. The availability of four of these (Deleatidium, Oligochaeta, Elmidae larvae and adults) differed among the streams occupied by different galaxiid species. Three differences in the frequency of prey consumption (Deleatidium, Oxyethira, and Elmidae larvae) were detected. Despite differences in diet it was concluded that all four fish species were generalist predators of invertebrates with similar diets that were influenced mainly by prey availability. Furthermore prey availability was not considered responsible for the allopatric distribution of the four species, rather, spawning habitat availability, physical nature of the streams, and prior geological history of the Taieri catchment are considered the controlling influences on galaxiid distribution.     

Vertebral abnormalities in free-living Chinook salmon (Oncorhynchus tshawytscha,Walbaum) in New Zealand

Spinal abnormalities are common in farmed Chinook Salmon in New Zealand. We report spinal abnormalities in adult Chinook salmon which were predominantly hatchery reared and released as smolts and which we term free-living. We compare these to rates seen in farmed New Zealand salmon. 101 free-living adult salmon were radiographically assessed for spinal curvatures (lordosis, kyphosis, scoliosis; LKS) and other vertebral abnormalities. Severity of abnormality was assessed on a three-point scale. Abnormal vertebral bodies were detected in 88.1% of free-living salmon. Spinal curvatures were the most common abnormality type with 83.2% of fish showing this abnormality but only one free-living Chinook had LKS of severity greater than 1. Farmed Chinook salmon are reported to have LKS rates of 29% with 18% of LKS abnormalities of severity greater than 1. These results suggest that free-living Chinook salmon frequently develop spinal abnormalities, but these abnormalities are less severe than those observed in farmed salmon.     

Variability in the nutritional value of the major copepods in Cape Cod Bay (Massachusetts, USA) with implications for right whales

The North Atlantic right whale, a seriously endangered species, is found in Cape Cod Bay (Massachusetts, USA) during the winter and early spring. During their residency in these waters, these whales are frequently observed feeding. This study evaluated spatial and temporal changes in the chemical composition (carbon weight and C/N ratio) of the food resource targeted by the right whales in Cape Cod Bay. The three taxa measured (Centropages typicus, Pseudocalanus spp., and Calanus finmarchicus) had highly variable chemical compositions resulting from the different life strategies and from fluctuations in their surrounding environment. The impact of seasonal variability in the energy densities of the food resource of right whales was calculated and compared to the energetic requirements of these whales. Calculations indicated that differences in the nutritional content of the zooplankton prey in Cape Cod Bay could have a considerable effect on the nutrition available to the right whales. Therefore, it is likely that using more precise estimates of the energetic densities of the prey of right whales would lead to a re‐evaluation of the adequacy of the food resource available to these whales in the North Atlantic.     

Factors Affecting Long-term Trends in the Estuarine Abundance of Pogge (Agonus cataphractus )

Long-term data assembled from a power station on the Thames estuary were used to model fluctuations in the abundance of pogge (Agonus cataphractus) as a function of estuarine environmental, seasonal and prey availability variables using multiple regression. Temperature was the most important determinant of abundance, with seasonal variables and changes in the relative abundance of age-0 dab, Limanda limanda, possibly as prey, also holding important consequences for pogge abundance. Pogge abundance was also moderately influenced by: salinity, flow, salinity-flow interactions, changes in the relative abundance of Dover sole, Solea solea, and a trend variable. Regular patterns of seasonal (spring/winter) occurrences were associated with the use of warmer estuarine waters to optimize growth and reproductive potential and/or correlated with inshore spawning migrations and the availability of prey species. Both mechanisms suggest opportunistic use of estuarine habitats by pogge. Available temperature-abundance relationships further suggest a well-defined estuarine thermal niche, with an optimum at 9·5 °C. There remains a requirement for improved information on the biology of pogge and factors governing the population dynamics of this important estuarine species.     

Forecasting the consequences of climate-driven shifts in human behavior on cetaceans

While climate change is expected to affect cetaceans primarily via loss of habitat and changes in prey availability, additional consequences may result from climate-driven shifts in human behaviors and economic activities. For example, increases in shipping, oil and gas exploration and fishing due to the loss of Arctic sea ice are highly likely to exacerbate acoustic disturbance, ship strikes, bycatch and prey depletion for Arctic cetaceans. In the tropics, climate change may result in increased hunting pressure on near-shore dolphins and whales off Asia, Latin America, Africa, and elsewhere as the availability of other marine resources diminishes. This study explores the range of potential consequences to cetaceans worldwide from predicted climate-driven shifts in human behavior, and evaluates the risks to particular species given their geographic ranges and habitat preferences. While concern about impacts of climate change on cetaceans has largely focused on polar species, the analysis presented here suggests tropical coastal and riverine cetaceans such as the Irawaddy dolphin, Indo-Pacific humpback dolphin, and finless porpoise are particularly vulnerable to those aspects of climate change that are mediated by changes in human behavior. Policy recommendations include the following: (1) information about cetacean populations should be incorporated into national, regional and international climate adaptation decisions wherever possible (for example, via GEF-sponsored adaptation initiatives); and (2) human-mediated impacts of climate change should be included in cetacean conservation and management plans, such as the management procedures of the International Whaling Commission (IWC), where possible. Because human responses to climate change are likely to evolve rapidly over the coming years and decades, it is important that local, regional and international cetacean conservation and management plans include regular reviews to allow them to adapt to new information.     

Evidence that whales (Balaenoptera borealis) visit drifting fish aggregating devices: do their presence affect the processes underlying fish aggregation?

Evidence of the presence of a group of sei whales (Balaenoptera borealis) detected around drifting fish aggregating devices (FADs) was provided by omnidirectional multi‐beam sonar during a survey off the Seychelles (Indian Ocean). The short visit by the sei whales produced a significant change in the behaviour of the fish assemblage associated with the FAD. There was first a significant increase in fish density when the whales approached the FAD, then a marked decrease after the whales had moved away from the FAD. Furthermore, the fish density was still low, 3 h after the whales had left the FAD. We assume that the presence and behaviour of the sei whales led some of the fish initially associated with the FAD to move away from it. There has been a considerable increase in the use of drifting artificial FADs in the Indian Ocean in recent decades. The frequency of cetacean visits to drifting FADs in the Indian Ocean is unknown, but they may have a major impact on assemblages of pelagic fish species around FADs. The effect of marine mammals on FAD‐associated fish could be relevant to the ecological trap theory (FAD acting as a trap for their associated fish) because of their impact on the dynamics of fish aggregation processes, through commensalism and/or predator–prey interactions.     

Ontogenetic shifts and temporal changes in the trophic patterns of the deep‐sea red shrimp,Aristaeomorpha foliacea (Decapods: Aristeidae), in the Eastern Ionian Sea (Eastern Mediterranean)

The purpose of this paper is to provide the first detailed data concerning the diet and feeding activity of the giant red shrimp, Aristaeomorpha foliacea, in the Eastern Ionian Sea (Eastern Mediterranean), in relation to season, size class and sex. Feeding activity in A. foliacea was intense, based on its low vacuity index and high prey diversity, with a diet dominated by mesopelagic prey and less frequent occurrence of benthic taxa. Giant red shrimp displayed a highly diversified diet that exhibited slight seasonal fluctuations. The diets of both sexes consisted of 60 different prey categories belonging chiefly to three groups: crustaceans (e.g. decapods, such as Plesionika spp. and Pasiphaeidae, amphipods), cephalopods (mainly Enoploteuthidae) and fishes (Myctophidae, Macrouridae). These three prey categories accounted for 72–82% of the relative abundance and total occurrence for males and 70–88% for females, respectively. Variation in food availability, as well as increased energy demands related to gonad development and breeding activity, appear to be critical factors driving temporal changes in feeding strategy. Feeding activity increased during spring and summer, which coincides with reproductive activities (mating, gonad maturation, egg‐laying). Females seem to be more active predators than males, consuming prey with greater swimming ability. However, ontogenetic shifts in diet were also apparent, despite high dietary overlap among small, medium and large females. Large individuals, which are more efficient predators, selected highly mobile prey (e.g. fishes), whereas small individuals consumed low‐mobility prey (e.g. copepods, ostracods, tanaids and sipunculans).     

Impacts of neoliberal policies on non-market fishing economies on the Yukon River,Alaska

This paper describes how and why a relatively minor, mostly native non-market economy founded on subsistence-caught Chinook salmon in Alaska has become the object of controversy and increasing regulatory pressure. Small-scale exchanges of cash for subsistence-caught fish conflict with a neoliberal emphasis on markets, profit maximization, and private property. This paper clarifies the role of neoliberal policies in shaping and even causing the controversy over these exchanges, and the consequences for management of Chinook salmon on the Yukon River.     

A year in Barkley Canyon: A time-series observatory study of mid-slope benthos and habitat dynamics using the NEPTUNE Canada network

Understanding long-term trends in species abundance and distribution represents an important challenge for future research in the deep sea, particularly as management of human impacts becomes a more important concern. However, until natural higher frequency variability is better understood, it will be difficult to interpret any long-term trends that may be apparent in data sets. We present here the results of the first year of observations at the NEPTUNE Canada cabled observatory site in Barkley Canyon, off the coast of Vancouver Island, in the northeast Pacific Ocean. Presence/absence and abundance data for 28 faunal groups were extracted from daily video records from an observatory camera. Concurrent CTD and current meter data were collected from co-located instruments. Water mass properties, currents and faunal community composition exhibited notable seasonal trends. Distinct seasonal faunal groupings were observed, together with summer and winter trends in temperature, salinity and current patterns. Variations in abundance of decapod crustaceans and fishes were responsible for most differences between faunal groups. We suggest that faunal composition may have been responding to seasonal variations in food availability, together with direct and indirect physical influences on predator and prey abundance.     

Predominance of parallel‐ and cross‐predation in anglerfish

Several studies in the last 20 years have revealed that morphological asymmetry in fish can be characterized as ‘antisymmetry’. Antisymmetry is a lateral dimorphism in which each population consists of individuals with well‐developed left sides (lefties) and well‐developed right sides (righties). This dimorphism influences predator–prey interactions. In some piscivorous fishes, it has been found that predators can catch more prey of the opposite morphological type to themselves (cross‐predation) than of the same morphological type (parallel‐predation). Our previous work clarified that the predominance of cross‐predation is caused by lateralized behaviors of predators and prey that correspond to their morphological antisymmetry. Moreover, based on the results of our behavioral observations, we hypothesized that parallel‐predation can predominate when predators encounter the potential prey frontally. To test this hypothesis, in the present study we investigated the relationship between lateral morphological types of anglerfish (Lophiomus setigerus) and those of the prey fishes found in their stomachs. Anglerfish attract potential prey using their first dorsal fin (illicium) as a lure, and their frontal encounters with potential prey fishes were photographed in situ and observed in an aquarium. The results of a stomach contents analysis indicated that parallel‐predation predominated in five benthopelagic prey fish species (perches and eels). By contrast, five benthic prey fishes (gobies and weevers) exhibited the predominance of cross‐predation. These results not only demonstrate the predominance of parallel‐predation in a natural fish community, but also suggest that the relationship between morphological types of predator and prey species can be reversed depending on the lifestyle of prey.     

Ecological aspects of fin whale and humpback whale distribution during summer in the Norwegian Sea

The Norwegian Sea is a migration and feeding ground for fin whales (Balaenoptera physalus) and humpback whales (Megaptera novaeangliae) in summer. During the last decade, significant structural changes in the prey community, including northerly expansion and movement in the distribution of pelagic fish species, have been reported from this ecosystem. However, little information on whale feeding ecology exists in the Norwegian Sea and surrounding waters. A total of 59 fin whales and 48 humpback whales were sighted during 864 h of observation over an observation distance of about 8200 nmi (15,200 km) in the Norwegian Sea from 15 July to 6 August 2006 and 2007. The fin whale group size, as mean (±SD), varied between one and five individuals (2.1 ± 1.2 ind.) and humpback whale group size varied between one and six individuals (2.5 ± 1.7 ind.). Fin‐ and humpback whales were observed mainly in the northern part of the study area, and were only found correlated with the presence of macro‐zooplankton in cold Arctic water. Humpback whales were not correlated with the occurrence of adult Norwegian spring‐spawning herring (Clupea harengus) except for the northernmost areas. Despite changes in the whale prey communities in the Norwegian Sea, no apparent changes in fin‐ or humpback whale distribution pattern could be found in our study compared to their observed summer distribution 10–15 years ago.     

Site-specific effects on productivity of an upper trophic-level marine predator: Bottom-up, top-down, and mismatch effects on reproduction in a colonial seabird

We investigated the relative roles of bottom-up and top-down factors in limiting productivity of an upper trophic level marine predator. Our primary working hypothesis was that the reproductive success of black-legged kittiwakes (Rissa tridactyla) a piscivorous, colonial-nesting seabird, was most limited by the abundance, distribution, and species composition of surface-schooling forage fishes. A secondary working hypothesis was that reproductive loss to kittiwake nest predators was greatest during years of reduced prey availability. We report on a broad-scale, integrated study of kittiwakes and their prey in Prince William Sound, Alaska. Our study spanned five breeding seasons (1995–1999) and focused on three colonies that differed in size (ranging from ca. 220 to ca. 7000 breeding pairs) and proximity to each other (50–135 km apart). Kittiwakes in PWS encountered a variety of aquatic habitats, creating a complex foraging environment for breeding birds. We measured kittiwake reproductive success and foraging activities, while simultaneously measuring the abundance of surface schooling forage fishes throughout the foraging range of breeding kittiwakes. The abundance of primary prey species for kittiwakes (Pacific herring Clupea pallasi, Pacific sand lance Ammodytes hexapterus, and capelin Mallotus villosus) varied both annually and regionally, with no one region consistently having the greatest abundance of prey. Likewise, kittiwake reproductive success varied considerably among colonies and years.We found that bottom-up, top-down, timing mismatch, and colony-specific effects were all important to kittiwake productivity. Although bottom-up effects appeared to be strongest, they were not evident in some cases until other effects, such as geographic location (proximity of colony to prey concentrations) and top-down predation, were considered. Important bottom-up effects on kittiwake reproductive success were not only total prey abundance and distribution, but also species, age composition, and chronology of prey occurrence (match/mismatch of timing with critical brood-rearing periods); these effects varied by colony.Top-down effects of predation on kittiwake nest contents (independent of prey abundance) confounded seabird-forage fish relationships. Ultimately, when confounding factors were minimized, non-linear asymptotic relationships were identified between kittiwakes and their prey, with an asymptotic threshold of fish school surface area density of ca. 5 m²/km², beyond which top-down, physiological, or phylogenetic constraints likely restrict further reproductive output. The integrated approach of our investigations provided a more thorough understanding of the mechanisms underlying predator–prey relationships in the complex marine environment. However, such mechanistic theories can only be tested and refined through long-term research and monitoring of much greater duration than the 5-year study reported herein.     

Variations in crab claw morphology and diet across contrasting inter‐tidal habitats

Crabs are important predators of inter‐tidal ecosystems, controlling the abundance and distribution of their prey populations. Often the same crab species occupies several habitats and, although their effects on prey have been quantified across habitats, crabs’ dietary and morphological responses to differing environmental influences have been overlooked. Here, we used the crabs Eriphia verrucosa and Pachygrapsus marmoratus as model species to examine differences in claw morphometry – size and wear – and diet between rocky shore and heterogeneous sand flat habitats. We predicted that, intra‐specifically, crabs from rocky shores would consume more hard‐shelled prey owing to their high availability and consequently, would display chelipeds with the following claw characteristics: a higher degree of claw damage, stronger musculature (higher propel height) and increased mechanical advantage (defined as the ratio of input lever length to output lever length) than crabs in the heterogeneous sand flat habitats. Sampling was performed in heterogeneous sand flat habitats and rocky shores of the Central Portuguese coast. For each crab species, carapace width, diet composition and several claw morphometric measures were recorded, revealing significant intra‐specific differences (using multivariate analysis) between shore types. We found that E. verrucosa and P. marmoratus consumed more hard prey on rocky shore than on sand flat habitats, which resulted in rocky shore crabs having more accentuated dentition wear and larger musculature than their sand flat habitat counterparts. We suggest that the strong response of crab claw morphometry to environmentally induced diet variations is an important mechanism in the successful adaptation of crab species to inhabit differing habitats. A major implication is that the impact of the same species on prey may vary largely with habitat type as a result of predation efficiency varying with claw condition.     

Environmental influences on the long-term fluctuations in the abundance of gadoid species during estuarine residence

Samples taken regularly from the intake screens of West Thurrock power station between January 1977 and November 1992 were used to examine physico-chemical factors influencing the abundance of Gadidae species (poor cod, Trisopterus minutus; pouting, Trisopterus luscus; and whiting, Merlanguis merlangus) in the Thames estuary. Most sampled fish were age-0 and followed a dominant pattern of seasonal occurrence. Multiple regression was used to model variations in sample abundance in relation to fluctuations in estuarine environmental variables, interactions between environmental variables and seasonal factors. Model results were used to examine hypotheses concerning the relative importance of temperature, salinity, prey availability and seasonal factors as determinants of estuarine gadoid abundance. Temperature was the most important determinant of species sample abundance and negatively related to sample abundance in all cases. Salinity was a major determinant of pouting abundance and a minor determinant of whiting abundance, with seasonal factors significantly influencing the occurrence of all species. Crangon abundance was a minor determinant of whiting and poor cod abundance. Interactions between environmental variables played a minor role in determining the sample abundance of a single species, pouting. Determinants of Gadidae species abundances in the Thames appear to be a complex mix of seasonal and environmental influences, with seasonal influences determining the dominant cyclical pattern and influences of temperature having the greatest effects on short-term variations in the pattern.     

Top-down and bottom-up factors affecting seabird population trends in the California current system (1985–2006)

To characterize the environmental factors affecting seabird population trends in the central portion of the California current system (CCS), we analyzed standardized vessel-based surveys collected during the late spring (May–June) upwelling season over 22 yr (1985–2006). We tested the working hypothesis that population trends are related to species-specific foraging ecology, and predicted that temporal variation in population size should be most extreme in diving species with higher energy expenditure during foraging. We related variation in individual species abundance (number km^?2) to seasonally lagged (late winter, early spring, late spring) and concurrent ocean conditions, and to long-term trends (using a proxy variable: year) during a multi-decadal period of major fluctuations in the El Niño-Southern oscillation (ENSO) and the Pacific decadal oscillation (PDO). We considered both remote (Multivariate ENSO Index, PDO) and local (coastal upwelling indices and sea-surface temperature) environmental variables as proxies for ocean productivity and prey availability. We also related seabird trends to those of potentially major trophic competitors, humpback (Megaptera novaeangliae) and blue (Balaenoptera musculus) whales, which increased in number 4–5-fold midway during our study. Cyclical oscillations in seabird abundance were apparent in the black-footed albatross (Phoebastria nigripes), and decreasing trends were documented for ashy storm-petrel (Oceanodroma homochroa), pigeon guillemot (Cepphus columbus), rhinoceros auklet (Cerorhinca monocerata), Cassin’s auklet (Ptychoramphus aleuticus), and western gull (Larus occidentalis); the sooty shearwater (Puffinus griseus), exhibited a marked decline before signs of recovery at the end of the study period. The abundance of nine other focal species varied with ocean conditions, but without decadal or long-term trends. Six of these species have the largest global populations in the CCS, and four are highly energetic, diving foragers. Furthermore, three of the diving species trends were negatively correlated with the abundance of humpback whales in the study area, a direct competitor for the same prey. Therefore, on the basis of literature reviewed, we hypothesize that the seabirds were affected by the decreasing carrying capacity of the CCS, over-exploitation of some prey stocks and interference competition from the previously exploited, but now increasing, baleen whale populations. Overall, our study highlights the complexity of the ecological factors driving seabird population trends in the highly variable and rapidly changing CCS ecosystem.     

多棘海盘车对太平洋牡蛎、紫贻贝、菲律宾蛤仔摄食选择性的研究

以多棘海盘车(Asterias amurensis)为试验对象,研究了多棘海盘车对3种双壳贝类的摄食选择性、摄食过程中用于搜寻和处理饵料生物的时间分配以及摄食机制等。试验以选择指数、捕食速率、日平均摄食率对多棘海盘车的摄食行为及搜寻和处理时间比进行了描述和统计。结果表明:各处理组多棘海盘车对菲律宾蛤仔的选择指数均为最高,占3种饵料生物的70%以上,显著高于太平洋牡蛎和紫贻贝(P<0.05),而各处理组对太平洋牡蛎的选择性均最低。多棘海盘车对菲律宾蛤仔的捕食速率和日平均摄食率显著高于太平洋牡蛎和紫贻贝。研究发现,在各处理组中多棘海盘车用于搜寻饵料生物的时间无显著差异,但随饵料密度的增加,用于处理饵料的时间逐渐减少。     

Food and feeding of small fish in the Rakaia River,New Zealand

Benthic macroinvertebrates and fish species were collected at monthly intervals during a 12‐month period, from 3 areas of the lower Rakaia River. The composition and abundance of the benthos and stomach contents of the fish were analysed for seasonal trends, food niche breadth and overlap between pairs of fish species, and overlap between the benthos and the diet of each fish species. The diets of bluegilled bully (Gobiomorphus hubbsi), upland bully (G. breviceps), juvenile longfinned eel (Anguilla dieffenbachii), Galaxias brevipinnis, G. paucispondylus, and juvenile brown trout (Salmo trutta) were similar to the proportions of prey species in the benthos. Deleatidium and chironomid larvae dominated the benthos in all seasons and formed the majority of food items in the diet of these species. Food niche overlap between these species was high, indicating potential competition, but preferred habitat and feeding habit differences plus low fish population density and abundance of main prey items probably eliminate the occurrence of any serious competition. The diet of the other fish species differed from this pattern. The common bully (Gobiomorphus cotidianus) ate proportionally more chironomid larvae and fish eggs than occurred in the benthos; torrentfish (Cheimarrichthys fosteri) ate proportionally more chironomid larvae than occurred in the benthos; and quinnat salmon (Oncorhynchus tshawytscha) depended largely upon prey species of terrestial origin, such as adult Deleatidium and dipterans.