The biology and ecology of lotic microcrustaceans

• 1 Copepoda, Ostracoda and ‘Cladocera’ are important meiobenthic Crustacea which can be both numerically abundant and species rich in running waters. Harpacticoids and ostracods are well adapted to benthic life because they are typical crawlers, walkers, and burrowers. Many cladocerans are substratum dwellers, but most benthic species among these can also swim. Cyclopoids which are generally good swimmers are nevertheless often bottom frequenters and actively colonise sediment interstices (the hyporheic zone).
  • 2 The subclass Copepoda includes 10 orders. With 53 families, the order Harpacticoida dominates the benthos. Only five of these families are represented in fresh waters (ca. 1 000 species and subspecies). The order Cyclopoida includes 12 families of which the Cyclopidae is well represented in freshwater habitats with 900 species and subspecies. Freshwater Ostracods belong to the order Podocopida (5 000 species) with three superfamilies occurring in running fresh waters. The group ‘Cladocera’ contains four orders, 12 families, more than 80 genera, and 450–600 freshwater species. Most of the benthic species are found in the families Chydoridae (39 genera), Macrothricidae, Ilyocryptidae and Sididae.
• 3 For each of the three major taxa, morphological characteristics are presented, specimen collection and preparation are described and references to available taxonomical keys are provided.
• 4 Biological characteristics are extremely diverse among and within the three taxa, resulting in a great variety of strategies in meiobenthic crustaceans. Characteristics of reproduction, sexual dimorphism, cyclomorphosis and population parameters (i.e. clutch size, lifespan, growth, moulting) are provided for some of the most common species.
• 5 Important differences between the three main taxa were found at the species level. Ecological requirements such as hydraulic microhabitats and geomorphologic features of the streambed are the major determinants of species diversity and abundance for benthic microcrustacea of lotic habitats. Many studies on the ecology of these communities are limited by a lack of knowledge of the life history characterisitics of lotic (especially interstitial) crustacean populations.

     

Faunal response to benthic and hyporheic sedimentation varies with direction of vertical hydrological exchange

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Spatial distribution patterns of the hyporheic invertebrate communities in a polluted river in Romania

The purpose of this study was to examine the sensitivity, in a field situation, of the hyporheic fauna to pollution by heavy metals and also to test the use of oxidative stress enzymes produced by this fauna as a sensitive indicator of oxidative stress generated by chemical contamination. This was done by surveying the patterns of distribution, structure, and composition of hyporheic invertebrate communities in one of the most polluted rivers in Romania. Twelve permanent sampling stations with differing water qualities were established along a 180 km transect of the Arie? River. Data on hyporheic invertebrate abundance and richness, chemistry of the surface and hyporheic water and interstitial suspended particles were analyzed via multifactorial analyses. In the downstream, more polluted stations, epigean species were less abundant and hyporheic communities, especially macrocrustaceans and oligochetes, became dominant. The higher levels of hyporheic invertebrate biodiversity in the moderately polluted stations compared to highly polluted, and the increase of the number of some hyporheos (especially macrocrustaceans) in the moderately polluted stations, suggested that the hyporheic fauna was more tolerant of heavy metal pollution than the surface water fauna of the area. However, the different richness and abundance of hyporheic fauna in sites of similar water chemistry suggested that additional factors, such as sediment structure are shaping the spatial distribution of hyporheic fauna. Strong correlations between superoxide dismutase (SOD) activity in pooled tissues extracts and some chemical parameters suggest that oxidative stress enzymes may prove to be sensitive indicators of chemical pollution in hyporheic zones.     

Influence of hyporheic zone characteristics on the structure and activity of microbial assemblages

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Cypriniform fish in running waters reduce hyporheic oxygen depletion in a eutrophic river

1. The hyporheic zone is an important habitat for benthic invertebrates and early-developmental stages of gravel spawning fish. However, the eutrophication of running waters and, in turn, the excessive periphyton biomass leads to its biological clogging. The result of these processes is oxygen depletion and a reduction in the habitat quality of the hyporheic zone.
2. This study assessed whether top-down effects of two important European river fish species, the large herbivorous cypriniform common nase (Chondrostoma nasus, L.) and the large omnivorous cypriniform European chub (Squalius cephalus, L.), can reduce eutrophication effects in the hyporheic zone. A 4-week mesocosm-based field experiment in a eutrophic river was conducted using cage enclosures stocked or not with either nase or chub.
3. The top-down control of periphyton was expected to reduce biological clogging and thereby increase oxygen availability in the hyporheic zone. Accordingly, we hypothesised that in enclosures stocked with either fish the concentrations of dissolved oxygen in the hyporheic zone would be higher and the periphyton biomass would be lower than in enclosures without fish stocking.
4. Hyporheic oxygen concentrations were significantly higher in enclosures stocked with either nase or chub than in enclosures without fish stocking. However, periphyton ash-free dry mass was significantly reduced only in enclosures stocked with nase, not in those stocked with chub. Thus, the positive effects of nase and chub on hyporheic oxygen availability were caused by different mechanisms.
5. Our results demonstrate that nase and chub can reduce eutrophication effects in the hyporheic zone of running waters. Hence, protecting and enhancing stocks of herbivorous and omnivorous fish will contribute to restoring the hyporheic zone in efforts to preserve biodiversity in eutrophic rivers.

     

Responses of microbial activity in hyporheic pore water to biogeochemical changes in a drying headwater stream

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Geomorphic influence on intraspecific genetic differentiation and diversity along hyporheic corridors

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Surface water–groundwater exchanges in three dimensions on a backwater of the Rhône River

1. Hydrological exchange between the surface stream and the hyporheic zone is well documented in the main channel of rivers, especially at the reach scale. Hydrological processes of advection/convection occur at different scales, and in secondary channels of large rivers little is known about these exchanges in the hyporheic zone on a broad scale (i.e. kilometres). This work studied exchanges of water and biota in a secondary channel on a large scale (4 km), using a three-dimensional framework. 2. The exchanges of water were described using physicochemical indicators of surface and groundwaters. Samples of water and biota were taken in three dimensions: (i) vertically from benthic (i.e. 0.20 m below the surface of the substratum) to hyporheic (0.50 m) and deep interstitial (1.0 m) zones; (ii) laterally from the right to the left bank (i.e. right, middle and left positions); and (iii) longitudinally from upstream to downstream (seven stations regularly distributed along the channel). 3. The physicochemical indicators clearly revealed hydrological heterogeneity in the longitudinal and vertical dimensions, whereas lateral variability was not significant. 4. Spatial distribution of biota exhibited strong longitudinal variations that were not gradual as predicted by an upstream/downstream continuum, but patchy and discontinuous. No significant differences were found between the three positions across the channel. 5. Analyses of both physicochemical and faunal data sets produced matched ordination of samples and stations, indicating that interstitial–surface flow relationships appear to be an important governing factor in the distribution of interstitial biota at this broad scale. 6. Results are discussed in relation to the hypothetical three-dimensional models of the hyporheic zone in rivers. Contrasting with other observations on the main channel (where advection/convection patterns are dominated by morphological changes of the river-bed morphology), it is proposed that water exchanges in backwaters are more likely to be related to local modifications of stream-bed porosity.     

Towards a biological and chemical definition of the hyporheic zone in two Canadian rivers

SUMMARY. 1. A series of samples of interstitial water and fauna was taken along transects from the channel into the bank in two small rivers in southern Ontario, Canada. These were examined for any discontinuities which might indicate the position of the hyporheic/groundwater interface. 2. There were several chemical discontinuities in Duffin Greek, with “break lines” occurring from about the river margin obliquely downwards under the bank for dissolved oxygen and carbon dioxide, B.O.D., alkalinity, suspended solids and amount of organic matter. Break lines in nitrate and sulphide concentration ran from near the margin obliquely down under the river bed. In the Rouge River, a discontinuity extending from a point approximately 1.5 m landwards from the margin obliquely down under the river was indicated by dissolved carbon dioxide, B.O.D., conductivity, suspended solids, organic matter, nitrate and alkalinity. 3. Ordination (DECORANA) and community classification (TWIN-SPAN) revealed that, in both rivers, linear distance from mid-river was the major factor associated with community structure. In both rivers the community under the bank was distinct from the river community and these two communities were separated by another community characteristic of the river margin. In Duffin Creek the classification procedure additionally discriminated surface and interstitial sub-sets within the river community. 4. Most taxa showed no significant correlations with the chemistry of the interstitial water but. in Duffin Creek, the densities of the copepod Diacyclops crasscaudis brachycercus (Kiefer) and Oligochaeta were positively correlated with nitrate: worms were also negatively correlated with sulphide. In the Rouge River, density of hyporheic animals was negatively correlated with both conductivity and alkalinity of interstitial water; oligochactes were positively correlated with both nitrate and organic matter; and density of nematodes was positively correlated with sulphide concentration. 5. The hyporheic faunas of these two rivers were dominated by insects, particularly chironomids. Compared with the hyporheos of rivers in Europe and Colorado, the two Ontario rivers lack significant numbers of harpacticoid copepods as well as bathynellacid, amphipod and isopod crustaceans.     

The effects of increased flow and fine sediment on hyporheic invertebrates and nutrients in stream mesocosms

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Nutrient and flow vector dynamics at the hyporheic/groundwater interface and their effects on the interstitial fauna

Environmental conditions in the interstices beneath streams and rivers with porous beds are unlike those found either on the bed surface or in the true groundwater. For most of the year, in many streams, the bulk of the water in the hyporheic zone is provided by baseflow but, as it passes across the hyporheic/groundwater interface, the physical and chemical nature of this groundwater changes, probably in response to mixing with surface water. Factors promoting the influx of surface water are associated with features of the bed and channel morphology. The upper and lower boundaries of the hyporheic zone are thought to vary in time, but at any instant they can be defined. As a habitat, the hyporheic zone fits the definition of an ecotone, although certain adverse features may result in reduced species diversity. There are limited, correlative, data available on the relationship of the fauna (hyporheos) to interstitial conditions and further study of the general biology of both species and populations is needed. In an attempt to stimulate future research on these systems, some preliminary models of hyporheic dynamics are proposed.     

Benthic macro- and meso-invertebrates of a sandy riverbed in a mountain stream, central Japan

Quantitative samples of benthic invertebrates were collected from a sandy riverbed of a mountainous stream (Kozu site of Takami-gawa stream, Nara Prefecture), central Japan by core samplers in five sampling occasions through the years 2008–2009. A total of 120 taxa were identified, representing 55 families and 97 genera. Insects formed about 92% of the total recorded taxa and 88% of individuals’ abundance. A total of 111 taxa of aquatic insects, belonging to 49 families and 92 genera, were identified and represented by ten orders. Oligochaeta and Acari were dominant non-insect invertebrates. Diptera was the most diverse insect group, followed by Trichoptera and Ephemeroptera. Dominant taxa were mesoinvertebrates, younger stages of macroinvertebrates, both of which predominantly inhabit the interstitial zone of a sandy riverbed. Both taxon richness and invertebrate abundance were higher in February 2009 and lower in April and August 2008. A few major invertebrate taxa demonstrated distinct seasonal trends; i.e. Nymphomyia alba, Rheosmittia, and Corynoneura were abundant in February 2009. Newly hatched larvae of Larcasia akagiae were abundant in May 2008. This study also demonstrated the effectiveness of core samplers to collect small-sized benthic fauna that inhabit the interstitial or hyporheic zone of the sandy riverbed.     

A new inexpensive suction apparatus for sampling arthropods in grassland

Abstract.

• 1 We describe a new technique for sampling arthropods from grassland and other vegetation, using a modified petrol-driven suction apparatus called a‘Blow & Vac’.
• 2 The new apparatus was tested by comparing its efficiency in extracting arthropods from vegetation with that of a conventional Dietrick, or‘D-Vac’, suction sampler.
• 3 Unimpeded air velocity inside the suction tube was approximately 4 times that of the D-Vac.
• 4 In field trials on semi-natural grasslands, the Blow & Vac caught more individuals of most spider and certain beetle species compared to the D-Vac and approximately equal numbers of Auchenorrhyncha.
• 5 We conclude that the‘Blow & Vac’has considerable potential for sampling arthropod populations. Its low cost and weight make it a suitable alternative to the conventional D-Vac apparatus.

     

The relative contributions of refugium types to the persistence of benthic invertebrates in a seasonal snowmelt flood

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Toxic benthic freshwater cyanobacterial proliferations: Challenges and solutions for enhancing knowledge and improving monitoring and mitigation

1. This review summarises knowledge on the ecology, toxin production, and impacts of toxic freshwater benthic cyanobacterial proliferations. It documents monitoring, management, and sampling strategies, and explores mitigation options.
2. Toxic proliferations of freshwater benthic cyanobacteria (taxa that grow attached to substrates) occur in streams, rivers, lakes, and thermal and meltwater ponds, and have been reported in 19 countries. Anatoxin- and microcystin-containing mats are most commonly reported (eight and 10 countries, respectively).
3. Studies exploring factors that promote toxic benthic cyanobacterial proliferations are limited to a few species and habitats. There is a hierarchy of importance in environmental and biological factors that regulate proliferations with variables such as flow (rivers), fine sediment deposition, nutrients, associated microbes, and grazing identified as key drivers. Regulating factors differ among colonisation, expansion, and dispersal phases.
4. New -omics-based approaches are providing novel insights into the physiological attributes of benthic cyanobacteria and the role of associated microorganisms in facilitating their proliferation.
5. Proliferations are commonly comprised of both toxic and non-toxic strains, and the relative proportion of these is the key factor contributing to the overall toxin content of each mat.
6. While these events are becoming more commonly reported globally, we currently lack standardised approaches to detect, monitor, and manage this emerging health issue. To solve these critical gaps, global collaborations are needed to facilitate the rapid transfer of knowledge and promote the development of standardised techniques that can be applied to diverse habitats and species, and ultimately lead to improved management.

     

A survey of stream invertebrates in the Cow Green basin (Upper Teesdale) before inundation

In 1967 a programme was initiated by the Freshwater Biological Association to study fish populations within the proposed Cow Green reservoir basin and in the Tees downstream of the dam, before and after impoundment. This paper describes the result of a supporting study on aquatic invertebrates covering the pre-impoundment period 1967–70. The benthic faunas of six streams in the reservoir basin, the Tees below Cauldron Snout and Maize Beck, a tributary ofthe Tees below the dam, were studied. Species list are presented for each habitat and changes in seasonal and annual abundance are discussed. All areas sampled lie at altitudes between 440 and 550 m O.D. and are situated amongst moorland and limestone grassland. Conditions in the streams ranged from slow-flowing peaty reaches to small streams with moss-covered bottoms and larger stony rivers and streams with relatively unstable bottoms. Samples were taken in riffles and pools using the ‘kick’ method wherever possible. An attempt was made to quantify kick-sample catches by comparing them with shovel-sample catches which cover a known area of stream bottom. It was found that 10 5 kicks gave a catch equivalent to the populations of 1 m², giving a population density of about 1200 animals/m for the reservoir-basin riffles. Over 120 taxa were recorded, 100 of which were at the species level. In the reservoir-basin streams, 116 taxa were found with seventy-one in Maize Beck and fifty-six in the Tees below Cauldron Snout. Ephemeroptera were the most abundant group in the reservoir basin and Maize Beck faunas with Rhithrogena semicohrata, Heptagenia lateralis and Baetidae being the most abundant forms, although Ecdyonurus spp. especially E. dispar were much more common in Maize Beck. In other groups Leuctra spp. and Gammarus pulex were very common. In the Tees below Cauldron Snout Limnaea peregra, Chironomidae and Baetidae formed the bulk of the fauna and Plecoptera were uncommon. Amongst the reservoir basin streams Weelhead Sike supported the largest number of species and species groups (eighty-one), and all the streams had forty-nine or more taxa represented. Information on seasonal changes in the numbers of those species or species groups composing 90% or more of the total fauna is presented. Faunal density was high in May with Ephemeroptera, particularly Ecdyonuridae, and the plecopteran Leuctra inermis being the most abundant forms. In August, the numbers of animals appeared t o fall and common members of the community were Baetidae, Diptera, Leuctra fusca and Ecdyonuridae. In October, Ecdyonuridae particularly R. semicolorata, were most abundant. A comparison ofthe bottom fauna of riffles and pools was made and more animals were found in riffles than in pools. The effect of gravel extraction on the bottom fauna of the Tees was examined. A severe drop in the numbers of animals was observed after extraction. Diptera were the first group to return to their pre-disturbance density. Elminthidae and Annelida were worst affected and slowest to recover. The fauna of the area is discussed and possible reasons for its relative species richness are put forward. Habitat diversity and chemical richness appear to be the most likely reasons for the relatively large number of species found.     

The boundaries of river systems: the metazoan perspective

1. This overview of metazoans associated with the riparian/groundwater interface focuses on the fauna inhabiting substratum interstices within the stream bed and in alluvial aquifers beneath the floodplain. The objective is to integrate knowledge of habitat conditions and ecology of the interstitial fauna into a broad spatiotemporal perspective of lotic ecosystems. 2. Most aquatic metazoans of terrestrial ancestry, secondarily aquatic forms including insects and water mites (Hydracarina), are largely confined to surface waters (epigean), most of the time penetrating only the superficial interstices of the stream bed. 3. Primary aquatic metazoans include crustaceans and other groups whose entire evolutionary histories took place in water. Some species are epigean, whereas other members of the primary aquatic fauna are true subterranean forms (hypogean ) , residing deep within the stream bed and in alluvial aquifers some distance laterally from the channel. 4. The hypogean/epigean affinities of interstitial animals are reflected in repetitive gradients of species distribution patterns along vertical (depth within the stream bed), longitudinal (riffle/pool), and lateral (across the floodplain) spatial dimensions, as well as along recovery trajectories following floods (temporal dimension). 5. Fluvial dynamics and sediment characteristics interact to determine hydraulic conductivity, oxygen levels, pore space, particle size heterogeneity, organic content and other habitat conditions within the interstitial milieu. 6. Multidimensional environmental gradients occur at various scales across riparian/groundwater boundary zones. The spatiotemporal variability of hydrogeomorphological processes plays an important role in determining habitat heterogeneity, habitat stability, and connectivity between habitat patches, thereby structuring biodiversity patterns across the riverine landscape. 7. The erosive action of flooding maintains a diversity of hydrarch and riparian successional stages in alluvial floodplains. The patchy distribution patterns of interstitial communities at the floodplain scale reflect, in part, the spatial heterogeneity engendered by successional processes. 8. Interstitial metazoans engage in passive and active movements between surface waters and ground waters, between aquatic and riparian habitats, and between different habitat types within the lotic system. Some of these are extensive migrations that involve significant exchange of organic matter and energy between ecosystem compartments. 9. The generally high resilience of lotic ecosystems to disturbance is attributable, in part, to high spatiotemporal heterogeneity. Habitat patches less affected by a particular perturbation may serve as ’refugia ‘; from which survivors recolonize more severely affected areas. Mechanisms of refugium use may also occur within habitats, as, for example, through ontogenetic shifts in microhabitat use. Rigorous investigations of interstitial habitats as refugia should lead to a clearer understanding of the roles of disturbance and stochasticity in lotic ecosystems. 10. Development of realistic ’whole river ‘; food webs have been constrained by the exclusion of interstitial metazoans, which may in fact contribute the majority of energy flow in lotic ecosystems. A related problem is failure to include groundwater/riparian habitats as integral components of alluvial rivers. A conceptual model is presented that integrates groundwater and riparian systems into riverine food webs and that reflects the spatiotemporal complexity of the physical system and connectivity between different components. 11. Interstitial metazoans also serve as ’ecosystem engineers, ‘; by influencing the availability of resouces to other species and by modifying habitat conditions within the sediment. For example, by grazing on biofilm, interstitial animals may markedly stimulate bacterial growth rates and nutrient dynamics. 12. Although there has been a recent surge of interest in the role of interstitial animals in running waters, the knowledge gaps are vast. For example, basic environmental requirements of the majority of groundwater metazoans remain uninvestigated. Virtually nothing is known regarding the role of biotic interactions in structuring faunal distribution patterns across groundwater/riparian boundary zones. Interstitial metazoans may contribute significantly to the total productivity and energy flow of the biosphere, but such data are not available. Nor are sufficient data available to determine the contribution of groundwater animals to estimates of global biodiversity. 13. Effective ecosystem management must include groundwater/riparian ecotones and interstitial metazoans in monitoring and restoration efforts. Evidence suggests that a ’connected ‘; groundwater/riparian system provides natural pollution control, prevents clogging of sediment interstices and maintains high levels of habitat heterogeneity and successional stage diversity. River protection and restoration should maintain or re-establish at least a portion of the natural fluvial dynamics that sustains the ecological integrity of the entire riverine–floodplain–aquifer ecosystem. Keywords: groundwater/riparian ecotones, hyporheic habitat, epigean, hypogean, interstitial fauna, biodiversity, food webs     

Reconstructing ancient ecological conditions from insect remains; some problems and an experimental approach

• 1 Analysis of insect ‘death assemblages’ from ancient natural and anthropogenic deposits can provide valuable information about conditions in the past but the theoretical basis of such investigations is weak, especially with respect to the quantification of habitats on archaeological sites.
• 2 Evidence is presented for the existence of a transported component, termed ‘background fauna’, in death assemblages, as well as the remains of insects derived from the immediate surroundings.
• 3 The background component is derived principally from airborne insects and from bird droppings and may seriously detract from the accuracy of reconstructions of past ecological conditions. Factors affecting its relative importance are discussed.
• 4 As well as including species from distant habitats, the background fauna may be biased in favour of species from temporary habitats.
• 5 Some solutions to the problems presented by the background fauna are discussed. A high proportion of a species or ecological group will usually be evidence that its habitat existed near to the deposit enclosing the fauna, although there may be exceptions.
• 6 The structure of the background rain of insects has probably altered with changes in town and countryside brought about by man. Attempts to recognize background fauna in ancient assemblages must take account of this.
• 7 Many ancient assemblages, especially if small, will give no useful ecological information.

     

Effects of buried leaf litter and vertical hydrologic exchange on hyporheic water chemistry and fauna in a gravel-bed river in northern New South Wales, Australia

1. Large amounts of coarse particulate organic matter (CPOM) are buried in the sand and gravel beds of many rivers during spates. The effects of these patchily distributed resources on hyporheic invertebrates and water chemistry are poorly understood. Buried CPOM may provide local ‘hot-spots’ of food for hyporheic detritivores and their predators, alter nutrient supply to nearby sediment biofilms, and generate habitat for some invertebrates. 2. To examine potential short-term effects on hyporheic water chemistry, nutrient concentrations and invertebrate assemblage composition, leaf packs were buried in downwelling (surface water infiltrating the hyporheic zone) and upwelling (hyporheic water emerging to the surface) zones at two sites along a gravel-bed river in northern New South Wales. At one site, pits were excavated to simulate leaf burial (procedural control) and plastic ‘leaves’ were buried to test whether invertebrates might respond to leaves as refuges rather than food. Hyporheic CPOM, sediment size fractions, and interstitial silt content were also quantified at these sites. 3. Dry weights of naturally buried CPOM (leaf litter and wood fragments) varied substantially (0.6–71.7 g L^–1 sediment). Amounts of CPOM did not differ between up- vs. downwelling zones or between sites. Hyporheic dissolved oxygen saturation was generally high (> 75%), and was lower in upwelling zones. The hyporheos was dominated taxonomically by water mites (≈ 20 species), whereas small oligochaetes were most abundant (40% of total abundance). Tiny instars of elmid beetle larvae and leptophlebiid mayfly nymphs were also common. Before experimental manipulation, faunal composition differed between up- and downwelling zones. In upwelling zones, bathynellaceans and blind peracarids were found, whereas small individuals of the surface benthos were common in samples from downwelling zones. This validated stratification of the experiment across zones of hydrologic exchange. 4. Twenty days after leaf burial, there was no effect of the treatments at either site on changes in most variables, including mean numbers of taxa and individuals per sample. Similarly, changes in faunal composition of the hyporheos in the treatments paralleled those in the controls except for a weak response in the buried leaves treatment in the upwelling zone at site 1. Artificially buried leaf litter does not seem to influence hyporheic water chemistry or fauna at these two sites. It is probable that naturally buried leaf litter is swiftly processed soon after entrainment and that repeating this experiment immediately after a flood may yield different results.     

The good,the bad,and the ugly: which Australian terrestrial mammal species attract most research?

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