Fish assemblage patterns in the littoral zone of a European reservoir

1. Although reservoirs are common aquatic habitats in Europe, there is little quantitative information on the spatial organisation of fish assemblages inhabiting their littoral zones. Consequently, we characterised fish assemblage structure in the littoral zone of a reservoir (Lake Pareloup) in SW France during late spring, summer and early autumn (the growing season).
2. We measured the relative abundance of fish weekly, from mid-May to mid-October, using point abundance sampling by electrofishing. We identified temporal patterns in assemblage structure using hierarchical cluster analysis, and then characterised the spatial distribution of 17 defined ecospecies using a Kohonen self-organising map (SOM, an unsupervised Artificial Neural Network).
3. Our analyses revealed three distinct faunal structures within the littoral zone. From mid-May to mid-July, adults and young-of-the-year (0+) occupied separate habitats, with most 0+ fish in vegetated habitats and adults in open water. From mid-July to late August, some 0+ co-occurred with adults, but most 0+ fishes remained in vegetated areas. Finally, from late August to mid-October, most fish (both 0+ and adults) left the vegetation for unvegetated littoral habitats, the exception being fish species known to be dependent on macrophytes.
4. Contrary to patterns for adult fishes, the 0+ fish assemblage was dynamic. These dynamics were driven by ontogenetic species-specific habitat changes. Consequently, there was little evidence of stable assemblages or strong assemblage–habitat relationships that would be expected of an 'interactive' assemblage. It is likely that the patterns observed are a result of species-specific response to habitat availability in the lake.     

Towards a better understanding of small scale distribution of littoral age-0 fish in a deep-valley reservoir: day or night surveys?

The effect of diel period and littoral habitats on the distribution of age-0 fish was tested in a deep-valley reservoir using boat-modified point abundance sampling by electrofishing (PASE). Day and night samplings unveiled differences in abundance of age-0 fish while recognizing most of the commonly present age-0 fish species in the littoral zone. Night survey provided better information about the abundance of age-0 fish since most species appeared in higher numbers at night. Alternatively, night sampling underestimated bleak (Alburnus alburnus) and gudgeon (Gobio gobio), which were predominantly found in the littoral zone in the daytime. The structure of the age-0 fish assemblage was determined primarily by the characteristics of the littoral habitats, i.e. slope steepness and structure, and three different patterns of habitat use were detected among the fish species. To obtain the most comprehensive assessment of a littoral age-0 fish assemblage and to register all species-specific patterns of habitat use, it is necessary that all littoral habitats in the reservoir are sampled during both day and night periods. The boat-modified PASE could be also used as a standard monitoring tool for routine age-0 fish sampling under difficult conditions of steep-sloped shorelines in large inland waterbodies.     

Habitats and Littoral Zone Fish Community Structure of Two Natural Lakes in Southeast Brazil

The fish community and habitat structure in the littoral zone of lakes Pedra and Hortência, middle Rio Doce Valley, Brazil, were investigated in three sampling periods from August 1992 to May 1993. A total of 9106 fishes were collected, including 11 species in Lake Hortência and 12 species in Lake Pedra. Diversity was higher in Lake Hortência, although total fish abundance was the same for both lakes, with no significant temporal differences. The length distribution of the majority of fish species was similar between lakes. Geophagus brasiliensis was the dominant species (number/biomass) in all habitats of the two lakes in all three sampling periods. A striking difference in total fish abundance in relation to habitat types, which we attribute to physical variables, especially the abundance of macrophytes was observed. Negative impacts attributable to the local Eucalyptus sp. plantations were not detected in the fish community structure of Lake Hortência, which is probably due to the maintenance of the riparian vegetation contributing to similar physical environmental conditions in the littoral zone and high similarity in fish community structure between the two lakes. However, further studies on fish community structure and potential Eucalyptus impacts are suggested as necessary to improve conservation measures of the lakes and their fish faunas.     

Structure of Littoral-zone Fish Communities in Relation to Habitat, Physical, and Chemical Gradients in a Southern Reservoir

How the distribution and abundance of organisms vary across environmental gradients can reveal factors important in structuring aquatic communities. We sampled the littoral-zone fish community in a large reservoir (Lake Texoma) on the Texas–Oklahoma (U.S.A.) border that has pronounced environmental gradients from up- to downlake and between major tributary arms. Our objective was to evaluate the predictability of the littoral-zone fish-community structure from a suite of environmental variables. A stepwise multiple-regression model, with environmental factors at independent variables, explained 64% of the variation in fish species richness across sample sites. The number of species was positively associated with water-column productivity and total Kjedahl nitrogen, and negatively associated with Secchi depth and benthic productivity. Canonical correspondence analysis, with environmental factors as independent variables, explained 63% of the variation in fish-community structure across sites. Equal proportions of the variation in community structure were explained by variables that have strong gradients within the reservoir (e.g., Secchi depth and water-column productivity) and those that represent local habitat variables (e.g., shoreline aspect and substrate type).     

Seasonal response of fish assemblages to habitat fragmentation caused by an impoundment in a Neotropical river

Changes in fish assemblages between the zones above and below Funil dam in Southeastern Brazil were investigated to evaluate the possible impacts of this impoundment in two contrasting seasons: summer/wet and winter/dry. We expect differences in fish assemblage structure and in environmental conditions between seasons and between the reservoir and the zone downriver of the dam. A total of 3,579 individuals comprising 38 species, including six non-natives, were collected. As expected, the comparatively high habitat complexity and water flow regime of the downriver zone favored a richer and more abundant fish assemblage compared with the reservoir, especially in the wet season. In this period, water covers part of the riparian vegetation, increasing habitat availability and nutrient input. Additionally, the dam prevents upriver migration of rheophilics fish species such as the Characiformes Prochilodus lineatus and Leporinus copelandii, and the Siluriformes Pimelodus fur and Pimelodus maculatus, thus increasing shoals below the dam. Although the reservoir represents a simplified ecosystem highly influenced by non-native top predator species (e.g. the Perciformes Cichla kelberi and Plagioscion squamosissimus), seasonal processes (e.g. water level fluctuations and flood pulses) seem to play a role in structuring of the fish assemblage. Environmental variables, mainly turbidity, temperature, and conductivity were significantly associated to spatial-temporal patterns of fish assemblage. In this freshwater tropical reservoir, the spatial scale, rather than the seasonal changes in environmental variables, was the dominant factor structuring fish assemblage in the reservoir and in the zone downriver of the dam.     

Macroinvertebrate communities in Phragmites australis (Cav.) Trin. ex Steud. reed beds and open bank habitats in central victorian streams in Australia

Reed invasion is a common phenomenon of open streams with disturbed riparian vegetation in river catchments. Knowledge of the effects of such vegetation change on aquatic communities is fundamental to river management. Macroinvertebrate fauna in Phragmites australis (Cav.) Trin. ex Steud. and open bank habitats were examined in three rivers in central Victoria in order to understand the effect of such littoral habitat on macroinvertebrates. Data were analysed using Partially Nested Factorial ANOVA with season, river and habitats as main effects. Habitat structure had a significant effect (p<0.05) on macroinvertebrate species richness, however this was not seasonally consistent across the three rivers. There was a significant increase (p<0.05) in macroinvertebrate taxa richness in Phragmites habitats during winter and spring seasons. Total abundance of taxa showed no consistent significant differences in the two habitats. Results of Canonical Analysis of Principle Coordinates indicated significant differences (p<0.05) in macroinvertebrate assemblages between Phragmites and bare bank habitats in all seasons. Habitat selection by taxa could be related to the microphysical environment of the habitats. This study suggests that reed beds create important littoral habitat structures which support diverse macroinvertebrate assemblages.     

Spatial Variation in Fish Assemblages across small Mediterranean Drainages: Effects of Habitat and Landscape Context

Much debate about assemblage organization in stream fish may stem from analysing the effects of both local and large-scale processes on assemblage attributes over whole geographic regions. This study addresses this issue, by examining the contribution of local habitat attributes and landscape context to fish assemblage variation across small Mediterranean drainages in southern Portugal. Fish abundance and species composition was estimated in 28 sites, across 10 drainages, in both a dry year (1999) and a wet year (2001), and related to two sets of variables reflecting habitat and landscape characteristics. Fish showed responses to both sets of variables with variance partitioning indicating that landscape context had important effects on species richness whereas habitat attributes were the primary determinants of local fish abundance. In general, high species richness was associated with larger drainage area and higher rainfall variability, whereas variation in species abundances mostly reflected the influence of width, depth, conductivity, current velocity, substrate size and emergent vegetation. The relative contributions of both landscape context and habitat attributes to species richness and abundance were generally lower in 1999 than in 2001, with much less diversified species–habitat relationships being found in the former dry year. These results point to the dynamic nature of assemblage organization, emphasizing the importance of innovative, multi-scale approaches in advancing our knowledge of fish assemblage structure in Mediterranean streams.     

The significance of littoral and shoreline habitat integrity to the conservation of lacustrine damselflies (Odonata)

Human development of pond and lake shorelines may significantly impact native lacustrine biota including a variety of aquatic macroinvertebrate groups. In an effort to better understand the habitat associations and sensitivities of lacustrine damselflies (Odonata: Zygoptera), we sampled adults in littoral macrophyte habitat during two flight periods at 35 randomly selected pond and lake sites in southern Maine during 2000 and 2001. Data were also collected to help characterize water body, shoreline disturbance, and aquatic vegetation at each study site. Nonmetric multidimensional scaling was used for ordination of damselfly assemblages, and coordinates from the most stable solution were related to site variables using forward stepwise multiple regression. Our results suggest that the diversity and composition of damselfly assemblages is related to the abundance and richness of littoral zone macrophytes, extent of riparian disturbance, benthic substrate granularity, and lake productivity; all variables subject to anthropogenic degradation on excessively developed waterbodies. Additionally, we developed a Habitat Tolerance Index useful for distinguishing between relative habitat specialists and generalists from among a diverse assemblage of 19 lacustrine species. Finally, species-specific damselfly associations with multiple genera of floating and emergent macrophytes were assessed using both nonparametric correlation and multiplicative regression yielding significant relationships for 17 species, including two damselflies of global conservation concern (Enallagma laterale and E. pictum). We conclude that the protection of littoral and shoreline habitat integrity, with special emphasis on emergent and floating macrophytes, is critical to the conservation of lacustrine biodiversity.     

Influences of low-head dams on the fish assemblages in the headwater streams of the Qingyi watershed, China

The influences of low-head dams on the fish assemblages were examined in this study, using fish data collected in six treatment and five reference sites at three low-head dams in the headwater streams of the Qingyi watershed, China. Comparing with those in the reference sites, local habitat variables were significantly altered by low-head dams in the treatment sites, involving wider channel (only in the impoundment area), deeper water and slower flow. Fish species richness varied significantly across seasons, not across site categories, suggesting that these low-head dams did not alter species richness. However, significant decreases in fish abundance and density were observed in the impoundment areas immediately upstream of dams, but not in the plunge areas downstream. Fish assemblage structures kept relative stability across seasons, and their significant difference between-site was only observed between the impoundment areas and the sites far from dams upstream. This variation in assemblage structures was due to the differing relative abundance of some co-occurring species; more lentic but less lotic fish was observed in the impoundment areas. The spatial and temporal patterns of fish assemblages were correlated with local habitat in this study area. Wetted width had negative correlation with fish species richness, abundance and density, respectively. Water temperature also positively affected species richness. In addition, wetted width, water depth, current velocity and substrate were the important habitat variables influencing assemblage structures. Our results suggested that, by modifying local habitat characteristics, low-head dams altered fish abundance and density in the impoundment areas immediately upstream of dam, not in the plunge areas immediately downstream, and thereby influenced fish assemblage structures in these stream segments.     

Habitat rehabilitation for juvenile fish in urban waterways: A case study from Berlin,Germany

This study examines the improvement of juvenile fish recruitment by technical rehabilitation measures implemented in two urban waterways in Berlin, Germany: the River Spree and the Teltow Canal. Rehabilitation sites provide new littoral habitats protected from navigation‐induced flow and wake wash and are intended to improve habitat quality for juvenile fish and to support urban fish diversity. Both sites and four control sites with common embankments have been sampled monthly by point electric fishing (875 dips in total) from May to September 2009. In total, 6,243 fish were caught representing 12 species. Two of the species, roach (Rutilus rutilus) and perch (Perca fluviatilis), dominated the fish community, comprising 81.5% and 16.1% of the total catch. Nine species were rare, with relative abundances <1%, which hampered more detailed species‐specific analyses. Of the dips 33% contained no fish, causing zero‐inflation of the dataset. A hurdle regression model confirmed positive effects of reduced hydraulic disturbances and resulting vegetation cover on juvenile fish occurrence. However, it also showed low oxygen concentrations limiting habitat quality, as observed at the older, densely vegetated rehabilitation site in the River Spree. The general improvement of the juvenile fish assemblage by the rehabilitation measure appeared rather low, as partially vegetated, rectangle trapeze and trapeze profile banks had even higher fish abundance. The study also revealed a need for adaptive maintenance to improve the long‐term performance of this type of rehabilitation structure. With the recovery of aquatic vegetation, the technical bank protection should be adaptively removed to retain the connectivity with the main channel for a sufficient oxygen supply for the long‐term functionality as a fish habitat.     

黄山陈村水库上游河源溪流的鱼类群落及其纵向梯度格局

确定鱼类群落的分布格局及其对人类活动的响应,是合理保护、恢复和管理鱼类多样性的基础。基于2011年5月和10月自黄山陈村水库上游3条河源溪流共39个样点的调查数据,比较研究了溪流间鱼类群落及其纵向梯度格局的异同,着重探讨了人类活动对溪流鱼类群落纵向梯度格局的影响。研究结果显示,同人为干扰较轻的舒溪相比,人为干扰严重的浦溪和麻溪中水宽、底质和植被覆盖率等局域栖息地条件显著变化,这造成了后者的鱼类多样性显著下降及物种组成的显著变化,主要表现为敏感性的地方物种(如宽鳍鱲、光唇鱼、原缨口鳅等)数量减少、耐受性的广布物种(如泥鳅、麦穗鱼、高体鰟鲏等)数量增多。舒溪的鱼类物种数及其组成均与海拔显著相关,但这种"海拔-鱼类群落"关系在麻溪和浦溪中削弱甚至消失。底质、植被覆盖率对舒溪鱼类群落具有重要影响,但对浦溪和麻溪鱼类群落却无显著影响。研究结果表明,在子流域空间尺度上,诸如城镇化发展、土地利用、河道治理等人类活动可通过对局域栖息地条件的影响,导致溪流鱼类多样性下降及其物种组成的变化,破坏鱼类群落的纵向梯度格局,并改变栖息地与鱼类群落之间的联系。     

Comparison of Day and Night Fyke Netting, Electrofishing and Snorkelling for Monitoring a Population of the Threatened Golden Galaxias (Galaxias auratus)

The littoral zone of small off-stream water storage containing a translocated population of Galaxias auratus was sampled fortnightly at day and night with fyke nets, electrofishing and snorkelling over 3 months. Variation in population data provided by each method, including relative abundance indices, size structure, and habitat preferences, were examined. Aspects of behaviour and activity patterns were also investigated. Night sampling using all methods consistently yielded larger catches than day sampling. The size structure of catches varied, with electrofishing at night and fyke netting during the day having higher proportions of juveniles, whilst snorkelling at night and electrofishing during the day had higher proportions of adults. Fyke netting at night yielded by far the largest catches (~3-fold more than other methods) and also captured balanced proportions of juveniles and adults. Galaxias auratus had a strong diel activity pattern and were most active at night. The majority of the population migrated into the littoral zone during the night and back into deeper water during the day. A small number of juveniles remained in the littoral zone and some adults sheltered in the dense cover of species-rich littoral vegetation during the day. Shores with shallow depth profiles appeared to be preferred due to higher catches in these areas using all methods. Based on the results of this study, fyke netting at night in littoral habitats is recommended for monitoring populations of G. auratus. Fyke netting is likely to be an effective method for monitoring other lacustrine galaxiid species; however, further work is required to investigate the effects of habitat variables and fish community structure on activity patterns of galaxiids, and hence their catchability with various methods, in more extensive lentic environments.     

Habitat Use by YOY in the Littoral Zone of an Artificially Heated Reservoir

Both temporal and spatial distributions of a young of the year (YOY) fish community were studied in the littoral zone of a French reservoir. The time survey indicated that littoral occupation highly depends on YOY time of appearance. During late spring and summer, the smallest white bream YOY (L_T < 20 mm) were restricted to complex habitats whereas those of pumpkinseed occupied all available habitats. Growing white bream progressively migrated to open water areas. At that time, pumpkinseed was the only species sampled in abundance in the littoral zone and was still collected in all habitats. The success of the introduction of pumpkinseed might be explained by both its late hatching and its opportunistic strategy in habitat selection enabling it to exploit physical spaces left vacant. (© 2008 WILEY‐VCH Verlag GmbH & Co. KGaA, Weinheim)     

Fish community structure response to major habitat changes within the littoral zone of an estuarine coastal lake

Synopsis The littoral environment and fish fauna of Swartvlei, an estuarine lake, was monitored for four years during which major habitat changes occurred. Initially (1979) the zone was dominated by the submerged macrophytes Potamogeton pectinatus, Chara globularis and Lamprothamnium papulosum. This plant community was replaced by filamentous algal mats during 1980 and with the disappearance of these mats in 1981 the littoral zone was transformed into a sandy habitat. There was a highly significant decline in the numbers of fishes in the littoral zone between the macrophyte and sand phases but no significant decrease in fish biomass between the two phases. Analysis of gill net catches revealed an increase in the CPUE of the family Mugilidae between the macrophyte and sand phases but a decline in the CPUE of vegetation associated species such as Monodactylus falciformis and Rhabdosargus holubi over the same period. The increase in mullet stocks during the sand phase was attributed to epipsammic micro-algal production and the input of allochthonous detritus during the 1981 floods. The three fish species diversity indices used in this study showed minor fluctuations between the habitat phases and these variations were related to changes in the equitability of distribution between the individual species within each habitat type. The numbers of fish species recorded during the macrophyte, algal mat and sand phases varied by less than 20%. The resilience of estuarine fishes to major alterations in their environment was illustrated by the fact that all fish species recorded at the beginning of 1979 were present at the end of 1982, despite major habitat and food resource changes.     

Diversity of medium and large mammals in the Loka Abaya National Park,southern Ethiopia

We evaluated the richness, diversity, and composition of the medium and large mammal community in the Loka Abaya National Park (LANP), southern Ethiopia, and how these parameters differ among four habitat types: wooded grassland, riverine forest, hilly scrubland and wetland, and between seasons. We recorded a total of 2,573 individual animals of 28 medium and large mammal species in the park. This included three globally threatened species: the endangered African wild dog (Lycaon pictus), the vulnerable Leopard (Panthera pardus), and Hippopotamus (Hippopothamus amphibius). Season had little effect on species richness, diversity, and composition both across and within habitat types. However, species richness across seasons was significantly different among the four habitat types, in the declining order of the following: wooded grassland > riverine forest > hilly scrubland > wetland. The strongest similarity in species composition, both across and within seasons, was found between wooded grassland and riverine forest. In terms of relative abundance, mammal assemblage of the wooded grassland and wetland habitats had more evenly distributed number of species with different relative abundance categories. Overall, Anubis Baboon (Papio anubis), Grivet Monkey (Chlorocebus aethiops), and Greater Kudu (Tragelephus strepsiceros) were the three most abundant species across habitat types. In conclusion, findings of our study reveal that LANP plays an important role in Ethiopia's mammal conservation. Our findings will serve as baseline information for managers of the park to make effective conservation decisions and as a baseline for researchers wishing to conduct related ecological studies.     

Body size and substrate association of littoral insects in relation to vegetation structure

Variation in substrate association types and maximum size of aquatic insects were studied in a vegetated littoral zone of three lake basins. The basins differed from each other in trophic status, biomass of benthivorous fish, and abundance of macrophytes. Four types of substrate association – swimmers, crawlers, semisessiles and burrowers, respectively – were assumed to represent decreasing vulnerability to fish predators. Large-sized species were also hypothesised to be more vulnerable to fish predators. The distributions of species traits were examined in relation to vegetation density. Inferring from ``predation hypothesis' opposite selection pressures on the species traits were expected along the vegetation density. Dense macrophyte beds were thought to be dominated by invertebrate predators and open water by fish predators, since the predation efficiency of fish decreases in complex environments. In the case of invertebrate predator domination, large size and higher activity should be favoured traits among the prey species. Distribution patterns of modes of the two studied traits were explored separately for predatory and non-predatory insects. As expected, swimmers and large-sized crawlers were characteristic of the insect assemblages of dense macrophyte beds. The densities of Odonata, Corixidae, Dytiscidae, Ephemeroptera and Sialidae were higher among macrophytes than in open water, where these insect taxa were possibly depleted by fish. On the other hand, the small-sized and fairly immobile Chironomidae were the most abundant group in open water. These results support the existence of a predator transition zone among littoral vegetation, ranging from domination of invertebrate predation among the dense beds to that of fish predation in open water.     

Seasonal abundance and habitat use of bird species in and around Wondo Genet Forest,south‐central Ethiopia

The habitat use and seasonal migratory pattern of birds in Ethiopia is less explored as compared to diversity studies. To this end, this study aimed at investigating the patterns of distribution related to seasonality and the effect of habitat characteristics (elevation, slope, and average vegetation height) on habitat use of birds of Wondo Genet Forest Patch. A stratified random sampling design was used to assess the avian fauna across the four dominant habitat types found in the study area: natural forest, wooded grassland, grassland, and agroforestry land. A point transect count was employed to investigate avian species richness and abundance per habitat type per season. Ancillary data, such as elevation above sea level, latitude and longitude, average vegetation height, and percent slope inclination, were recorded with a GPS and clinometers per plot. A total of 33 migratory bird species were recorded from the area, of which 20 species were northern (Palearctic) migrants while 13 were inter‐African migrants. There was a significant difference in the mean abundance of migratory bird species between dry and wet seasons (t = 2.13, p = .038, df = 44). The variation in mean abundance per plot between the dry and wet seasons in the grassland habitat was significant (t = 2.35, p = .051, df = 7). In most habitat types during both dry and wet seasons, omnivore birds were the most abundant. While slope was a good predictor for bird species abundance in the dry season, altitude and average vegetation height accounted more in the wet season. The patch of forest and its surrounding is an important bird area for migratory, endemic, and global threatened species. Hence, it is conservation priority area, and the study suggests that conservation coupled with ecotourism development is needed for its sustainability.     

Temporal and spatial distribution of microcrustacean zooplankton in relation to turbidity and other environmental factors in a large tropical lake (L. Tana, Ethiopia)

The spatial and seasonal distribution of microcrustacean zooplankton of Lake Tana (Ethiopia) was monthly studied for 2 years. Concurrently, various environmental parameters were measured and related to zooplankton distribution. Canonical Correspondence Analysis (CCA) was used to estimate the influence of abiotic factors and chlorophyll a content in structuring the zooplankton assemblage. Among the environmental factors, zooplankton abundance correlated most strongly with turbidity. Turbidity was negatively correlated with species abundance, especially for Daphnia spp. and to the least extent for Diaphanosoma spp. Analysis of variance (ANOVA) was used to determine spatial (littoral, sublittoral and pelagic zone) and temporal (four seasons) variation in zooplankton abundance. We observed significant temporal differences in zooplankton abundance, with highest densities during dry season (November–April). Only cladocerans showed significant differences in habitat use (highest densities in the sublittoral zone). %     

Habitat preferences of small mammals in tropical open-forest of the Northern Territory

Pattern analysis was used to investigate the habitat preferences of five small mammal species in tropical open-forest of the Northern Territory. Fifty-one sites were classified by faunal abundance and the groups examined for significant differences in vegetation structural attributes and plant species in both dry and wet seasons. The omnivore Isoodon macrourus showed strong association with floristic and vegetation structural attributes only in the dry season, when areas with a dense understorey of small trees and shrubs and a high percentage of leaf litter cover were favoured. Of the two primarily carnivorous species, Antechinus bellus was related strongly with floristic and structural attributes in both seasons and showed a consistent preference for areas with relatively dense low-level foliage (< 2m). By contrast, Dasyurus hallucatus was associated more with the structurally simple open-forest types. Of the two mainly herbivorous species Mesembdomys gouldii showed associations only withfioristics in both seasons, while the habitat relationships of Trichosurus arnhemensis were very weak due to its low abundance in the study area. The number and strength of animal/habitat relationships were greatest in the dry season. Forest types with dense mid-level foliage and abundant hollow logs and leaf litter had the greatest mammal richness and abundance; these areas may be critical to the survival of local mammal populations. A comparison of site-groups, defined independently on the basis of fauna, floristics or structure, showed that animal groups overlapped one to six of the habitat groups. The animal's perception of s‘habitat’ may thus differ from that of humans, or that defined by measurement of habitat attributes.