Patterns in the distribution of fish assemblages in Río Amacuzac,Mexico: influence of abiotic factors and biotic factors

Patterns in spatial and seasonal distribution of fish communities were analyzed in the Río Amacuzac, Mexico, and their relationship to environmental variables and habitat characteristics. The PCA biplot of the study sites and environmental factors showed the first two axes accounting for 52.93% of the variance. The diagram showed the study sites ordination in environmental gradients. The first axis explained variables related to habitat characteristics and temperature (36.30%) and second axis arranged the sites in physicochemical and water quality environmental gradients (conductivity, dissolved oxygen, orthophosphates, ammonium, pH) displaying seasonal variation. Fifteen species were recorded, eight of them are exotic. One new record appeared for the Río Amacuzac: Pterygoplichthys disjunctivus is exotic. Study sites with highest species richness were: 5, 7 and 9 (twelve species each one); while the study sites with low species richness were 1, 2, 3 and 6 (eight species each one). Six of the species were distributed throughout the whole river. Based on the composition of the fish fauna, the study sites form two groups and the analysis of fish species displays three groups according to the Bray–Curtis index. The diagram of the canonical correspondence analysis relates environmental parameters to the abundance of fish species and showed that the first two axes exhibit 78.31% of the explained variance. Species richness had a spatial pattern associated to the introduction of exotic species for ornamental uses. According to the results of the importance value index (IVI), the dominant species were the poecilids Poeciliopsis gracilis and Heterandria bimaculata, small fishes that were introduced in the river. Río Amacuzac has a biotic alteration in the structure of fish communities due to the invasion of exotic species, representing risks to the integrity of the native fish fauna.     

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.     

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.     

Juvenile coral reef fish use sound to locate habitats

There is limited knowledge of the orientation cues used by reef fish in their movement among different habitats, especially those cues used during darkness. Although acoustic cues have been found to be important for settlement-stage fish as they seek settlement habitats, only a small number of studies support the possible role of acoustic cues in the orientation of post-settled and adult reef fish. Therefore, the aim of this study was to determine whether habitat-specific acoustic cues were involved in the nocturnal movements of juvenile reef fish to small experimental patch reefs that were broadcasting sound previously recorded from different habitats (Fringing Reef, Lagoon, Silent). Juvenile fish arriving at each patch reef were caught the next morning by divers and were identified. There were a greater number of occasions when juvenile fish (from all species together) moved onto the patch reefs broadcasting Fringing Reef and Lagoon sound (43 and 38%, respectively) compared to Silent reefs (19%) (χ² = 33.5; P < 0.05). There were significantly more occasions when juvenile fish from the family Nemipteridae were attracted to the patch reefs broadcasting Lagoon sound (63%) versus those reefs broadcasting either Fringing Reef sound (31%) or Silent (6%). In contrast, there were more occasions when juveniles from the family Pomacentridae were attracted to the patch reefs broadcasting Fringing Reef sound (56%) than either Lagoon (24%) or Silent patch reefs (20%) (χ² = 19.5; P < 0.05). These results indicate that some juvenile fish use specific habitat sounds to guide their nocturnal movements. Therefore, the fish are able to not only use the directional information contained in acoustic cues, but can also interpret the content of the acoustic signals for relevant habitat information which is then used in their decision-making for orientation.     

New records of the restinga antwren Formicivora littoralis Gonzaga and Pacheco (Aves, Thamnophilidae) in the state of Rio de Janeiro, Brazil: inland extended range and threats

The Restinga Antwren (Formicivora littoralis) has a narrow distribution range in southeastern Brazil, and it is a typical species of restinga habitat (sandy coastal plain vegetation). In this paper, we describe two new records for the species (22 degrees 51' 45" S and 42 degrees 14' 13" W; 22 degrees 51' 14" S and 42 degrees 11' 47" W) in the northern margin of the Araruama Lagoon, which represent a new inland limit for its distribution (11 km), besides assessing the current state of its habitat. We recorded supposed isolated subpopulations, most of them due the accelerated human-made fragmentation. The Massambaba Environmental Protection Area comprises the larger continuous extent of the suitable habitat for the Restinga Antwren, being essential to its long-term existence. However, the region lacks effective protected areas and, besides urgent practical measures, we recommend an accurate mapping and populational studies on this species.     

Day and night trophic variations of dominant fish species in a lagoon influenced by freshwater seeps

The aim of this study was to determine the trophic structure and nycthemeral variations in the diet of dominant fish species (Ariopsis felis, Bairdiella chrysoura, Micropogonias undulatus, Eucinostomus gula, Eucinostomus argenteus, Lagodon rhomboides and Sphoeroides testudineus) in Celestun Lagoon, a biosphere reserve located in the southern Gulf of Mexico, and influenced by freshwater seeps. A total of 1473 stomachs were analysed and nine trophic groups were recorded. Bray–Curtis analyses with analyses of similarity (ANOSIM) statistical tests were used to determine two groups of feeding guilds: zoobenthivores and omnivores, with significant differences between time and habitat. The relationships between fish feeding habits, size class and environmental variables were investigated using canonical correspondence analysis (CCA). Most of the species showed a low niche breadth with high specialization towards amphipod consumption, with the exception of L. rhomboides (0·60), which indicated generalist feeding. This study in a protected area is an important source of information for drawing up conservation policies in relation to the management of aquatic resources, and will aid in the establishment of priority areas for conservation.     

Gravel dredging alters diversity and structure of riverine fish assemblages

1. Human activities affect fish assemblages in a variety of ways. Large‐scale and long‐term disturbances such as in‐stream dredging and mining alter habitat and hydrodynamic characteristics within rivers which can, in turn, alter fish distribution. Habitat heterogeneity is decreased as the natural riffle–pool–run sequences are lost to continuous pools and, as a consequence, lotic species are displaced by lentic species, while generalist and invasive species displace native habitat specialists. Sediment and organic detritus accumulate in deep, dredged reaches and behind dams, disrupting nutrient flow and destroying critical habitat for habitat specialist species. 2. We used standard ecological metrics such as species richness and diversity, as well as stable isotope analysis of δ¹³C and δ¹⁵N, to quantify the differences in fish assemblages sampled by benthic trawls among dredged and undredged sites in the Allegheny River, Pennsylvania, U.S.A. 3. Using mixed‐effects models, we found that total catch, species richness and diversity were negatively correlated with depth (P < 0.05), while species richness, diversity and proportion of species in lithophilic (‘rock‐loving’) reproductive guilds were lower at dredged than at undredged sites (P < 0.05). 4. Principal components analysis and manova revealed that taxa such as darters in brood hider and substratum chooser reproductive guilds were predominantly associated with undredged sites along principal component axis 1 (PC1 and manova P < 0.05), while nest spawners such as catfish and open substratum spawners including suckers were more associated with dredged sites along PC2 (P < 0.05). 5. Stable isotope analysis of δ¹³C and δ¹⁵N revealed shifts from reliance on shallow water and benthic‐derived nutrients at undredged sites to reliance on phytoplankton and terrestrial detritus at deep‐water dredged sites. Relative trophic positions were also lower at dredged sites for many species; loss of benthic nutrient pathways associated with depth and dredging history is hypothesised. 6. The combination of ecological metrics and stable isotope analysis thus shows how anthropogenic habitat loss caused by gravel dredging can decrease benthic fish abundance and diversity, and that species in substratum‐specific reproductive guilds are at particular risk. The effects of dredging also manifest by altering resource use and nutrient pathways within food webs. Management and conservation decisions should therefore consider the protection of relatively shallow areas with suitable substratum for spawning for the protection of native fishes.     

Diet reveals links between morphology and foraging in a cryptic temperate reef fish

Predators select prey so as to maximize energy and minimize manipulation time. In order to reduce prey detection and handling time, individuals must actively select their foraging space (microhabitat) and populations exhibit morphologies that are best suited for capturing locally available prey. We explored how variation in diet correlates with habitat type, and how these factors influence key morphological structures (mouth gape, eye diameter, fin length, fin area, and pectoral fin ratio) in a common microcarnivorous cryptic reef fish species, the triplefin Helcogrammoides cunninghami. In a mensurative experiment carried out at six kelp‐dominated sites, we observed considerable differences in diet along 400 km of the Chilean coast coincident with variation in habitat availability and prey distributions. Triplefins preferred a single prey type (bivalves or barnacles) at northern sites, coincident with a low diversity of foraging habitats. In contrast, southern sites presented varied and heterogeneous habitats, where triplefin diets were more diverse and included amphipods, decapods, and cumaceans. Allometry‐corrected results indicated that some morphological structures were consistently correlated with different prey items. Specifically, large mouth gape was associated with the capture of highly mobile prey such as decapods, while small mouth gape was more associated with cumaceans and copepods. In contrast, triplefins that capture sessile prey such as hydroids tend to have larger eyes. Therefore, morphological structures co‐vary with habitat selection and prey usage in this species. Our study shows how an abundant generalist reef fish exhibits variable feeding morphologies in response to the distribution of potential habitats and prey throughout its range.     

Habitat Use by Fishes in Coral Reefs,Seagrass Beds and Mangrove Habitats in the Philippines

Understanding the interconnectivity of organisms among different habitats is a key requirement for generating effective management plans in coastal ecosystems, particularly when determining component habitat structures in marine protected areas. To elucidate the patterns of habitat use by fishes among coral, seagrass, and mangrove habitats, and between natural and transplanted mangroves, visual censuses were conducted semiannually at two sites in the Philippines during September and March 2010–2012. In total, 265 species and 15,930 individuals were recorded. Species richness and abundance of fishes were significantly higher in coral reefs (234 species, 12,306 individuals) than in seagrass (38 species, 1,198 individuals) and mangrove (47 species, 2,426 individuals) habitats. Similarity tests revealed a highly significant difference among the three habitats. Fishes exhibited two different strategies for habitat use, inhabiting either a single (85.6% of recorded species) or several habitats (14.4%). Some fish that utilized multiple habitats, such as Lutjanus monostigma and Parupeneus barberinus, showed possible ontogenetic habitat shifts from mangroves and/or seagrass habitats to coral reefs. Moreover, over 20% of commercial fish species used multiple habitats, highlighting the importance of including different habitat types within marine protected areas to achieve efficient and effective resource management. Neither species richness nor abundance of fishes significantly differed between natural and transplanted mangroves. In addition, 14 fish species were recorded in a 20-year-old transplanted mangrove area, and over 90% of these species used multiple habitats, further demonstrating the key role of transplanted mangroves as a reef fish habitat in this region.     

Distribution status and habitat characteristics of the endangered freshwater fish,Microphysogobio rapidus (Cyprinidae)

To investigate distribution, habitat characteristics, and current conservation status of the endangered endemic species, rapid small gudgeon Microphysogobio rapidus (Cyprinidae), we surveyed a total of 79 sites from the historic records (20 sites) plus additional sites (59 sites) with good habitat conditions, analyzed their sites, and compared them with historic recorded sites to reveal the factors of extinction threats and causes. We found only eight out of 79 sites in the Nam River areas. The habitats were greatly reduced and restricted compared with the historic sites, which mainly cause from habitat modification, such as various types of river renovations at the main stream and tributary streams of the Nakdong River. The present habitats are higher water temperature and more number of fish species than the absent ones, but conductivity, total nitrogen, and number of weir are lower. In addition, the present sites are lower low velocity at pool and higher mean substrate at pool. From this study, we suggest that maintaining good water quality and preventing anthropogenic impacts greatly aid conservation of the M. rapidus in South Korea.     

Habitat use,feeding and territorial behavior of a Brazilian endemic damselfish <Emphasis Type="Italic">Stegastes rocasensis</Emphasis> (Actinopterygii: Pomacentridae)

This paper presents the results of the first field study of the Brazilian endemic damselfish Stegastes rocasensis. Underwater data were collected at the Fernando de Noronha Archipelago (NE Brazil), where four sites were visually assayed for local territorial and feeding behaviors, as well as habitat use. S. rocasensis preferentially fed on turf and macroalgae, and during the afternoon period. Its territory size ranged from 0.26 to 13.86 m² (1.61 ± 0.23 m²). Our findings suggest that adults preferably inhabit shallow, turf-rich areas, while sub-adults and juveniles occupy deeper areas with higher percent cover of invertebrates and hard corals, respectively. With regards to agonistic behaviors, pursuit of intruders was the most frequent reaction recorded (p < 0.001); the highest level of aggression was elicited by territorial herbivores (p < 0.001). Nevertheless, analysis of attacks elicited by different fish species within S. rocasensis’ territory showed that more attacks were launched on conspecifics than on other fish species.     

Using river color to predict Amazonian floodplain forest avifaunas in the world's largest blackwater river basin

Despite the importance of rivers in Amazonian biogeography, avian distribution patterns in river‐created habitats (i.e., floodplain forests) have been sparsely addressed. Here, we explore geographic variation in floodplain forest avifaunas, specifically regarding one of the most striking aspects of the Amazon: the diversity of river “colors” (i.e., types, based on the color of the water). We sampled the avifauna at 30 sites, located in 17 different rivers (nine black‐ and eight whitewater), in the Rio Negro basin, northwestern Brazil. Our sampling comprised ten 15‐min point‐counts per site, distributed every 500–1000 m along the river. We recorded a total of 352 bird species, many of which occurred in both river types. Although bird species richness was similar among rivers, we found significant differences in species composition. Nearly 14 percent of the species were significantly associated with one or the other river type. Most floodplain forest specialists occurred predominantly in whitewater rivers, whereas species that are typically associated with white‐sand habitats occurred in blackwater. Despite significant distinctions between river types, occurrence patterns and levels of habitat association differed among indicator species and may vary in the same species throughout its global distribution. There were also “intermediate” avifauna in some of our sites, suggesting that continuous parameters characterizing river types structure species turnover. The water color‐based classification of Amazonian rivers represents a simple and powerful predictor of the floodplain forest avifauna, offering a stimulating starting point for understanding patterns of floodplain bird distributions and for prioritizing conservation efforts in these overlooked habitats. Abstract in Portuguese is available with online material.     

Characterizing fish–habitat associations in streams as the first step in ecological restoration

Abstract Habitat availability is often regarded as the primary factor that limits population and community recovery in degraded ecosystems, and physical habitat is thus often targeted in restoration. The identification of which habitat(s) to attempt to restore is a critical step in the restoration process, but one for which there is often a paucity of useful information. Here we examine the distribution of fish in three lowland streams in Victoria, Australia, that have been degraded by severe sedimentation. We aim to identify habitats that are associated with high abundances of native fish, and that thus might be appropriate to target in habitat restoration. Associations between native fish abundances and physical habitat characteristics were examined at three spatial scales (among streams, among sites and within sites) to determine the types of habitat to which fish respond, and the scales over which these responses occur. Of the four species of native fish found in the streams, three (Galaxias olidus Günther, Gadopsis marmoratus (Richardson) and Nanoperca australis Günther) showed significant habitat associations at small spatial scales (i.e. within sites). In particular, these species were generally found in deeper water, and in close proximity to cover (typically either coarse or fine woody debris or vegetation). Differences in habitat availability among sites and streams were less influential, except in the case of G. marmoratus, which was completely absent from both the ephemeral streams. Although our results suggest that these species collectively respond to habitat at several spatial scales, fish distributions were allied to the presence of habitat structure at the scale of metres, the smallest spatial scale examined. We hypothesize that fish abundances are currently limited by the low availability of habitat at these small spatial scales. It may therefore be possible to increase fish abundances in these creeks by augmenting the amount of available habitat via stream restoration.     

Local and ecoregion effects on fish assemblage structure in tributaries of the Rio Paraíba do Sul, Brazil

1.  We examined the effects of physical and chemical habitat variables and ecoregions on species occurrence and fish assemblage structure in streams of the Paraíba do Sul basin, in southeast Brazil.
2.  Fish and environmental data were collected from 42 sites on 26 first to fourth order streams (1 : 50 000 map scale) in three ecoregions. The sites occurred in one valley and two plateau ecoregions at altitudes of 40–1080 m and distances of 0.1–188 km from the main channel of the Rio Paraíba do Sul. Physical habitat (substratum, riparian cover, habitat types) and water quality (dissolved oxygen, pH, temperature and conductivity) variables were measured at each sampling site.
3.  A total of 2684 individuals in 16 families and 59 species were recorded.
4.  Ecoregion was a better predictor of the fish assemblage than the other environmental variables, according to the differences between the mean within-class and mean between-class similarities in assemblage data.
5.  Differing landscape characteristics were associated with differing local variables and thereby with differing fish assemblage structures. Riffles, shrub, grass, dissolved oxygen, conductivity and temperature were closely related to fish assemblage structure.
6.  Fish assemblages in sites far from the main river and at higher altitudes also differed from those near the Paraíba do Sul main channel, presumably as a result of differences in connectivity, covarying environmental factors and anthropogenic influence.
7.  These results reinforce the importance of understanding how stream communities are influenced by processes and patterns operating at local and regional scales, which will aid water resource managers to target those factors in their management and rehabilitation efforts.     

Gastropod diversity,distribution and abundance in habitats with and without anthropogenic disturbances in Lake Victoria,Kenya

We investigated freshwater gastropod diversity, abundance and distribution in habitats with and without anthropogenic disturbance in two localities, Ndere in the Winam Gulf and Mbita Point, Lake Victoria, Kenya, from May 2002 to January 2004. A total of 133 984 gastropod specimens belonging to 15 species were recorded, 14 from Mbita and 12 from Ndere. Two species, Ferrissia kavirondica and Cleopatra cridlandi, which were recorded only from undisturbed habitats, could be indicators of least disturbed habitats. Water chemistry did differ between fish landing sites and undisturbed habitats at some sampling times, indicating that differences due to human impact do exist, but these are dependent on periods of calm weather. The study shows that anthropogenic disturbances cause ecological changes that can be exploited by some snail species, especially Biomphalaria choanomphala and Melanoides tuberculata, while other species may not tolerate these changes. In order to protect gastropod diversity and avoid dominance of intermediate hosts, such as B. choanomphala, a management plan for the use of these fish landing sites should be developed. This could include rules on how to dispose of fish remnants and other wastes so as to reduce local eutrophication, thereby reducing risks associated with transmission of potential snail-borne diseases.     

Temporal and Spatial Variations in Fish Catches in the Fly River System in Papua New Guinea and the Possible Effects of the Ok Tedi Copper Mine

Biological monitoring of fishes in the Fly River system in Papua New Guinea has been carried out in relation to the input into the system of mine wastes from the Ok Tedi copper mine. A total of 86 fish species representing 32 families has been recorded from sites in the main river channel since the commencement of monitoring operations in 1983. Catfish in the families Ariidae (16 spp.) and Plotosidae (9 spp.) were the dominant groups overall, although Nematolosa herrings were the most numerous species, forming over 37% of the catch. However, barramundi, Lates calcarifer, comprised the greatest biomass, forming over 30% of the overall catch. Fish catch biomass at most sites showed considerable temporal and spatial variation over the period of sampling. However, significant reductions in biomass, ranging from 65% to 96%, were recorded at most sites in the Ok Tedi, middle and upper Fly. The greatest declines in biomass were recorded in the Ok Tedi at sites closest to the mine, although reductions up to 73% were also recorded at sites in the middle Fly. Barramundi, which formed a high proportion of catch biomass at many sites, particularly in the middle Fly, declined in number at most sites following peak numbers in the early 1990's. The main causative factors involved in the overall declines in fish catches, including both mine-related and non-mine-related factors, are discussed. It is concluded that loss of fish habitat through increased river bed aggradation, due to the input of mined waste rock and tailings, is likely to be the main causal factor. However, other mine-related factors, such as elevated levels of dissolved and particulate copper, and other non-mine-related factors, such as introduced species, may also be involved in declining fish catches.     

Thermal response of demersal and pelagic juvenile fishes from the surf zone during a heat‐wave simulation

Experimental measurements were collected in the laboratory to evaluate the maximum thermal limit and thermal plasticity of Neotropical juvenile fish with different life habitats (demersal and pelagic) from surf zone in response to a “heat‐wave experiment”. Trials were conducted using two temperature acclimations (T_a), including the current average temperature of Southeastern Brazil (T_a: 14 days at 25°C) and the “heat‐wave experiment” (T_a: 14 days at 30°C), simulating a heat‐wave event that occurs when the daily maximum temperature of more than five consecutive days exceeds the average maximum temperature by 5°C. Typical species of the surf zone were used: the demersal White sea catfish (Genidens barbus) and Gulf kingcroaker (Menticirrhus littoralis), and the pelagic fishes Great pompano (Trachinotus goodei) and Long‐fin mullet (Mugil brevirostris). The thermal range and plasticity values for the both life‐habitats species were verified through current and heat‐wave acclimation. The thermal tolerance at high temperatures (CT_max) of these species differed between T_a, habitat and species. Fish showed a species‐specific response to temperature increase, regardless of their habitat even under similar abiotic conditions. However, at the heat‐wave simulation, the demersal fish presented a greater thermal plasticity in relation to the pelagic fish. Despite the higher thermal tolerance when exposed to heat‐wave simulation, all fish species displayed a lower thermal edge safety that is markedly close to their maximum thermal limits.     

Spatial and temporal variation in fish assemblage structure in the Mary River,south-eastern Queensland: the influence of habitat structure

Synopsis Fish assemblages at eleven sites within the Mary River were sampled over a 15 month period by back-pack electrofishing. The study took place during a period of abnormally low flows but included two high flow events one of which approached record levels. Spatial and temporal variation in fish assemblage structure was pronounced. Spatial variation was related to position within the catchment and correlated with habitat attributes related to substrate composition and water depth. The absence of a strong effect due to water velocity was probably due to overall low water flows throughout the study period. Temporal variation at some sites, particularly headwater sites, was correlated with changes in the amount and type of cover available to fishes. Cover was significantly correlated with the mean species richness of each site. Flooding had very little effect on both fish assemblage structure and the physical structure of the habitat but resulted in substantial reduction in the amount and type of cover available to fishes. The flow regime of the Mary River was highly variable and consequently both the timing and duration of high and low flows was unpredictable.