Structural Habitat Heterogeneity Research Articles

Quantifying the impacts of an invasive weed on habitat quality and prey availability for tiger snakes (Notechis scutatus) in urban wetlands

Invasive plants are a threat to natural ecosystems worldwide, with urban wetlands being some of the most susceptible and highly modified environments of all. The tiger snake (Notechis scutatus) is a top predator that persists in urban wetlands of south-western Australia, many of which have been degraded by introduced kikuyu grass (Cenchrus clandestinus). To evaluate the potential impact of kikuyu grass on habitat quality for tiger snakes we quantified the structural features of habitats within wetlands degraded by kikuyu grass and compared them to wetlands with native vegetation. We also examined tiger snake prey availability, assessed predation risk for juvenile snakes using clay models, and measured the thermal quality of the vegetation. Proliferation of kikuyu grass has reduced habitat structural heterogeneity by reducing available bare ground and increasing vegetation density. This homogenisation of habitat structure had little effect on the predation risk for juveniles or the thermal properties of tiger snake shelter sites; however, one key prey species, the motorbike frog, had significantly lower abundance in the most impacted habitat. Habitat types with more structural complexity also offered tiger snakes more stable thermal regimes and lower predation risk. These findings indicate that the current extent of kikuyu grass invasion offers overall similar habitat quality for tiger snakes to native vegetation and may contribute to their persistence in urban wetlands; however, both tiger snakes and their anuran prey may benefit from increased habitat structural complexity.

Bird alpha, beta and functional diversities across three peri-urban woodland stands along an anthropogenic disturbance gradient: is formal protection a guarantee for ecological integrity?

Urban woodlands are significant as refugia for species of wild birds which contribute to provisioning a wide range of ecological services beneficial to nature and to humans. Due to the increasingly small and fragmental structure of these habitats, arising from constant expansion of city infrastructure, these birds are in increasing conservation peril with serious ramifications to the ecological and touristic values they represent. We assessed the alpha, beta and functional diversities of birds in three small woodland stands within the metropolis of Nairobi, Kenya and compared their variations along an anthropogenic disturbance gradient (Low, Moderate; High) based on relative openness, vegetation cover and rate of human visitation. Birds were surveyed using timed species counts during six sampling periods and grouped into three diet-guilds and three vertical foraging strata to assess compositional and assemblage turnover responses to disturbance. Functional overlaps in birds’ ecological roles were also evaluated, and potential candidate species for monitoring disturbance trends were identified. Alpha diversities were determined using Chao1 species estimator, and beta diversity evaluated through the PERMDISP function and main-test Permanova in PERMANOVA+ add-on of Primer-E, to assess species turnovers. Parameters were compared cross-site using general linear models while indicator species were identified using Indval function in R. Significant responses to impacts of anthropogenic disturbance were observed in abundance of birds overall, abundance of upper-stratum species, and in terms of alpha diversity of carnivores but not for other guilds. On the other hand, species turnovers occurred along the disturbance gradient for birds overall, in frugivores and also for both upper- and lower-stratum species. Species functional redundancies, depicting diminished habitat complexities, were detected in the most disturbed site for birds overall, amongst carnivores, all herbivores combined, and among the lower and middle-stratum species, but were not evident for birds overall in the low- or moderately-disturbed sites, nor among upper-stratum bird species. Only two species, both carnivorous, qualified as potential candidates for monitoring disturbance trends across the three woodland sites. These results demonstrate negative effects of anthropogenic disturbance on peri-urban woodland bird assemblage and composition, and greater resilience of taxonomic over functional diversity in response to such disturbance, but also that while formal protection may enhance bird’s community-level functional roles, not all guilds may benefit. Mitigating effects of disturbance requires measures for improving habitat structural heterogeneity to broaden birds’ assemblage composition, diversify their ecosystem roles at all levels and promote their site-based avi-tourism potential.

Management of ecosystems alters vector dynamics and haemosporidian infections

The presence of insect vectors is a key prerequisite for transmission of vector-borne disease such as avian haemosporidians. In general, the effects of land use change on Diptera vectors are not well studied; the response of vectors to forest management depends on vector species, as has been shown previously for the birds. We tested if abundance of insects from different Diptera families and haemosporidian infection are affected through alteration of habitat structural variables (measured by LiDAR) and forest management intensities. We identified higher large-scale variation of female insect abundance in northeastern than in southwestern Germany. Unmanaged forest stands had higher Diptera insect abundances. We found that abundance of female Diptera increased with the amount of forest gaps but decreased in forest plots with more south facing aspect, higher habitat structural heterogeneity, temperature and humidity. We found that haemosporidian infections in Diptera insects increased with increased management intensity and more canopy structural diversity (e.g., amount of edge habitat), but decreased with a denser shrub layer, deeper leaf litter and higher humidity (characteristics for unmanaged forest stands). Although higher forest management intensity decreased vector abundance, the haemosporidian infections in the vectors increased, indicating a significant effect of forest management on disease dynamics.

Increasing ecological multifunctionality during early plant succession

Ecological multifunctionality quantifies the functional performance of various important plant traits and increases with growing structural habitat heterogeneity, number of plant functional traits, and species richness. However, the successional changes in multifunctionality have not been traced so far. We use quantitative plant samples of 1 m2 plots from the first 6 years of initial vegetation dynamics in a German created catchment to infer the temporal changes in plant functional trait space and multifunctionality. Multifunctionality at the plot level was in all study years lower than expected from a random sample of the local pool of potential colonizers and was lowest at intermediate states of succession. In each year species containing a specific set of traits occurred with limited but focused functionality. The observed average low degree of multifunctionality contrasts with recent models predicting a tendency towards maximum multifunctionality during plant community development. However, variability in multifunctionality among plots increased during succession and the respective multifunctionality distribution among plots was increasingly right skewed indicating an excess of plots with relatively high multifunctionality. This relative excess of plots with high multifunctionality might act as an important trigger of community development paving the way for new species and functions to become established.

Exploring structural habitat heterogeneity in sustainable urban drainage systems (SUDS) for urban biodiversity support

Implementation of sustainable urban drainage systems (SUDS) as part of cities’ transition towards climate resilience encompasses unique opportunities to enhance urban nature qualities. SUDS include individual elements integrated in the urban landscape, e.g. rain gardens or wet basins, which are usually constructed with low structural heterogeneity. Structural heterogeneity of habitats is, however, associated with a high degree of biodiversity. This study explored potentials of habitat heterogeneity enhancement in SUDS using structural heterogeneity as a proxy for biodiversity potential. In an interdisciplinary workshop, landscape architects and urban ecologists designed individual SUDS elements with enhanced structural habitat heterogeneity, hereafter bio-SUDS. The redesigned SUDS elements were infiltration trench, curb extension, rain garden, swale, wet basin, and dry basin. To evaluate the bio-SUDS designs, we developed an index of habitat heterogeneity in SUDS based on registration of 45 structurally important components. The conversion from standard SUDS to bio-SUDS entailed substantial increases in structural heterogeneity ranging from 4 to 18 additional components within elements. Larger bio-SUDS elements returned higher index scores, but the relative score increases were substantial for all bio-SUDS and in the range of a factor of 2–5.5. Increased terrain differences, meandering edges, stones, gravel, boulders, deadwood, and brown roof-like vegetation were the components most often employed in the bio-SUDS designs. In conclusion, implementation of bio-SUDS has potential to increase structural habitat heterogeneity thereby providing habitat for biodiversity in urban green areas. The developed bio-SUDS index can assist in future assessments of ecological qualities in designs and real-world SUDS elements.

Patterns of landscape seasonality influence passerine diversity: Implications for conservation management under global change

The importance of environmental heterogeneity for biodiversity across scales is widely recognized in ecological theory and profusely supported by evidence. However, our understanding of the effects of spatiotemporal patterns of landscape functional properties on biodiversity is still rather limited. We examined the relationship between common passerine species richness and ecosystem functioning dynamics, namely seasonality, measured by satellite remote sensing. We focused on rural landscapes of a mountain National Park in Portugal undergoing rapid reshaping from agro-pastoral mosaics to early successional landscapes. We applied multi-model inference to compare the hypothesis of landscape seasonality as a driver of species richness with three competing hypotheses representing structural habitat heterogeneity, disturbance, and availability of food resources. We found support for landscape seasonality and its spatial heterogeneity in explaining passerine richness in mountain rural landscapes. Conversely, no significant support of the remaining hypotheses was found. These results highlight the role of ecosystem functioning variability in space and time. They also stress the importance of considering species-energy relationships for conservation at the landscape level. Specifically, they provide support and guidance to the identification of meaningful functional attributes of the landscape that shape its biodiversity. Our results further demonstrate the utility of remote sensing approaches and products to measure those attributes and follow their trends through time. Spatially-explicit measures of energy variability, such as the functional amplitude between winter and summer retrieved from earth observations, can link global socio-environmental change to species’ responses and support the inclusion of landscape seasonality on conservation and monitoring frameworks.

Constraints to restoring fire and grazing ecological processes to optimize grassland vegetation structural diversity

Extirpation of the ecosystem engineer (bison) and its interaction with fire, coupled with the utilitarian concept of moderate grazing, have contributed to homogenization of grassland habitat in North America. Although cattle may serve as a proxy for bison, combining fire with cattle grazing has been uncommon and to date managers have not always successfully applied cattle and controlled burns as tools to manipulate grassland vegetation heterogeneity and increase habitat diversity. Using an information-theoretic approach, we assessed factors constraining the fire-grazing interaction ecological process to engineer habitat structure of grasslands via patch-burn grazing. We assessed how grazing, fire, and biotic and abiotic features in tallgrass prairie influenced establishment and maintenance of low vegetative structure in burned patches, the positive feedback driving the fire-grazing interaction, and subsequent structural heterogeneity across a pasture. Four pastures were divided into three patches with a different patch burned annually in March/April from 2007 to 2013. Cattle were stocked from light to heavy (1.1–4.4AUM/ha) from May to October (∼150days) with access to the entire pasture. We hypothesized that the exotic C3 grass tall fescue (Schedonorus arundinaceus), lag-time between burning date and the date cattle were put into experimental pastures, and burn date would be the constraining factors. However, the most informative model included stocking rate, date of burn completion, and precipitation. The lightest cattle stocking rate did not establish low vegetative structure in the burn patch, which resulted in the lowest heterogeneity among patches. The heaviest cattle stocking rate established but did not maintain low vegetative structure in the burn patch. The intermediate cattle stocking rate maintained the lowest vegetative structure in the burn patch and the greatest heterogeneity among patches, i.e., the best efficacy of patch-burn grazing to engineer habitat structural heterogeneity. The relationships of stocking rate to burn patch vegetative structure and to landscape heterogeneity were both quadratic and were both optimized at intermediate stocking rate.

Relative Cost/Benefit Trade‐Off Between Cover‐Seeking and Escape Behaviour in an Ancestral Fish: The Importance of Structural Habitat Heterogeneity

AbstractFleeing from predators and moving into protective habitats are two common antipredator behaviours in the animal kingdom. Surprisingly, the relative cost/benefit trade‐off of each behavioural option has rarely been examined empirically. Here, we investigate the interplay between decisions surrounding escape behaviour and rocky microhabitat occupancy in lake sturgeon, Acipenser fulvescens. In high‐risk clear‐water environments, sturgeon responded to danger by evoking an escape response and by seeking cover in rocky microhabitats. However, in low‐risk turbid environments, we found that sturgeon responded to danger by seeking cover in rocky microhabitats, but not fleeing to a significant degree. Cover‐seeking behaviour may therefore be a relatively low‐cost/high‐benefit antipredator strategy. These findings highlight the importance of structural habitat heterogeneity for prey animals in predator‐dominated landscapes.

Bias in the use of broadscale vegetation data in the analysis of habitat selection

Effective conservation efforts often depend on accurate identification of habitat requirements. Studies that identify habitat requirements for animals typically investigate use of structural habitat (vegetation) instead of functional habitat (conditions for biological fitness). The spatial scale of data could bias the match between functional and structural habitat because broadscale structural habitat maps exclude small habitat patches (inclusions) and broadscale location error can bias estimates of habitat use. To investigate how spatial scale affects the match between functional and structural habitat, we backtracked American marten (Martes americana) and fisher (Pekania [formerly Martes] pennanti) movement paths during winter and compared results from selection and tortuosity analyses conducted with broadscale (4 ha) and fine-scale (0.02 ha) structural habitat data. Functional habitat (rest sites and prey kill sites) occurred disproportionately in hemlock–cedar. Finescale structural habitat data detected greater selection and tortuosity within hemlock–cedar by traveling martens, but broadscale structural habitat data did not, which demonstrates that combining fine-scale location data with fine-scale structural habitat data improves the match between functional and structural habitat and understanding of habitat requirements. Selection and tortuosity indexes were poorly correlated, indicating that factors other than structural habitat influenced movement patterns. Within-stand structural habitat heterogeneity is important to martens and fishers, especially when heterogeneity includes mature conifer inclusions within primarily deciduous forests. Broadscale data may identify structural habitat associated with required types, rather than required habitat itself, when functional habitat corresponds with landscape features such as inclusions.

Does richness of Oligochaeta (Annelida) follows a linear distribution with habitat structural heterogeneity in aquatic sediments?

Sediment of aquatic environments supports several local communities, among them aquatic invertebrates. Habitat structural heterogeneity in sediments of aquatic ecosystems is related to a number of factors, including variability in the particle size of the substrate and availability of different food resources, the two structures that comprise the aquatic sediment. These structures are necessary for the stability of zoobenthic community, including the Oligochaeta assemblage, favoring richness as habitat heterogeneity increases. To analyze the habitat structural heterogeneity, we use these following structures: granulometric composition and organic matter composition, and we tested these structures together (habitat heterogeneity) and separately (granulometric and organic matter heterogeneity). This study investigated the relationship between habitat structural heterogeneity and richness of the Oligochaeta assemblage. We hypothesized that there is a positive linear relationship between habitat structural heterogeneity and richness of Oligochaeta, and predicted that the granulometric composition is more important than organic matter composition for increasing richness. There was no linear relationship between Oligochaeta richness and habitat heterogeneity; but the analysis showed an asymmetric triangular dispersion pattern, being granulometric heterogeneity more important than organic matter heterogeneity for richness of Oligochaeta. These results indicated that habitat structural heterogeneity was beneficial for richness of Oligochaeta until certain threshold and after this, the relationship between structural heterogeneity of habitats and richness of Oligochaeta is negative with decrease of diversity due to the filling of interstitial spaces which gradually reduces the ability to colonize these habitats.

Does mollusc shell debris determine patterns of macrofaunal recolonisation on a tidal flat? Experimental evidence from reciprocal transplantations

Mollusc shells introduce complexity and heterogeneity into soft-sediment habitats and might regulate the structure and functioning of benthic assemblages. Two areas of an unvegetated tidal flat located at the mouth of Paranaguá Bay (southern Brazil) largely differ in substrata characteristics and macrofaunal assemblage structure. Sediments of the inner part of the studied site are structurally more complex than those of the peripheral portion, due to high densities of empty shells of the bivalve Anomalocardia brasiliana. The effects of this biogenic matrix on macrofaunal recolonisation during a three-month period were assessed through a reciprocal transplantation experiment of defaunated sediment between shelled and bare habitats. Hypotheses were tested to distinguish between three possible causes of variation: (i) habitat complexity, (ii) intrinsic factors of each area (regardless of the presence/absence of shells), and (iii) their interaction. Habitat structural heterogeneity provided by empty shells does not explain the large differences between macrofauna in shelled and bare habitats. Assemblages that colonised transplanted sediments became similar to those in controls at the area to which they were moved, but differed from those in controls at the area where they originated. The lack of a consistent response of assemblages to the presence of shells can be explained not only by the extremely fast recovery following small-scale experimental disturbances, but also by the dispersal mechanisms related to reproductive traits of the dominant taxa. Changes in assemblages were greatly influenced by the tanaid Monokalliapseudes schubarti, the numerically-dominant species that showed consistent differences among treatments. The reduced dispersal capacity due to the lack of planktonic larval stages associated with high fecundity, continuous reproduction and fast growth were determinant features for the rapid colonisation of defaunated sediments, regardless of habitat complexity. Despite the well-recognised impact of mollusc shell production on soft-sediment habitats, our study showed that its effects are scale-dependent, both spatially and temporally, and might be regulated by the structure of local assemblages and species-specific responses related to dispersal mechanisms.

Why do lizards avoid weeds?

Invasion of native habitats by exotic plants often causes reductions in faunal diversity. However, there is little direct evidence of native fauna actively avoiding invaded habitat and few quantitative studies on the mechanisms underlying such avoidance. We quantified alterations made to the composition and physical structure of an Australian tropical savanna by grader grass (Themeda quadrivalvis); an understudied invasive grass that is associated with reduced faunal abundance and diversity. We found that grader grass profoundly changed the physical structure and floral composition of tropical savanna, forming dense lawn-like monocultures unlike the native savanna. Second, we investigated the habitat preferences of small ectotherms in partially invaded habitat, using a rainbow skink (Carlia schmeltzii) as a model system and discovered that they actively avoided grader grass. Finally, we experimentally tested predictions regarding mechanisms that may have driven the avoidance of grader grass. Predation rates and food availability were not likely the cause of grader grass avoidance, because experiments using models deployed in the field showed that predation rates were higher in native grass, and collection of invertebrates in both habitats indicated that prey availability was similar. However, mesocosm experiments on habitat selection in relation to vegetation structure, along with field measures of available operative environmental temperatures, suggested that small ectotherms probably avoid grader-grass-invaded savanna due to a suboptimal thermal environment and lack of appropriate habitat structural heterogeneity.

Rhabdomyolysis associated with hydroxymethylglutaryl-coenzyme-a reductase inhibitors

Extirpation of the ecosystem engineer (bison) and its interaction with fire, coupled with the utilitarian concept of moderate grazing, have contributed to homogenization of grassland habitat in North America. Although cattle may serve as a proxy for bison, combining fire with cattle grazing has been uncommon and to date managers have not always successfully applied cattle and controlled burns as tools to manipulate grassland vegetation heterogeneity and increase habitat diversity. Using an information-theoretic approach, we assessed factors constraining the fire-grazing interaction ecological process to engineer habitat structure of grasslands via patch-burn grazing. We assessed how grazing, fire, and biotic and abiotic features in tallgrass prairie influenced establishment and maintenance of low vegetative structure in burned patches, the positive feedback driving the fire-grazing interaction, and subsequent structural heterogeneity across a pasture. Four pastures were divided into three patches with a different patch burned annually in March/April from 2007 to 2013. Cattle were stocked from light to heavy (1.1–4.4 AUM/ha) from May to October (∼150 days) with access to the entire pasture. We hypothesized that the exotic C3 grass tall fescue (Schedonorus arundinaceus), lag-time between burning date and the date cattle were put into experimental pastures, and burn date would be the constraining factors. However, the most informative model included stocking rate, date of burn completion, and precipitation. The lightest cattle stocking rate did not establish low vegetative structure in the burn patch, which resulted in the lowest heterogeneity among patches. The heaviest cattle stocking rate established but did not maintain low vegetative structure in the burn patch. The intermediate cattle stocking rate maintained the lowest vegetative structure in the burn patch and the greatest heterogeneity among patches, i.e., the best efficacy of patch-burn grazing to engineer habitat structural heterogeneity. The relationships of stocking rate to burn patch vegetative structure and to landscape heterogeneity were both quadratic and were both optimized at intermediate stocking rate.

Effects of environmental parameters on the distribution of bullhead Cottus gobio with particular consideration of the effects of obstructions

SUMMARY 1. The bullhead Cottus gobio is a small‐sized fish that occurs in stony streams and lakes in Switzerland. Its numbers have declined and its distribution has become narrower over the last 30 years, most probably due to chemical pollution and/or structural habitat deterioration, and at present it is classified as a potentially endangered species. This study evaluated whether chemical parameters and/or structural habitat heterogeneity have an effect on the occurrence and density of bullhead in a river system, in order to assess the potential usefulness of this species as an indicator for these parameters. Ten sites were chosen, where chemical parameters were measured monthly over a period of 2 years. Habitat heterogeneity was assessed with respect to river width and maximum depth. Correlations were drawn between the density of bullhead, the chemical water constituents and the structural habitat diversity. The longitudinal distribution of bullhead in the whole river system was determined and all obstructions with a height of at least 10 cm were investigated. In two sites, the total population density of bullhead older than O+ was assessed using both multiple removal collecting and a mark‐recapture method. Structural habitat diversity appeared not to be important for bullhead, and the only chemical parameter that correlated with the density of this species was dissolved organic carbon. Obstructions with a height of 18–20 cm were impassable for upstream movement of bullhead and thus acted as migration barriers. The longitudinal connectivity of the main river was interrupted by 48 such obstructions, which were all human‐made. Multiple removal collecting of bullhead resulted in small catching probabilities of 0.17 and 0.34. The total estimated population densities of bullhead were high, with 118 and 327 in 100 m2. The mark‐recapture method revealed even higher estimated densities, of 121 and 444 in 100 m2. It is concluded that the bullhead is a useful indicator species for determining to what extent the longitudinal connectivity of a river is interrupted by obstructions. For restoration of rivers, it is recommended that any human‐made obstructions with a height of 20 cm or above are removed, starting at confluences with joining rivers and moving upstream.

Patterns of Seasonal Availability and Habitat Use by Fishes and Decapod Crustaceans in a Southern New Jersey Estuary

We examined the community structure of fish and selected decapod crustaceans and tested for within estuary differences among habitats at depths of 0.6 m to 7.9 m, in Great Bay and Little Egg Harbor in southern New Jersey. Several habitat types were identified a priori (e.g., eelgrass, sea lettuce, and marsh creeks) and sampled by trawl (4.9 m headrope, 19-mm mesh wings, 6.3-mm mesh liner), monthly, from June 1988 through October 1989. Repetitive (n=4) 2-min trawl tows were taken at each habitat type from 13 locations. The fishes and decapod crustaceans collected were typical of other Mid-Atlantic Bight estuaries but varied greatly inseasonal abundance and species. In the years sampled, bay anchovy (Anchoa mitchilli) was the dominant species (50.5% of the total number), followed by spot (Leiostomus xanthurus) (10.7%), Atlantic silverside (Menidia menidia) (9.7%), fourspine stickleback (Apeltes quadracus) (5.9%), blue crab (Callinectes sapidus) (4.6%), and northern pipefish (Syngnathus fuscus) (4.2%). The biota were examined by multi-dimensional scaling (MDS) for habitat associations and “best abiotic predictor” of community structure. Percent silt combined with salinity was the most important abiotic determinant of the faunal distributions among habitats. Temperature was a major factor influencing seasonal occurrence of the biota but had less effect on habitat comparisons. The analysis confirmed the distinct nature of the assemblages associated with the habitats, that is, eelgrass, upper estuary subtidal creeks, channels, and open bay areas. Several species were associated with specific habitats: for example,A. quadracus andS. fuscus with eelgrass, clupeids with subtidal creek stations,L. xanthurus with marsh channels, and black sea bass (Centropristis striata) and spotted hake (Urophycis regia) with sponge-peat habitat. Species richness appeared to be positively related to habitat structural heterogeneity. Thus, the best predictors for these estuarine fish and decapod crustacean assemblages were seasonal temperature, percent silt and salinity combined, and the physical heterogeneity of the habitat.