Regional occupancy in unicellular eukaryotes: a reflection of niche breadth,habitat availability or size‐related dispersal capacity?

1. The distribution patterns of unicellular and multicellular organisms have recently been shown to differ profoundly, with the former probably being mostly cosmopolitan, whereas the latter are mostly restricted to certain regions. However, the within‐region distribution patterns of these two organism groups may be rather similar. 2. We predicted that the degree of regional occupancy in unicellular eukaryotes would be related to niche characteristics, dispersal ability and size, as has been found previously for multicellular organisms. The niche characteristics we considered were niche position, that measures marginality in species habitat distribution, and niche breadth, that measures amplitude in species habitat distribution. Niche characteristics were determined using Outlying Mean Index (OMI) analysis. 3. We found that the regional occupancy in our model group of unicellular eukaryotes, stream diatoms, was primarily a reflection of the niche position of a species or, more generally, habitat availability. Thus, non‐marginal species (i.e. species that occupied common habitat conditions across the region) tended to be more widely distributed than marginal species (i.e. species that were restricted to a limited range of rare habitat conditions). This finding was further supported by the general linear model, with niche position, niche breadth, maximum size and attachment mode as explanatory variables: niche position was by far the most important variable accounting for variability in regional occupancy, with significant amounts of additional variation related to niche breadth and maximum size of diatoms. 4. Thus, the degree of regional occupancy among unicellular eukaryotes may be primarily governed by habitat availability, supporting former findings for multicellular organisms.     

Specialization and habitat: spatial and environmental effects on abundance and genetic diversity of forest generalist and specialist Carabus species

Habitat specialist species are supposed to be more susceptible to variations in local environmental characteristics than generalists. To test this hypothesis, we conducted a comparative analysis on abundance and genetic diversity of forest carabids differing in their habitat requirements. Four species were sampled in forests characterized by abiotic, landscape and biotic environmental variables. A statistical framework based on canonical correspondence analysis was used for one habitat generalist and one habitat specialist species to determine the relative contribution of environmental variables in structuring inter- and intrapopulational genetic diversity depicted by microsatellites. Our results showed that sympatric species differed in their sensitivity to environmental variables. The same variables were found to be important in analyses of abundance and genetic data. However, specialization was not related to a greater sensitivity to local environmental characteristics. The strong impact of spatial variables on genetic data suggested that genetic variation among populations would largely reflect the response of individual species to dispersal opportunities more than the effect of habitat quality.     

Competition,niche specialization and the evolution of brain size in the genus Peromyscus

Previous studies of relative brain size in mammals have suggested an association with complex habitats and with low reproductive rate. In order to examine the causal relationships more thoroughly, a detailed examination of relative brain size variation in the genus Peromyscus was undertaken. Endocranial volumes were used to estimate brain weight for 32 species including 161 subspecies, and relative brain size calculated as the species deviation from the allometric relationship between brain and body size. The intrageneric allometric coefficient was higher than most values previously reported from low taxonomic levels, but intraspecific coefficients were generally lower than this. Island species, and relict species isolated on mountain tops, which may be ecological ‘islands’, had consistently small relative brain sizes, but peninsular species were large brained. Among the remaining species there were significant correlations between litter size and relative brain size, and between the number of competitor species and relative brain size. Species with many competitor species have relatively large brains and small litters. It is concluded that the nature of the geographical distribution, the pattern of species formation and habitat complexity all influence relative brain size in existing forms.     

Competition, niche specialization and the evolution of brain size in the genus Peromyscus

Previous studies of relative brain size in mammals have suggested an association with complex habitats and with low reproductive rate. In order to examine the causal relationships more thoroughly, a detailed examination of relative brain size variation in the genus Peromyscus was undertaken. Endocranial volumes were used to estimate brain weight for 32 species including 161 subspecies, and relative brain size calculated as the species deviation from the allometric relationship between brain and body size. The intrageneric allometric coefficient was higher than most values previously reported from low taxonomic levels, but intraspecific coefficients were generally lower than this.
Island species, and relict species isolated on mountain tops, which may be ecological 'islands', had consistently small relative brain sizes, but peninsular species were large brained. Among the remaining species there were significant correlations between litter size and relative brain size, and between the number of competitor species and relative brain size. Species with many competitor species have relatively large brains and small litters. It is concluded that the nature of the geographical distribution, the pattern of species formation and habitat complexity all influence relative brain size in existing forms.     

Niche partitioning and niche filtering jointly mediate the coexistence of three closely related spider species (Araneae,Philodromidae)

1. Several mechanisms can mediate the coexistence of species, such as the neutral dynamic, niche filtering and niche partitioning. The present study investigated which of these mechanisms mediate the coexistence of closely related spider species at the scale of one locality. 2. The coexistence of three spider species of the genus Philodromus (Philodromidae) (Philodromus albidus, Philodromus aureolus and Philodromus cespitum) was studied. The study area comprised three habitat types, including a deciduous forest, a scrub and a plum tree stand. The spatial niche properties of the Philodromus species were explored by comparing their micro/macrohabitat preferences. The natural diet of these species was analysed. Laboratory experiments involving prey acceptance were conducted to investigate the trophic niche properties. The species' phenologies were studied to compare their temporal niches. 3. The philodromids had differentiated their trophic and habitat niches. The coexistence mechanisms were therefore assessed by studying the relationships between niche overlaps and the pairwise product of relative abundances. A negative relationship was observed between functional niche overlaps and the product of relative abundances, whereas a positive relationship was observed between spatial niche overlaps and the product of relative abundances. 4. The present results suggest that different mechanisms influence different niche dimensions. Niche partitioning influenced functional niches, whereas niche filtering influenced spatial niches. The results also suggest that the filtering process in one dimension was facilitated by niche differentiation in another dimension.     

The niche variation hypothesis and the evolution of colour polymorphism in birds: a comparative study of owls, nightjars and raptors

We studied the evolution of colour polymorphism in diurnal raptors, owls and nightjars, the avian taxa in which this trait is most widespread, in relation to species ecological niche width and diet. Two main mechanisms have been put forward to explain the maintenance of polymorphism, namely apostatic selection and disruptive selection. The niche variation hypothesis states that species with broader ecological niches should be more variable compared with those with narrow niches because of the action of disruptive selection; the apostatic selection hypothesis conversely suggests that intraspecific colour variation should be promoted in predators by prey forming an avoidance image for the more common colour morph. Our aim was to determine if colour polymorphism occurrence was associated with broad ecological niches as predicted by the niche variation hypothesis, or with predation on intelligent and sharp‐sighted prey as predicted by the avoidance image hypothesis. Pairwise comparisons were made between pairs of closely related species differing in variables expected to influence the occurrence of polymorphism. We found that polymorphic species of all three groups showed wider and more continuous distribution ranges, frequented many different habitats, both open and closed, and lived in seasonally alternating dry/wet climates. Polymorphic species were more migratory compared with monomorphic ones, and they showed an activity pattern covering both day and night. Conversely, colour polymorphism was not higher in species preying on birds and mammals. All these findings support the hypothesis that colour polymorphism evolved in bird species with wider niche breadth and not in species preying on intelligent prey. Therefore, we propose that disruptive selection may be the main mechanism maintaining colour polymorphism in these bird groups by favouring different morphs in different environmental conditions. © 2004 The Linnean Society of London, Biological Journal of the Linnean Society, 2004, 82 , 237–248.     

Responses of 'resistant' vertebrates to habitat loss and fragmentation: the importance of niche breadth and range boundaries

Abstract. An ability to predict species' sensitivities to habitat loss and fragmentation has important conservation implications, and numerous hypotheses have been proposed to explain interspecific differences observed in human-dominated landscapes. We used occupancy data collected on 32 species of vertebrates (16 mammals and 16 amphibians) in an agricultural landscape of Indiana, USA, to compare hypotheses that focus on different causal mechanisms underlying interspecific variation in responses to habitat alteration: (1) body size; (2) morphology and development; (3) behaviour; (4) niche breadth; (5) proximity to range boundary; and multiple-process models combining main effects and interactions of hypotheses (1)–(2) and (4)–(5). The majority of habitat alteration occurred over a century ago and coincided with extinction of several species; thus, our study dealt only with variation in responses of extant species that often are considered 'resistant' to human modifications of native habitat. Corrected Akaike scores and Akaike weights provided strongest support for models incorporating niche breadth and proximity to range boundary. Measures of dietary and habitat breadth obtained from the literature were negatively correlated with sensitivity to habitat alteration. Additionally, greater sensitivity was observed for species occurring at the periphery of their geographical ranges, especially at northern or western margins. Body size, morphological, developmental and behavioural traits were inferior predictors of tolerance to fragmentation for the species and landscape we examined. Our findings reinforce the importance of niche breadth as a predictor of species' responses to habitat alteration. They also highlight the importance of viewing the effects of habitat loss and fragmentation in a landscape within a biogeographical context that considers a species' level of adaptation to local environmental conditions.     

Niche dynamics of Memecylon in Sri Lanka: Distribution patterns,climate change effects,and conservation priorities

Recent climate projections have shown that the distribution of organisms in island biotas is highly affected by climate change. Here, we present the result of the analysis of niche dynamics of a plant group, Memecylon, in Sri Lanka, an island, using species occurrences and climate data. We aim to determine which climate variables explain current distribution, model how climate change impacts the availability of suitable habitat for Memecylon, and determine conservation priority areas for Sri Lankan Memecylon. We used georeferenced occurrence data of Sri Lankan Memecylon to develop ecological niche models and assess both current and future potential distributions under six climate change scenarios in 2041–2060 and 2061–2080. We also overlaid land cover and protected area maps and performed a gap analysis to understand the impacts of land‐cover changes on Memecylon distributions and propose new areas for conservation. Differences among suitable habitats of Memecylon were found to be related to patterns of endemism. Under varying future climate scenarios, endemic groups were predicted to experience habitat shifts, gains, or losses. The narrow endemic Memecylon restricted to the montane zone were predicted to be the most impacted by climate change. Projections also indicated that changes in species’ habitats can be expected as early as 2041–2060. Gap analysis showed that while narrow endemic categories are considerably protected as demonstrated by their overlap with protected areas, more conservation efforts in Sri Lankan forests containing wide endemic and nonendemic Memecylon are needed. This research helped clarify general patterns of responses of Sri Lankan Memecylon to global climate change. Data from this study are useful for designing measures aimed at filling the gaps in forest conservation on this island.     

The genetic architecture of a niche: variation and covariation in host use traits in the Colorado potato beetle

The genetic basis of host plant use by phytophagous insects can provide insight into the evolution of ecological niches, especially phenomena such as specialization and phylogenetic conservatism. We carried out a quantitative genetic analysis of multiple host use traits, estimated on five species of host plants, in the Colorado potato beetle, Leptinotarsa decemlineata (Coleoptera: Chrysomelidae). Mean values of all characters varied among host plants, providing evidence that adaptation to plants may require evolution of both behavioral (preference) and post-ingestive physiological (performance) characteristics. Significant additive genetic variation was detected for several characters on several hosts, but not in the capacity to use the two major hosts, a pattern that might be caused by directional selection. No negative genetic correlations across hosts were detected for any 'performance' traits, i.e. we found no evidence of trade-offs in fitness on different plants. Larval consumption was positively genetically correlated across host plants, suggesting that diet generalization might evolve as a distinct trait, rather than by independent evolution of feeding responses to each plant species, but several other traits did not show this pattern. We explored genetic correlations among traits expressed on a given plant species, in a first effort to shed light on the number of independent traits that may evolve in response to selection for host-plant utilization. Most traits were not correlated with each other, implying that adaptation to a novel potential host could be a complex, multidimensional 'character' that might constrain adaptation and contribute to the pronounced ecological specialization and the phylogenetic niche conservatism that characterize many clades of phytophagous insects.     

Lizard habitat partitioning on islands: the interaction of local and landscape scales

Aim This study addresses how species resolve environmental differences into biological habitats at multiple, interacting spatial scales. How do patterns of local habitat use change along an elevation gradient? How do patterns of local habitat partitioning interact with partitioning at a landscape scale? Location Northern and southern Lesser Antilles islands, West Indies. Methods We document how Anolis Daudin, 1802 lizards partition habitat locally at sites along a landscape‐scale elevation gradient. We examine habitat partitioning both with and without interspecific interactions in the predominately flat northern Lesser Antilles islands and in the more mountainous southern islands. Results Anoles partition local habitat along perch‐height and microclimate axes. Northern‐group sympatric anoles partition local habitat by perch height and have overlapping distributions at the landscape scale. Southern‐group sympatric anoles partition local habitat by microclimate and specialize in particular habitats at the landscape scale. In both the northern and southern groups, species use different perch heights and microclimates only in areas of species overlap along the elevation gradient. Main conclusions We demonstrate the interaction between local‐ and landscape‐scale habitat partitioning. In the case of microclimate partitioning, the interaction results from the use of thermal physiology to partition habitat at multiple scales. This interaction prompts the question of whether habitat partitioning developed ‘local‐out’ or ‘landscape‐in’. We pose this dichotomy and present a framework for its resolution.     

Genetic response to human‐induced habitat changes in the marine environment: A century of evolution of European sprat in Landvikvannet,Norway

Habitat changes represent one of the five most pervasive threats to biodiversity. However, anthropogenic activities also have the capacity to create novel niche spaces to which species respond differently. In 1880, one such habitat alterations occurred in Landvikvannet, a freshwater lake on the Norwegian coast of Skagerrak, which became brackish after being artificially connected to the sea. This lake is now home to the European sprat, a pelagic marine fish that managed to develop a self‐recruiting population in barely few decades. Landvikvannet sprat proved to be genetically isolated from the three main populations described for this species; that is, Norwegian fjords, Baltic Sea, and the combination of North Sea, Kattegat, and Skagerrak. This distinctness was depicted by an accuracy self‐assignment of 89% and a highly significant F _ST between the lake sprat and each of the remaining samples (average of ≈0.105). The correlation between genetic and environmental variation indicated that salinity could be an important environmental driver of selection (3.3% of the 91 SNPs showed strong associations). Likewise, Isolation by Environment was detected for salinity, although not for temperature, in samples not adhering to an Isolation by Distance pattern. Neighbor‐joining tree analysis suggested that the source of the lake sprat is in the Norwegian fjords, rather than in the Baltic Sea despite a similar salinity profile. Strongly drifted allele frequencies and lower genetic diversity in Landvikvannet compared with the Norwegian fjords concur with a founder effect potentially associated with local adaptation to low salinity. Genetic differentiation (F _ST) between marine and brackish sprat is larger in the comparison Norway‐Landvikvannet than in Norway‐Baltic, which suggests that the observed divergence was achieved in Landvikvannet in some 65 generations, that is, 132 years, rather than gradually over thousands of years (the age of the Baltic Sea), thus highlighting the pace at which human‐driven evolution can happen.     

The experimental evolution of specialists,generalists, and the maintenance of diversity

Environmental heterogeneity may be a general explanation for both the quantity of genetic variation in populations and the ecological niche width of individuals. To evaluate this hypothesis, I review the literature on selection experiments in heterogeneous environments. The niche width usually – but not invariably – evolves to match the amount of environmental variation, specialists evolving in homogeneous environments and generalists evolving in heterogeneous environments. The genetics of niche width are more complex than has previously been recognized, particularly with respect to the magnitude of costs of adaptation and the putative constraints on the evolution of generalists. Genetic variation in fitness is more readily maintained in heterogeneous environments than in homogeneous environments and this diversity is often stably maintained through negative frequency‐dependent selection. Moreover environmental heterogeneity appears to be a plausible mechanism for at least two well‐known patterns of species diversity at the landscape scale. I conclude that environmental heterogeneity is a plausible and possibly very general explanation for diversity across the range of scales from individuals to landscapes.     

The distribution of ground spiders (Araneae, Gnaphosidae) along the altitudinal gradient of Crete, Greece: species richness, activity and altitudinal range

Aim To study the altitudinal variation of ground spiders (Araneae, Gnaphosidae) of Crete, Greece, as far as species composition, species richness, activity and range of distribution are concerned. Location Altitudinal zones (0–2400 m) along the three main mountain massifs of the island of Crete. Methods Thirty‐three sampling sites were located from 0 to 2400 m a.s.l. on Crete, and sampled using pitfall traps. Material from the high‐activity period of Gnaphosidae (mid‐spring to mid‐autumn) was analysed. Sampling sites were divided into five altitudinal zones of 500 m each. Statistical analysis involved univariate statistics (anova ) and multivariate statistics, such as multidimensional scaling (MDS) and cluster analysis (UPGMA) using binomial data of species presence or absence. Results Species richness declines with altitude and follows a hump‐shaped pattern. The activity pattern of the family, as a whole, is not correlated with altitude and is highly species‐specific. In the highest zone, both species richness and activity decline dramatically. The altitudinal range of species distribution increases with altitude. On the Cretan summits live highly tolerant lowland species and isolated residents of the high mountains of Crete. Two different patterns of community structure are recorded. Main conclusions Communities of Gnaphosidae on Crete present two distinct structures following the altitudinal gradient, these being separated by a transitional zone between 1600 and 2000 m. This study supports previous results which show a hump‐shaped decline in species richness of Gnaphosidae along altitudinal gradients, leading to a peak at 400–700 m, where an optimum of environmental factors exists. This makes this zone the meeting point of the often opportunistic lowland species with the older and most permanent residents of the island. Rapoport's rule on the positive correlation of the altitudinal range of species distributions with altitude is also supported. The high activity recorded for the species that persist on the high mountains of Crete is indicative of a tolerant arachnofauna, and is considered to result from relaxation of competitive interactions with other species. This is related to a reduction in species numbers, shortening of the activity period on high mountains and the unique presence of high mountain species that thrive only there. As shown in our study, strategies to cope with altitude are species‐specific. Therefore, there cannot exist one single model to describe how animals react to the change in altitude, even under the same environmental conditions.     

Diversity and endemism in deglaciated areas: ploidy,relative genome size and niche differentiation in the Galium pusillum complex (Rubiaceae) in Northern and Central Europe

Background and Aims

Plants endemic to areas covered by ice sheets during the last glaciation represent paradigmatic examples of rapid speciation in changing environments, yet very few systems outside the harsh arctic zone have been comprehensively investigated so far. The Galium pusillum aggregate (Rubiaceae) is a challenging species complex that exhibits a marked differentiation in boreal parts of Northern Europe. As a first step towards understanding its evolutionary history in deglaciated regions, this study assesses cytological variation and ecological preferences of the northern endemics and compares the results with corresponding data for species occurring in neighbouring unglaciated parts of Central and Western Europe.

Methods

DNA flow cytometry was used together with confirmatory chromosome counts to determine ploidy levels and relative genome sizes in 1158 individuals from 181 populations. A formalized analysis of habitat preferences was applied to explore niche differentiation among species and ploidy levels.

Key Results

The G. pusillum complex evolved at diploid and tetraploid levels in Northern Europe, in contrast to the high-polyploid evolution of most other northern endemics. A high level of eco-geographic segregation was observed between different species (particularly along gradients of soil pH and competition) which is unusual for plants in deglaciated areas and most probably contributes to maintaining species integrity. Relative monoploid DNA contents of the species from previously glaciated regions were significantly lower than those of their counterparts from mostly unglaciated Central Europe, suggesting independent evolutionary histories.

Conclusions

The aggregate of G. pusillum in Northern Europe represents an exceptional case with a geographically vicariant and ecologically distinct diploid/tetraploid species endemic to formerly glaciated areas. The high level of interspecific differentiation substantially widens our perception of the evolutionary dynamics and speciation rates in the dramatically changing environments of Northern Europe.     

Linking ecology,morphology, and metabolism: Niche differentiation in sympatric populations of closely related species of the genus Littorina (Neritrema)

Divergence of ecological niches in phylogenetically closely related species indicates the importance of ecology in speciation, especially for sympatric species are considered. Such ecological diversification provides an advantage of alleviating interspecies competition and promotes more efficient exploitation of environmental resources, thus being a basis for ecological speciation. We analyzed a group of closely related species from the subgenus Neritrema (genus Littorina, Caenogastropoda) from the gravel‐bouldery shores. In two distant sites at the Barents and Norwegian Sea, we examined the patterns of snail distribution during low tide (quantitative sampling stratified by intertidal level, presence of macrophytes, macrophyte species, and position on them), shell shape and its variability (geometric morphometrics), and metabolic characteristics (metabolomic profiling). The studied species diversified microbiotopes, which imply an important role of ecological specification in the recent evolution of this group. The only exception to this trend was the species pair L. arcana / L. saxatilis, which is specifically discussed. The ecological divergence was accompanied by differences in shell shape and metabolomic characteristics. Significant differences were found between L. obtusata versus L. fabalis and L. saxatilis / L. arcana versus L. compressa both in shell morphology and in metabolomes. L. saxatilis demonstrated a clear variability depending on intertidal level which corresponds to a shift in conditions within the occupied microhabitat. Interestingly, the differences between L. arcana (inhabiting the upper intertidal level) and L. compressa (inhabiting the lower one) were analogous to those between the upper and lower fractions of L. saxatilis. No significant level‐dependent changes were found between the upper and lower fractions of L. obtusata, most probably due to habitat amelioration by fucoid macroalgae. All these results are discussed in the contexts of the role of ecology in speciation, ecological niche dynamics and conservatism, and evolutionary history of the Neritrema species.     

The evolution of vocalisations in the genus Corvus: effects of phylogeny, morphology and habitat

In this study we investigated the phylogenetic, morphological and ecological factors affecting the caw calls of 28 species of the genus Corvus, spanning the worldwide range of the taxon. The three phylogeographic groups identified by Goodwin (1986, Crows of the World, British Museum (Natural History), St Edmudsbury Press, Bury St Edmunds), i.e. the American stock, the Palearctic-African stock and the Oriental-Australian stock, were differentiated by some of the acoustic features of their calls, suggesting that historical factors may have played an important role in the evolution of vocalisations in this group. To assess the effects of morphology (body size and bill length) and environment (open vs. closed habitat) and to simultaneously take into account the phylogenetic effects, we used the phylogenetically independent contrast method. This manner of analysis revealed that body size was important in shaping the acoustic attributes of the caw call, as it influenced two temporal and two spectral variables, whereas the effect of bill length was far weaker. Notably, our results did not confirm the negative correlation between call frequency and body size that resulted in a phylogeny-free analysis of the same data in many studies on passerines. Larger Corvus species, in fact, utter calls with higher fundamental frequency than those of smaller species. Hence, these results show that incorporating phylogeny in analyses can substantially alter the conclusions reached by studies carried out with non-phylogenetic approaches. The acoustic environment, considered one of the most important forces driving the evolution of vocalisations in passerines, slightly influenced only two acoustic parameters in the Corvusgenus, call fundamental frequency and duration of pulsed units, both of which increased in the calls of forest species.     

Influence of volcanic activity on the population genetic structure of Hawaiian Tetragnatha spiders: fragmentation, rapid population growth and the potential for accelerated evolution

Volcanic activity on the island of Hawaii results in a cyclical pattern of habitat destruction and fragmentation by lava, followed by habitat regeneration on newly formed substrates. While this pattern has been hypothesized to promote the diversification of Hawaiian lineages, there have been few attempts to link geological processes to measurable changes in population structure. We investigated the genetic structure of three species of Hawaiian spiders in forests fragmented by a 150-year-old lava flow on Mauna Loa Volcano, island of Hawaii: Tetragnatha quasimodo (forest and lava flow generalist), T. anuenue and T. brevignatha (forest specialists). To estimate fragmentation effects on population subdivision in each species, we examined variation in mitochondrial and nuclear genomes (DNA sequences and allozymes, respectively). Population subdivision was higher for forest specialists than for the generalist in fragments separated by lava. Patterns of mtDNA sequence evolution also revealed that forest specialists have undergone rapid expansion, while the generalist has experienced more gradual population growth. Results confirm that patterns of neutral genetic variation reflect patterns of volcanic activity in some Tetragnatha species. Our study further suggests that population subdivision and expansion can occur across small spatial and temporal scales, which may facilitate the rapid spread of new character states, leading to speciation as hypothesized by H. L. Carson 30 years ago.