Phylogeographical patterns and genetic diversity in three species of Eurasian boreal forest beetles

Many species that occur in formerly glaciated areas of Fennoscandia have reached their current ranges from glacial refugial areas in Eurasia. Little is known of the refugia and postglacial colonization routes of insect species that are confined to boreal forests. Here, we investigate the phylogeography of three species of saproxylic beetles distributed across Eurasia: two rare boreal forest specialists, Pytho kolwensis and Pytho abieticola , and a common, less specialized species, Pytho depressus . In all species, there were two well-defined haplotype clades based on 645 bp of cytochrome oxidase subunit I gene sequence. In each species one clade was found only in China. The other clade occurred from China to north-western Europe in both P. kolwensis and P. depressus , but was apparently absent from China in P. abieticola. In spite of common phylogeographical patterns, the distribution of genetic variation differed markedly between the three species. In P. kolwensis , a highly-threatened species in old-growth forests in Fennoscandia, there was an extremely low level of genetic variation throughout Eurasia. One common haplotype, represented by 86% of the samples, dominated in all sampling localities. Levels of genetic variation were higher in both P. abieticola and P. depressus , with 31% and 58%, respectively, of the samples representing a unique haplotype. In each species, relationships between haplotypes were not well resolved, and haplotypes from one sampling locality were generally not clustered in either Neighbour-joining trees or statistical parsimony networks. These patterns in the distribution of genetic variation can be attributed to differences in the species' population sizes, ecologies, glacial refugial areas, and postglacial colonization dynamics.  © 2007 The Linnean Society of London, Biological Journal of the Linnean Society , 2007, 91 , 267–279.     

Spatial structure and genetic diversity of three tropical tree species with different habitat preferences within a natural forest

Analyses of the spatial distribution pattern, spatial genetic structure and genetic diversity were carried out using a 33-ha plot in a hill dipterocarp forest for three dipterocarps with different habitat preferences, i.e. Shorea curtisii on the ridges, Shorea leprosula in the valleys and Shorea macroptera both on the ridges and in the valleys. The significant spatial aggregation in small-diameter trees of all the three species was explained by limited seed dispersal. At the large-diameter trees, only S. macroptera showed random distribution and this might further prove that S. macroptera is habitat generalist, whilst S. curtisii and S. leprosula are habitat specific. The levels of genetic diversity estimated based on five microsatellite loci were high and comparable in all the three studied species. As the three studied species reproduced mainly through outcrossing, the observed high levels of genetic diversity might support the fact that the plant mating system can be used as guideline to infer the levels of genetic diversity, regardless of whether the species is habitat specific or habitat generalist. The lack of spatial genetic structure but significant aggregation in the small-diameter trees of all the three species might indicate limited seed dispersal but extensive pollen flow. Hence, if seed dispersal is restricted but pollen flow is extensive, significant spatial aggregation but no spatial genetic structure will be observed at the small-diameter trees, regardless of whether the species is habitat specific or habitat generalist. The inferred extensive pollen flow might indicate that energetic pollinators are involved in the pollination of Shorea species in the hill dipterocarp forests.     

Spatio-environmental determinants of the genetic structure of three steppe species in a highly fragmented landscape

Habitat fragmentation can lead to substantial genetic depletion. As a consequence, restoration schemes often involve the introduction of propagules into isolated plant populations to improve genetic diversity. To avoid introducing maladapted seed material, such measures need to account for landscape genetic processes. However, surprisingly little is known as to whether different species within a distinct fragmented ecosystem respond similarly or idiosyncratically to eco-geographical variation.Using AFLP markers, we studied the population genetic structure in three species of the highly fragmented Kulunda steppe (South Siberia): Adonis villosa, Jurinea multiflora and Paeonia hybrida. In each population, we conducted a vegetation survey. We performed Mantel tests and an RDA approach to investigate how genetic structure was affected by three spatio-environmental variables: spatial distance, floristic composition and climate.Despite strong fragmentation, genetic diversity was moderate (A. villosa, J. multiflora) to high (P. hybrida), while differentiation was weak (A. villosa) to moderate (P. hybrida, J. multiflora). Mantel tests showed that spatial distance correlated with genetic distance in A. villosa and P. hybrida. Floristic composition was significantly associated with genetic differentiation in A. villosa. Climate did not have an effect on genetic structure in any species.All three species are long-lived, which may contribute to explaining why genetic effects of recent fragmentation are still limited. We highlight that floristic composition can be a powerful predictor of population differentiation in species that show rather stable conditions in their recent population histories (e.g. A. villosa). This can have important implications for identifying source populations where restoration actions involve the (re)introduction of propagules. In contrast, for P. hybrida and J. multiflora, we could not identify deterministic drivers of differentiation. We advocate that future studies should aim at disentangling the interactive effects of varying life cycles, eco-evolutionary population histories and spatio-environmental heterogeneity in fragmented landscapes.     

Patch isolation and shape predict plant functional diversity in a naturally fragmented forest

Aims It is known that taxonomic diversity can be predicted by the spatial configuration of the habitat, in particular by its area and degree of isolation. However, taxonomic diversity is a poor predictor of ecosystem functioning. While functional diversity is strongly linked to the functionality and stability of ecosystems, little is known about how changes in the spatial configuration of the habitat affect functional diversity. In this study, we evaluated whether the spatial configuration of forest patches predicts the functional diversity of plants in a fragmented forest.Methods Five functional leaf traits (leaf dry matter content, leaf punch force, specific leaf area, leaf size and leaf thickness) were measured for 23 dominant plant species in 20 forest patches in a naturally fragmented forest on the Yucatan Peninsula. Abundance-weighted multivariate and individual trait metrics of functional diversity were calculated and correlated with size, degree of isolation and the shape of forest patches.Important findings Patch shape was negatively correlated with multivariate and individual trait (leaf dry matter content and leaf size) metrics of functional diversity. Patch isolation measures were also negatively correlated with individual trait (leaf dry matter content, leaf punch force and leaf size) metrics of functional diversity. In other words, greater patch shape irregularity and isolation degree impoverish plant functional variability. This is the first report of the negative effects of patch shape irregularity and isolation on the functional diversity of plant communities in a forest that has been fragmented for a long time.     

Species and genetic diversity are not congruent in fragmented dry grasslands

Biological diversity comprises both species diversity (SD) and genetic diversity (GD), and it has been postulated that both levels of diversity depend on similar mechanisms. Species‐genetic diversity correlations (SGDC) are therefore supposed to be generally positive. However, in contrast to theory, empirical data are contradictory. Furthermore, there is a pronounced lack of multispecies studies including also the ecological factors potentially driving species and genetic diversity. We analyzed the relationship between the species diversity of dry grasslands and the genetic diversity of several dry grassland plant species, therefore, in the context of habitat fragmentation and habitat conditions. Our study revealed a lack of correlation between species and genetic diversity. We demonstrated previously that SD mainly depends on habitat conditions (vegetation height and cover of litter), whereas GD is significantly affected by habitat fragmentation (distance to the nearest dry grassland in 1830 and connectivity in 2013). This seems to be the main reason why SD and GD are not congruent in fragmented grasslands. Our results support, hence, the observation that positive SGDCs can mainly be found in natural, island‐like study systems in equilibrium and at similar levels of heterogeneity. In fragmented dry grassland ecosystems, which differ in heterogeneity, this state of equilibrium may not have been reached mitigating the positive relationship between SD and GD. From our study, it can be concluded that in fragmented dry grasslands, the protection of SD does not necessarily ensure the conservation of GD.     

Changes in tree reproductive traits reduce functional diversity in a fragmented Atlantic forest landscape

Functional diversity has been postulated to be critical for the maintenance of ecosystem functioning, but the way it can be disrupted by human-related disturbances remains poorly investigated. Here we test the hypothesis that habitat fragmentation changes the relative contribution of tree species within categories of reproductive traits (frequency of traits) and reduces the functional diversity of tree assemblages. The study was carried out in an old and severely fragmented landscape of the Brazilian Atlantic forest. We used published information and field observations to obtain the frequency of tree species and individuals within 50 categories of reproductive traits (distributed in four major classes: pollination systems, floral biology, sexual systems, and reproductive systems) in 10 fragments and 10 tracts of forest interior (control plots). As hypothesized, populations in fragments and control plots differed substantially in the representation of the four major classes of reproductive traits (more than 50% of the categories investigated). The most conspicuous differences were the lack of three pollination systems in fragments--pollination by birds, flies and non-flying mammals--and that fragments had a higher frequency of both species and individuals pollinated by generalist vectors. Hermaphroditic species predominate in both habitats, although their relative abundances were higher in fragments. On the contrary, self-incompatible species were underrepresented in fragments. Moreover, fragments showed lower functional diversity (H' scores) for pollination systems (-30.3%), floral types (-23.6%), and floral sizes (-20.8%) in comparison to control plots. In contrast to the overwhelming effect of fragmentation, patch and landscape metrics such as patch size and forest cover played a minor role on the frequency of traits. Our results suggest that habitat fragmentation promotes a marked shift in the relative abundance of tree reproductive traits and greatly reduces the functional diversity of tree assemblages in fragmented landscapes.     

Assessing the effects of elephant foraging on the structure and diversity of an Afrotropical forest

African forest elephants (Loxodonta cyclotis) are ecosystem engineers that browse and damage large quantities of vegetation during their foraging and movement. Though elephant trail networks and clearings are conspicuous features of many African forests, the consequences of elephant foraging for forest structure and diversity are poorly documented. In this study in northeastern Gabon, we compare stem size, stem density, proportional damage, species diversity, and species relative abundance of seedlings and saplings in the vicinity of seven tree species that produce elephant-preferred fruits (“elephant trees”) relative to control trees that do not. Across 34 survey trees, with a combined census area of 2.04 ha, we recorded data on 26,128 woody stems in three sizes classes. Compared with control trees, the area around elephant trees had the following: (a) a significantly greater proportion of damaged seedlings and a marginally greater proportion of damaged saplings (with 82% and 24% greater odds of damage, respectively); (b) no significant difference in stem density or species diversity; and (c) a significantly greater relative abundance of seedlings of elephant tree species. Increasing distance away from focal elephant trees was associated with significantly reduced sapling stem damage, significantly increased sapling stem density, and significantly increased sapling species diversity. Considered in sum, our results suggest that elephants can affect the structure and diversity of Afrotropical forests through their foraging activities, with some variation based on location and plant size class. Developing a more complete understanding of elephants’ ecological effects will require continued research, ideally with manipulative experiments.     

High genetic diversity and contrasting fine-scale spatial genetic structure in four seasonally dry tropical forest tree species

In this study we compared population structure, genetic diversity and fine-scale spatial genetic structure (SGS) in four Bignoniaceae tree species, Handroanthus chrysotrichus, H. impetiginosus, Tabebuia roseoalba and H. serratifolius in a remnant of seasonally dry tropical forest in Central-West Brazil, based on polymorphisms at six microsatellite loci. All species, except T. roseoalba, presented the inverted ‘J’ population structure indicating recruitment of juveniles. Juveniles presented a clumped distribution suggesting limitation in dispersal or patchy distribution of suitable microhabitat for recruitment. All species showed high levels of polymorphism and genetic diversity but without a clear pattern of distribution among life stages. The SGS was significant for all species, except T. roseoalba, but the pattern and strength of the spatial genetic structure differed among species. Handroanthus serratifolius had stronger SGS with significant kinship until 77 m. For H. impetiginosus and H. chrysotrichus, kinship was significant just until 23 and 6 m, respectively. Despite the high genetic diversity, all species showed low number of adults and high fixation indices suggesting that habitat fragmentation and disturbance have been affecting these populations in Central-West Brazil.     

Mass effects,clonality, and phenology but not seed traits predict species success in colonizing restored grasslands

Preserved grasslands commonly host species‐rich plant communities but a large part of the grasslands were plowed up in the past. Their restoration often requires a long time and initial restoration measures might trigger ecosystem recovery, which is then followed by spontaneous colonization. We evaluate the establishment success of target grassland species, which were not sown but established spontaneously in the restored grasslands of Bílé Karpaty Mts., SE Czech Republic. According to their key functional traits and incidence in the landscape (mass effect; acquired from the results of a grid mapping project in the region), we examined the frequency of species and their mean cover in 82 restored grasslands. The best predictor of species frequency in the grasslands was their mass effect, followed by a high capacity for clonal growth and late phenology. Seed dispersal traits (seed mass, terminal velocity, epizoochory ranking index) and plant height had no significant effect. Specific leaf area was positively correlated with mass effect. Species having a high cover in the restored grasslands had a high capacity for clonal growth. In the preparation of seed mixtures, we should therefore consider that nonclonal species relying on regeneration from seeds will be generally less able to reproduce and should be promoted by artificial sowing. At the same time, species common in the landscape, which spread well clonally, and those with a late phenology, might be expected to colonize restored meadows on their own, so that sowing them is not necessary.     

Species traits and abundances predict metrics of plant–pollinator network structure,but not pairwise interactions

Plant–pollinator mutualistic networks represent the ecological context of foraging (for pollinators) and reproduction (for plants and some pollinators). Plant–pollinator visitation networks exhibit highly conserved structural properties across diverse habitats and species assemblages. The most successful hypotheses to explain these network properties are the neutrality and biological constraints hypotheses, which posit that species interaction frequencies can be explained by species relative abundances, and trait mismatches between potential mutualists respectively. However, previous network analyses emphasize the prediction of metrics of qualitative network structure, which may not represent stringent tests of these hypotheses. Using a newly documented temporally explicit alpine plant–pollinator visitation network, we show that metrics of both qualitative and quantitative network structure are easy to predict, even by models that predict the identity or frequency of species interactions poorly. A variety of phenological and morphological constraints as well as neutral interactions successfully predicted all network metrics tested, without accurately predicting species observed interactions. Species phenology alone was the best predictor of observed interaction frequencies. However, all models were poor predictors of species pairwise interaction frequencies, suggesting that other aspects of species biology not generally considered in network studies, such as reproduction for dipterans, play an important role in shaping plant–pollinator visitation network structure at this site. Future progress in explaining the structure and dynamics of mutualistic networks will require new approaches that emphasize accurate prediction of species pairwise interactions rather than network metrics, and better reflect the biology underlying species interactions.     

Clonal structure and genetic diversity of three desert phreatophytes

The objective of this paper was to assess clone sizes of three perennial desert plant species with AFLP markers and to relate them to clonal and genetic diversity and to hydroecology. The study was carried out at the southern rim of the Taklamakan Desert, where sexual regeneration is only possible shortly after rare flooding events, resulting in rarely established cohorts with subsequent extensive vertical growth and horizontal clonal spread. In this environment, repeated seedling establishment is excluded. We expected decreasing clonal and genetic diversity with increasing clone size and increasing distance to the groundwater table and a common response pattern among all study species. Maximum sizes of Populus euphratica and Alhagi sparsifolia clones were 121 ha and 6.1 ha, respectively, while Tamarix ramosissima clones reached a maximum size of only 38 m(2). In P. euphratica and A. sparsifolia, clonal diversity declined with increasing clone size and increasing distance to the groundwater table, while genetic diversity remained unaffected. Tamarix ramosissima differed from the other species because of a much smaller clonality. Clone size and clonal diversity were found to be good proxy variables for clone age. Despite the considerable age of the clones, genetic diversity is maintained in the populations.     

Tree species influences soil microbial community diversity but not biomass in a karst forest in southwestern China

中国西南喀斯特森林树种对土壤微生物群落多样性和生物量的影响陆地生态系统中植物种对土壤微生物群落结构的影响不一,而喀斯特生态系统中植物种对土壤微生物群落结构影响的研究尚未见报道。本研究利用磷酸脂肪酸(PLFA)法,分析了黔中高原型喀斯特常绿落叶阔叶混交林5个优势树种—窄叶石栎(Lithocarpus confinis Huang)、圆果化香(Platycarya longipes Wu)、滇鼠刺(Itea yunnanensis Franch.)、安顺润楠(Machilus cavaleriei H. Lév.)、云贵鹅耳枥(Carpinus pubescens Burkill)—与土壤理化性质对土壤微生物群落组成和生物量的影响。在测试的212个土壤样品 中共检测出132种PLFA,每个样品土壤微生物平均PLFA数量和生物量分别为65.97和11.22 µg g^–1。土壤表层(0–10 cm)的土壤微生物PLFA数量与下层(10–20 cm)土壤接近,但前者土壤微生物生物量显著高于后者(P < 0.05)。树种影响土壤微生物PLFA数量,但对土壤微生物生物量没有影响。云贵鹅耳枥附近的土壤微生物PLFA数量显著高于其他树种(P < 0.05),而其他树种土壤微生物PLFA数量接近。土壤微生物 生物量与表层土壤的理化性质无显著相关,但与下层土壤的有机碳、全氮和全磷含量呈显著正相关 (P < 0.05)。总之,黔中高原型喀斯特森林真菌/细菌生物量比率低,微生物总生物量低,但微生物群落多样性高。树种对土壤微生物群落多样性产生影响。     

Linking traits to species diversity and community structure in phytoplankton

In addition to answering Hutchinson’s question “Why are there so many species?”, we need to understand why certain species are found only under certain environmental conditions and not others. Trait-based approaches are being increasingly used in ecology to do just that: explain and predict species distributions along environmental gradients. These approaches can be successful in understanding the diversity and community structure of phytoplankton. Among major traits shaping phytoplankton distributions are resource utilization traits, morphological traits (with size being probably the most influential), grazer resistance traits, and temperature responses. We review these trait-based approaches and give examples of how trait data can explain species distributions in both freshwater and marine systems. We also outline new directions in trait-based approaches applied to phytoplankton such as looking simultaneously at trait and phylogenetic structure of phytoplankton communities and using adaptive dynamics models to predict trait evolution.     

Species distribution models predict temporal but not spatial variation in forest growth

Bioclimate envelope models have been widely used to illustrate the discrepancy between current species distributions and their potential habitat under climate change. However, the realism and correct interpretation of such projections has been the subject of considerable discussion. Here, we investigate whether climate suitability predictions correlate to tree growth, measured in permanent inventory plots and inferred from tree‐ring records. We use the ensemble classifier RandomForest and species occurrence data from ~200,000 inventory plots to build species distribution models for four important European forestry species: Norway spruce, Scots pine, European beech, and pedunculate oak. We then correlate climate‐based habitat suitability with volume measurements from ~50‐year‐old stands, available from ~11,000 inventory plots. Secondly, habitat projections based on annual historical climate are compared with ring width from ~300 tree‐ring chronologies. Our working hypothesis is that habitat suitability projections from species distribution models should to some degree be associated with temporal or spatial variation in these growth records. We find that the habitat projections are uncorrelated with spatial growth records (inventory plot data), but they do predict interannual variation in tree‐ring width, with an average correlation of .22. Correlation coefficients for individual chronologies range from values as high as .82 or as low as ?.31. We conclude that tree responses to projected climate change are highly site‐specific and that local suitability of a species for reforestation is difficult to predict. That said, projected increase or decrease in climatic suitability may be interpreted as an average expectation of increased or reduced growth over larger geographic scales.     

Leaf habit does not predict leaf functional traits in cerrado woody species

Plant species with a high leaf life span (LLS) commonly have a low specific leaf area (SLA), leaf nitrogen per unit mass (N), and phosphorous concentration (P), whereas species with low LLS have a high SLA, N and P. However, LLS tends to be longer in species growing in low-nutrient soils and, therefore, differences in LLS and other leaf traits may not be consistent with a plant classification according to leaf habit. Here we investigated whether leaf habit is consistent with leaf economic spectrum trade-offs in cerrado (a Neotropical savanna) woody species. We analyzed the SLA, N and P of 125 woody species with a distinct leaf habit (deciduous, semideciduous, brevideciduous or evergreen). We also gathered data on the LLS (33 species), maximum net photosynthesis per leaf area (A_area, 56 species) and per leaf mass (A_mass, 31 species), comprising the most extensive database analyzed so far for the cerrado. Differences among leaf habit groups were tested using generalized linear mixed models and ANOVA. We did not find differences in SLA and N among species with a distinct leaf habit, but deciduous species had a higher leaf P concentration than evergreens. Species did not differ in LLS and A_mass, but A_area varied among groups. Semideciduous species had higher A_area values than deciduous and brevideciduous species, but all other groups had similar A_area values. Because of the small difference in the LLS, SLA, leaf N, leaf P and maximum net photosynthesis, we argue that deciduous, brevideciduous, semideciduous and evergreen species may not constitute different functional groups in cerrado woody species.     

Parallel declines in species and genetic diversity in tropical forest fragments

The potential for parallel impacts of habitat change on multiple biodiversity levels has important conservation implications. We report on the first empirical test of the 'species-genetic diversity correlation' across co-distributed taxa with contrasting ecological traits in the context of habitat fragmentation. In a rainforest landscape undergoing conversion to oil palm, we show that depauperate species richness in fragments is mirrored by concomitant declines in population genetic diversity in the taxon predicted to be most susceptible to fragmentation. This association, not seen in the other species, relates to fragment area rather than isolation. While highlighting the over-simplification of extrapolating across taxa, we show that fragmentation presents a double jeopardy for some species. For these, conserving genetic diversity at levels of pristine forest could require sites 15-fold larger than those needed to safeguard species numbers. Importantly, however, each fragment contributes to regional species richness, with larger ones tending to contain more species.     

Impacts of forest gaps on butterfly diversity in a Bornean peat-swamp forest

Forest degradation is leading to widespread negative impacts on biodiversity in South-east Asia. Tropical peat-swamp forests are one South-east Asian habitat in which insect communities, and the impacts of forest degradation on them, are poorly understood. To address this information deficit, we investigated the impacts of forest gaps on fruit-feeding butterflies in the Sabangau peat-swamp forest, Central Kalimantan, Indonesia. Fruit-baited traps were used to monitor butterflies for 3 months during the 2009 dry season. A network of 34 traps (n_gap = 17, n_shade = 17) was assembled in a grid covering a 35 ha area. A total of 445 capture events were recorded, comprising 384 individuals from 8 species and 2 additional species complexes classified to genera. On an inter-site scale, canopy traps captured higher species richness than understory traps; however, understory traps captured higher diversity within each site. Species richness was positively correlated with percent canopy cover and comparisons of diversity indices support these findings. Coupled with results demonstrating morphological differences in thorax volume and forewing length between species caught in closed-canopy traps vs. those in gaps, this indicates that forest degradation has a profound effect on butterfly communities in this habitat, with more generalist species being favored in disturbed conditions. Further studies are necessary to better understand the influences of macro-habitat quality and seasonal variations on butterfly diversity and community composition in South-east Asian peat-swamp forests.