Subsampling Herbarium Collections to Assess Geographic Diversity Gradients: A Case Study with Endemic Orchidaceae and Rubiaceae in Cameroon

We compiled herbarium specimen data to provide an improved characterization of geographic patterns of diversity using indices of species diversity and floristic similarity based on rarefaction principles. A dataset of 3650 georeferenced plant specimens belonging to Orchidaceae and Rubiaceae endemic to Atlantic Central Africa was assembled to assess species composition per half‐degree or one‐degree grid cells. Local diversity was measured by the expected number of species (S_k) per grid cell found in subsamples of increasing size and compared with raw species richness (S_R). A nearly unbiased estimator of the effective number of species per grid cell was also used, allowing quantification of ratios of ‘true diversity’ between grid cells. Species turnover was measured using a presence/absence‐based similarity index (Sørensen) and an abundance‐based index that corrects for sampling bias (NNESS). Our results confirm that the coastal region of Cameroon is more diverse in endemic species than those more inland. The southern part of this coastal forest is, however, as diverse as the more intensively inventoried northern part, and should also be recognized as an important center of endemism. A strong congruence between Sørensen and NNESS similarity matrices lead to similar delimitations of floristic units. Hence, heterogeneous sampling seems to confer more bias when measuring patterns of local diversity using raw species richness than species turnover using Sørensen index. Overall, we argue that subsampling methods represent a useful way to assess diversity gradients using herbarium specimens while correcting for heterogeneous sampling effort. Abstract in French is available in the online version of this article.     

Thematic and Spatial Resolutions Affect Model-Based Predictions of Tree Species Distribution

Subjective decisions of thematic and spatial resolutions in characterizing environmental heterogeneity may affect the characterizations of spatial pattern and the simulation of occurrence and rate of ecological processes, and in turn, model-based tree species distribution. Thus, this study quantified the importance of thematic and spatial resolutions, and their interaction in predictions of tree species distribution (quantified by species abundance). We investigated how model-predicted species abundances changed and whether tree species with different ecological traits (e.g., seed dispersal distance, competitive capacity) had different responses to varying thematic and spatial resolutions. We used the LANDIS forest landscape model to predict tree species distribution at the landscape scale and designed a series of scenarios with different thematic (different numbers of land types) and spatial resolutions combinations, and then statistically examined the differences of species abundance among these scenarios. Results showed that both thematic and spatial resolutions affected model-based predictions of species distribution, but thematic resolution had a greater effect. Species ecological traits affected the predictions. For species with moderate dispersal distance and relatively abundant seed sources, predicted abundance increased as thematic resolution increased. However, for species with long seeding distance or high shade tolerance, thematic resolution had an inverse effect on predicted abundance. When seed sources and dispersal distance were not limiting, the predicted species abundance increased with spatial resolution and vice versa. Results from this study may provide insights into the choice of thematic and spatial resolutions for model-based predictions of tree species distribution.     

Conspecific density dependence in seedlings varies with species shade tolerance in a wet tropical forest

Density-dependent seedling mortality could increase with a species relative abundance, thereby promoting species coexistence. Differences among species in light-dependent mortality also could enhance coexistence via resource partitioning. These compatible ideas rarely have been considered simultaneously. We developed models of mortality as functions of irradiance and local conspecific density (LCD) for seedlings of 53 tropical woody species. Species varied in mortality responses to these factors, but mortality consistently increased with shading and LCD. Across species, density-dependent mortality on a per-neighbour basis was inversely related to species community abundance, but higher LCD in more common species resulted in a weak relationship between species abundance and density-dependent mortality scaled to species maximum LCD. Species mortality responses to shading and maximum LCD were strongly and positively correlated. Our results suggest that species differences in density-dependent mortality are more strongly related to physiologically based life-history traits than biotic feedbacks related to community abundance.     

Species number, species abundance and body length relationships of arboreal beetles in Bornean lowland rain forest trees

ABSTRACT.

• 1 The relationships between number of species, abundance per species, and body length are examined for 859 species of beetles in samples of arthropods collected from ten Bornean lowland forest trees by insecticide fogging. Similar relationships are examined for different feeding guilds of these beetles, and for those beetles from different species of trees.
• 2 The data are used to construct four interrelated graphs, namely species: abundance, species: body length, population abundance: body length and total number of individuals: body length distributions.
• 3 In contrast to a number of previous studies, no consistent linear relationship between population density and body length was found for the Bornean beetles and it is suggested that, as in birds, the added dispersal ability of flight reduces critical population densities necessary for persistence in small species. Previous relationships between body weight and population abundance may also be artefacts of the way in which data were gathered.
• 4 Despite large samples, we failed to locate the mode in plots of the number of species in each abundance category (species: abundance distribution).
• 5 Species: body length and total number of individuals: body length plots were similar to those found in previous studies, although using data for Coleoptera alone may have produced a steeper decline in the total number of individuals as body size increases than is apparent in samples of all arthropods.
• 6 We present the first three-dimensional graph relating numbers of species, body lengths and population abundances. The surface of this three-dimensional relationship is relatively simple.

     

Sampling procedures and species estimation: testing the effectiveness of herbarium data against vegetation sampling in an oceanic island

Questions: What is the relationship between species assemblages in herbarium collections and species abundances in the field, and how trustworthy are herbarium data in vegetation science? Location: Guadalupe Island, Baja California, Mexico. Methods: We compared species‐abundance distribution and evenness in 110 vegetation plots in Guadalupe Island against data from four herbaria. We tested whether the relative frequencies derived from herbarium specimens differed significantly from species frequencies in the field. We compared the rarefaction curves for both field and herbarium data sets, and tested whether taxonomic collectors accumulated new species at a higher rate than that observed in ecological plot sampling. Results: At any given sampling effort, the total number of observed species was higher in herbarium data. The relative abundance of common species in the field was higher, and the evenness of the distribution was lower, than in herbarium data. There was no significant correlation between species abundances in the field and in the herbaria. By selectively targeting rare species, collectors accumulate previously unseen species much faster than through ecological sampling. Conclusions: Because collectors aim for the rarer species and avoid the more common ones, the relative abundance of species in herbarium collections cannot be interpreted as a predictor of their true abundance in the field. Any statistical procedure that requires the sample to be representative of the true abundance distribution is likely to show errors when applied to herbarium data. However, because collectors actively search for rare species their rate of species accumulation is higher and their floristic lists are more complete than those obtained through ecological field sampling.     

Plant biodiversity inventory and conservation of two tropical dry evergreen forests on the Coromandel coast, south India

Species diversity, population structure, abundance and dispersion patterns of all woody plants 10cm gbh were inventoried in two 1-ha plots of tropical dry evergreen (sacred grove or temple) forests at Kuzhanthaikuppam (KK) and Thirumanikkuzhi (TM) on the Coromandel coast of south India. Site KK is a stunted forest (average tree height ca 6 m) and TM a tall forest (average tree height ca 10 m). A total of 54 species (in 47 genera and 31 families) were recorded. Species richness and stand density were 42 and 38 species and 1367 and 974 individuals ha–1 respectively for the sites KK and TM. About 50% of the total species were common to both the sites. Site TM is twofold more voluminous (basal area 29.48 m2 ha–1) than KK (basal area 15.44 m2 ha–1). Nearly one third of the individuals are multi-stemmed in the low-statured site KK whereas one fourth of the tree density is multi-stemmed in TM. Species abundance pattern varied between the two sites. The abundance of three species in KK and two species in TM is pronounced. Memecylon umbellatum, the most abundant species contributing to one third of total stand density in KK, is least represented in TM. Species richness, density and diversity indices decreased with increasing girth threshold. Most species exhibited clumped dispersion of individuals both at 0.25 and 1-ha scales. Population structure for girth frequency is an expanding one for both the sites, except for basal area distribution in KK. Variations in plant diversity and abundance are related to site attributes and human impacts. In the light of habitat uniqueness, species richness and sacred grove status, the need for conservation is emphasized.     

Changes in Forest Structure and Species Composition during Secondary Forest Succession in the Bolivian Amazon1

Changes in forest structure and species diversity throughout secondary succession were studied using a chronosequence at two sites in the Bolivian Amazon. Secondary forests ranging in age from 2 to 40 years as well as mature forests were included, making a total of 14 stands. Fifty plants per forest layer (understory, subcanopy, and canopy) were sampled using the transect of variable area technique. Mean and maximum height, total stem density, basal area, and species number were calculated at the stand level. Species diversity was calculated for each stand and for each combination of forest layer and stand. A correspondence analysis was performed, and the relationship between relative abundance of the species and stand age was modeled using a set of hierarchical models. Canopy height and basal area increased with stand age, indicating that secondary forests rapidly attain a forest structure similar in many respects to mature forests. A total of 250 species were recorded of which ca 50 percent made up 87 percent of the sampled individuals. Species diversity increased with stand age and varied among the forest layers, with the lowest diversity in the canopy. The results of the correspondence analysis indicated that species composition varies with stand age, forest layer, and site. The species composition of mature forests recovered at different rates in the different forest layers, being the slowest in the canopy layer. Species showed different patterns of abundance in relation to stand age, supporting the current model of succession.     

Distribution and abundance of vines along the light gradient in a southern temperate rain forest

Question: Are vines light‐demanding species? Location: Temperate evergreen rain forest of southern Chile (40°39′S, 72°11′W). Methods: In 45 plots of 25 m² distributed in treefall canopy gaps, secondary forest stands and old‐growth forest (15 plots per light environment), all climbing and non‐supported vines were counted and identified to species level, and canopy openness was quantified using hemispherical photographs. Vine abundance and diversity (species richness and Simpson's index) were compared in the three light environments and similarity between vine communities was estimated using Jaccard's similarity coefficient. We also determined the relationship between light niche breadth and local dominance at the species level. Results: In total there were 2510 vine individuals of 14 species. Canopy openness was significantly different in the three light environments. Species richness, diversity, community composition and density of vines were similar in treefall gaps, secondary and old‐growth forest. Of the seven more common vine species, which accounted for 91% of all vines, three had even distribution, two were more abundant in the shaded understorey, and two had higher density in well‐lit sites. Local dominance of vine species and niche breadth were not significantly associated. Conclusions: Our study in a temperate rain forest questions the widespread notion of vines as pioneer‐like species, which may be a consequence of the abundance of some lianas in disturbed sites of tropical forests. Functional arguments are needed to justify a general hypothesis on light requirements of vines, which constitute a vast group of species.     

Non-Indigenous Woody Invasive Plants in a Rural New England Town

We investigated the abundance of non-indigenous woody invasive plants in Farmington, Maine, a rural New England town in a forested landscape. We found 12 invasive species and more than 7 patches per km from surveys on 33 transects (54.3 km) along field edges, abandoned railroad right-of-ways, roadsides, and riparian zones. Invasive abundance was apparently lower than for more developed areas of the northeastern US, where, in contrast to western Maine, invasives have extensively penetrated forest interiors. Invasive abundance increased with the amount of landscaping and proximity to town, suggesting a close association between local horticulture and the spread of woody invasives. Invasive abundance and diversity were highest in riparian areas, probably due to relatively high levels of propagule pressure. Species differed in the extent of invasiveness, ranging from those still dependent on planted parent trees to fully invasive populations. The invasive species recorded in this study have caused environmental and economic damage elsewhere. The lower levels of invasiveness in Farmington are likely a result of the isolation, small human population, and forested landscape rather than low levels of invasibility. This suggests the potential for future risks, and the importance of intervention while populations can still be eradicated or controlled.     

Structure of the species--energy relationship

The relationship between energy availability and species richness (the species-energy relationship) is one of the best documented macroecological phenomena. However, the structure of species distribution along the gradient, the proximate driver of the relationship, is poorly known. Here, using data on the distribution of birds in southern Africa, for which species richness increases linearly with energy availability, we provide an explicit determination of this structure. We show that most species exhibit increasing occupancy towards more productive regions (occurring in more grid cells within a productivity class). However, average reporting rates per species within occupied grid cells, a correlate of local density, do not show a similar increase. The mean range of used energy levels and the mean geographical range size of species in southern Africa decreases along the energy gradient, as most species are present at high productivity levels but only some can extend their ranges towards lower levels. Species turnover among grid cells consequently decreases towards high energy levels. In summary, these patterns support the hypothesis that higher productivity leads to more species by increasing the probability of occurrence of resources that enable the persistence of viable populations, without necessarily affecting local population densities.     

Thornbush invasion in a savanna ecosystem in eastern Botswana

The paper describes the thornbush invasion (bush encroachment) found in the savanna at Olifants Drift in eastern Botswana. Overgrazing by cattle is responsible for the opening up of the grass sward and has enabled woody species to establish. A quantification of this process between 1950 and 1975 in an area of ca. 100 km² is given in terms of plant density as well as aboveground biomass. The loss of herbaceous biomass which generally accompanies bush encroachment and the possibilities for pasture restoration are mentioned.Species nomenclature follows the Flora of Southern Africa and the system in use at the herbarium of the Botanical Research Institute at Pretoria, South Africa.     

The influence of increasing population size and vegetation productivity on elephant distribution in the Kruger National Park

Decisions to reduce the impacts of large herbivores on biodiversity in protected areas are often based on controlling their numbers. However, numbers per se may not be the foremost consideration when managing impacts. This is because density‐related changes in distribution can also affect habitat utilization and hence, impact. In this study we tested whether changes in the distribution of African elephants are associated with increasing population size. We used spatially explicit count data collected during the dry seasons from 1998 to 2004 in South Africa's Kruger National Park. We did this at five spatial scales and in landscapes defined by vegetation, geology, climate and soils. We then investigated whether observed distributions and grid‐cell‐specific densities were associated with the remotely sensed Normalized Difference Vegetation Index (NDVI) as a measure of productivity and therefore food resource availability at the landscape scale. Consistent with density mediated changes, we found that elephant grid‐cell occupancy increased with population size, while grid‐cell‐specific density became less variable. In addition, the combined distribution of bull groups and breeding herds became less clumped with increasing population size. We further found that within landscapes elephants were present on grid‐cells with higher NDVI values, but that the influence of NDVI during the dry season on densities among landscapes may be weak. These results suggest that NDVI was more indicative of structural habitat choices such as woody vegetation than food availability per se. Our study highlights the need to consider factors other than population size alone when formulating management decisions to reduce large herbivore impacts on biodiversity in protected areas.     

Decline of birds in a human modified coastal dune forest landscape in South Africa

Previous studies demonstrate that old-growth forest remnants and vegetation regenerating after anthropogenic disturbance provide habitat for birds in a human modified coastal dune forest landscape in northern KwaZulu-Natal, South Africa. However, occurrence does not ensure persistence. Based on a 13-year monitoring database we calculated population trends for 37 bird species and general trends in overall bird density in different vegetation types. We evaluated species' characteristics as covariates of population trend and assessed changes in rainfall and proportional area and survey coverage per vegetation type. 76% of species assessed have declined, 57% significantly so at an average rate of 13.9% per year. Overall, bird density has fallen at 12.2% per year across old-growth forest and woody regenerating vegetation types. Changes in proportional area and coverage per vegetation type may partly explain trends for a few species but are unlikely to account for most. Below average rainfall may have contributed to bird declines. However, other possibilities warrant further investigation. Species with larger range extents tended to decline more sharply than did others, and these species may be responding to environmental changes on a broader geographical scale. Our results cast doubt on the future persistence of birds in this human modified landscape. More research is needed to elucidate the mechanisms driving population decline in the study area and to investigate whether the declines identified here are more widespread across the region and perhaps the continent.     

Variance in composition of inquiline communities in leaves of Sarracenia purpurea L. on multiple spatial scales

A survey of the abundances of species that inhabit the water-bearing leaves of the pitcher plant Sarracenia purpurea was conducted at several different spatial scales in northern Florida. Individual leaves are hosts to communities of inquiline species, including mosquitoes, midges, mites, copepods, cladocerans, and a diverse bacterial assemblage. Inquiline communities were quantified from four pitchers per plant, three plants per subpopulation, two subpopulations per population, and three populations. Species varied in abundance at different spatial scales. Variation in the abundances of mosquitoes and copepods was not significantly associated with any spatial scale. Midges varied in abundance at the level of populations; one population contained significantly more midges than the other two. Cladocerans varied at the level of the subpopulation, whereas mites varied at the level of the individual plants. Bacterial communities were described by means of Biolog plates, which quantify the types of carbon media used by the bacteria in each pitcher. Bacterial communities were found to vary significantly in composition among individual plants but not among populations or subpopulations. These results suggest that independent factors determining the abundances of individual species are important in determining community patterns in pitcher-plant inquilines.     

Lack of tree layer control on herb layer characteristics in a subtropical forest,China

Question: Knowledge of the interaction between understorey herb and overstorey tree layer diversity is mostly restricted to temperate forests. How do tree layer diversity and environmental variables affect herb layer attributes in subtropical forests and do these relationships change in the course of succession? Do abundance and diversity of woody saplings within the herb layer shift during succession? Location: Subtropical broad‐leaved forests in southeast China (29°8′18″‐29°17′29″N, 118°2′14″118°11′12″E). Methods: A full inventory of the herb layer including all plants below 1‐m height was done in 27 plots (10 × 10 m) from five successional stages (<20, <40, <60, <80 and ≥80 yr). We quantified the contribution of different life forms (herbaceous, woody and climber species) to herb layer diversity and productivity and analysed effects of environmental variables and tree layer diversity on these attributes. Results: Herb layer composition followed a successional gradient, as revealed by non‐metric multidimensional scaling (NMDS), but diversity was not correlated to the successional gradient. There was no correlation of diversity across layers. Herb layer productivity was neither affected by tree layer diversity nor by herb layer diversity. Although abundance of woody species in the herb layer decreased significantly during succession, woody species contributed extraordinarily to herb layer species diversity in all successional stages. All environmental factors considered had little impact on herb layer attributes. Conclusions: The subtropical forest investigated displays an immense richness of woody species in the herb layer while herbaceous species are less prominent. Species composition of the herb layer shows a clear successional pattern, however, the presence or absence of certain species appears to be random.     

Roles of Spatial Scale and Rarity on the Relationship between Butterfly Species Richness and Human Density in South Africa

Wildlife and humans tend to prefer the same productive environments, yet high human densities often lead to reduced biodiversity. Species richness is often positively correlated with human population density at broad scales, but this correlation could also be caused by unequal sampling effort leading to higher species tallies in areas of dense human activity. We examined the relationships between butterfly species richness and human population density at five spatial resolutions ranging from 2'' to 60'' across South Africa. We used atlas-type data and spatial interpolation techniques aimed at reducing the effect of unequal spatial sampling. Our results confirm the general positive correlation between total species richness and human population density. Contrary to our expectations, the strength of this positive correlation did not weaken at finer spatial resolutions. The patterns observed using total species richness were driven mostly by common species. The richness of threatened and restricted range species was not correlated to human population density. None of the correlations we examined were particularly strong, with much unexplained variance remaining, suggesting that the overlap between butterflies and humans is not strong compared to other factors not accounted for in our analyses. Special consideration needs to be made regarding conservation goals and variables used when investigating the overlap between species and humans for biodiversity conservation.     

The fitness of chemotrophs increases when their catabolic by‐products are consumed by other species

Chemotrophic microorganisms synthesise biomass by utilising energy obtained from a set of chemical reactions that convert resources to by‐products, forming catabolic interactions. The amount of energy obtained per catabolic reaction decreases with the abundance of the by‐product named as the ‘abundant resource premium’. Consider two species, Species 1 and 2, Species 1 obtains energy from a reaction that converts resource A to by‐product B. Species 2 then utilises B as its resource, extracting energy from a reaction that converts B to C. Thus, the presence of Species 2 reduces the abundance of B, which improves the fitness of Species 1 by increasing the energy acquisition per reaction of A to B. We discuss the population dynamic implication of this effect and its importance in expanding a realised niche, boosting material flow through the ecosystem and providing mutualistic interactions among species linked by the material flow. Introducing thermodynamics into population ecology could offer us fundamental ecological insights into understanding the ecology of chemotrophic microorganisms dominating the subsurface realm.     

Seasonal distribution of dinoflagellate resting cysts in surface sediments from Changjiang River Estuary

In order to understand the distribution of dinoflagellate cysts, surface sediments were collected from 15 stations in Changjiang River Estuary from 122°E to 123.5°E and from 29°N to 32°N in four cruises from May 2002 to February 2003. In the present study, 38 different cyst morphotypes representing 21 genera and 6 groups were identified, while 1 type was not identified into genus level. Species number and cell density of dinoflagellate cysts ranged from 10 to 25 species and from 12 to 587 per gram of dry weight, respectively. There were no obvious differences in cyst composition and density among seasons. However, the highest cyst species number and density were recorded in summer and winter, respectively. Cysts of heterotrophic dinoflagellates, which held 55.7% of the overall cyst density averagely, dominated cyst assemblages. Cyst density and species number increased from the west to the east, from the north to the south within the study area. Cysts of toxic dinoflagellates Alexandrium cat‐enella and Alexandrium tamarense complex distributed widely and were observed in almost all stations, with the maximum cell density of 81 per gram of dry weight.     

Life-history correlates of maximum population growth rates in marine fishes

Theory predicts that populations of animals with late maturity, low fecundity, large body size and low body growth rates will have low potential rates of population increase at low abundance. If this is true, then these traits may be used to predict the intrinsic rate of increase for species or populations, as well as extinction risks. We used life-history and population data for 63 stocks of commercially exploited fish species from the northeast Atlantic to test relationships between life-history parameters and the rate of population increase at low abundance. We used cross-taxonomic analyses among stocks and among species, and analyses that accounted for phylogenetic relationships. These analyses confirmed that large-bodied, slow-growing stocks and species had significantly lower rates of recruitment and adult production per spawning adult at low abundance. Furthermore, high ages at maturity were significantly correlated with low maximum recruit production. Contrary to expectation, fecundity was significantly negatively related to recruit production, due to its positive relationship with maximum body size. Our results support theoretical predictions, and suggest that a simply measured life-history parameter can provide a useful tool for predicting rates of recovery from low population abundance.