Species‐time‐area and phylogenetic‐time‐area relationships in tropical tree communities

The species‐area relationship (SAR) has proven to be one of the few strong generalities in ecology. The temporal analog of the SAR, the species‐time relationship (STR), has received considerably less attention. Recent work primarily from the temperate zone has aimed to merge the SAR and the STR into a synthetic and unified species‐time‐area relationship (STAR) as originally envisioned by Preston (1960). Here we test this framework using two tropical tree communities and extend it by deriving a phylogenetic‐time‐area relationship (PTAR). The work finds some support for Preston's prediction that diversity‐time relationships, both species and phylogenetic, are sensitive to the spatial scale of the sampling. Contrary to the Preston's predictions we find a decoupling of diversity‐area and diversity‐time relationships in both forests as the time period used to quantify the diversity‐area relationship changes. In particular, diversity‐area and diversity‐time relationships are positively correlated using the initial census to quantify the diversity‐area relationship, but weakly or even negatively correlated when using the most recent census. Thus, diversity‐area relationships could forecast the temporal accumulation of biodiversity of the forests, but they failed to “back‐cast” the temporal accumulation of biodiversity suggesting a decoupling of space and time.     

Fragmentation,grazing and the species–area relationship

Habitat loss is one of the greatest threats to species persistence. Gauging the scale of this problem requires quantitative methods that can predict the number of extinctions resulting from habitat loss. For the past three decades, the species–area relationship, an empirical relationship between the number of species present in an area and the size of that area, has been this tool. However, it fails to incorporate threats to species aside from habitat loss and the heterogeneous distribution of these threats across habitats. Recent studies have improved species–area predictions by incorporating not only direct effects of area on richness, but also indirect effects of area (through area‐mediated predator abundance), on prey species richness. We extend this work to test the hypotheses that the indirect effects of the multiple threats of grazing and trampling in addition to fragmentation will amplify the effect of area on species richness and that this effect will be greatest in zones closest to the fragment edge. Further, we test for species and population level effects of fragmentation and grazing, including the non‐random pattern of species loss and the decline in population sizes. We test our hypotheses with a field study of land snail richness in fragments with and without the additional threats of grazing and trampling. Our study supports the hypotheses that fragments with multiple threats in addition to habitat loss harbour fewer species than fragments without these threats, and that this effect is non‐uniform across fragments, populations and species.     

Habitat heterogeneity overrides the species–area relationship

Aim The most obvious, although not exclusive, explanation for the increase of species richness with increasing sample area (the species–area relationship) is that species richness is ultimately linked to area-based increases in habitat heterogeneity. The aim of this paper is to examine the relative importance of area and habitat heterogeneity in determining species richness in nature reserves. Specifically, the work tests the hypothesis that species–area relationships are not positive if habitat heterogeneity does not increase with area. Location Sixteen nature reserves (area range 89–11,030 ha) in central Hungary. Methods Four-year faunistic inventories were conducted in the reserves involving c. 70 fieldworkers and 65 taxonomists. CORINE 50,000 land-cover maps were used for calculating the heterogeneity of the reserve landscape (number of habitat types, number of habitat patches and total length of edges). Results Large reserves were less heterogeneous than small reserves, probably because large reserves were established in large blocks of unproductive land whereas small reserves tended to be in more fertile land. In total, 3975 arthropod species were included in the analysis. The slope of the species–area relationship was positive only for Neuroptera and Trichoptera. There was no significant relationship in the other nine taxa examined (Collembola, Acari, Orthoptera, Thysanoptera, Coleoptera, Araneae, Diplopoda, Chilopoda, Diptera). The density (number of species ha⁻¹) of all species, however, showed a positive correlation with heterogeneity. Main conclusions The general lack of fit of species–area relationships in this study is inconsistent with most previous published studies. Importantly, and unlike many other studies, habitat heterogeneity was not correlated with reserve area in the studied system. In the absence of this source of covariation, stronger relationships were identified that suggested a fundamental link between species richness and habitat heterogeneity. The results indicate that habitat heterogeneity rather than area per se is the most important predictor of species richness in the studied system.     

Evolutionary species–area curves as revealed by single-island endemics: insights for the inter-provincial species–area relationship

The species–area relationship (SAR) between different biological provinces is one of the most interesting, but least explored aspects of the well-known species–area pattern. Following the usage that a biological province is a region whose species have for the most part evolved within it, rather than immigrating from somewhere else, we propose that islands can be considered equivalent to biological provinces for single island endemic species (SIEs). Hence, analyses of the relationships between numbers of SIEs and island area can be used as model systems to explore the form of inter-provincial SARs. We analyzed 13 different data sets derived from 11 sources, using the power (log–log) model. Eleven of the SIE–area relationships were statistically significant, explaining high proportions of the variance in SIE numbers (R² 0.57–0.95). The z-values (slopes) of the statistically significant SIE–area relationships ranged from 0.47 to 1.13, with a mean value of 0.80 (SD±0.24).
All the island systems in which SIE represent >50% of species exhibited z-values for the SARs of native species higher than those deemed typical of archipelagic SARs. The SIE–area slopes are quite similar to those previously calculated for inter-provincial SARs, indicating that, when speciation becomes the dominant process adding to the species richness of assemblages, high z-values should be anticipated, regardless of the biogeographical scale of the study system.     

Negative density–area relationships: the importance of the zeros

Aim Estimates of abundances and densities of birds and mammals have often been shown to be scale dependent, in that population sizes over large areas are overestimated if extrapolated from surveys of small plots. Previous tests of the mechanisms suggested to cause this decelerating scaling pattern found evidence of a biased choice of small plots in patches of homogeneous habitat. Here we show that negative density–area relationships can also arise as result of not considering plots where individuals of the species or assemblage of interest are absent in surveys of differing spatial resolution. Location We took a complete census of violets (Viola spp.) in 800 m² of chalk grassland in Wye, Kent, UK, and used human population censuses for Finnish, Swiss and Italian municipalities, English districts, states of the USA and European countries. Methods We used mixed models of logarithmically transformed number of individuals or densities as a function of area. Results The census of violets shows that by increasing the survey resolution and by not considering plots without individuals, the effectively occupied area diminishes and a negative density–area relationship arises. The finding that negative density–area relationships are also common for people is evidence that the non‐random choice of plots in population surveys of varying areas can be responsible for many observed negative density–area relationships. The shallower slope of the people–administrative area relationship for Switzerland and Finland compared with Italy, as well as for England and the USA compared with Europe, confirms that less than proportionate individuals–area relationships can be the consequence of larger plot areas containing a higher proportion of areas without individuals. Main conclusions Densities should be reported together with the effective areas for which they were estimated. It should be clearly conveyed whether or not plots where the surveyed species was absent were included in the density estimation.     

Abscesses of the breast; recurring lesions in the areolar area

Subareolar abscesses beginning either in infected skin glands or in breast ducts have an extraordinary tendency to recur and to be resistant to treatment. About three-fourths of 64 patients observed had from one to many recurrences of abscess after either spontaneous or surgical drainage, and many even after wide excision of scar in an interval of quiescence.The most successful of a number of methods of treatment used was wide removal of scar and underlying chronic abscess cavity combined with removal of the ampulla and mouth of a connecting duct. In a substantial number, after either drainage or unsuccessful excision, the process gradually subsided over a period of months or years. Cancer has not been observed in any of the 64 patients.     

Species–area and species–sampling effort relationships: disentangling the effects

Species numbers tend to increase with both the area surveyed (species–area relationship, SAR) and the number of samples taken (species–sampling effort relationship, SSER). These two relationships differ in their nature and underlying mechanisms but are not clearly distinguished in field studies. To discriminate the effects of area (spatial extent) and sampling effort (SE) on species richness, several models explicitly involving both variables were proposed and tested against 13 datasets from marine micro‐, meio‐ and macrobenthos. A combination of power SSER and piecewise power SAR terms was found to have the best fit. The effects of area and SE were both significant, but the former one was noticeably weaker. The SSERs were roughly linear in log‐log space, whereas the SARs demonstrated scale‐dependent behavior with a noticeable threshold (slope breakpoint). Species richness was almost area‐independent below this threshold (the “small area effect”, SAE) but followed typical power‐law SAR beyond the threshold. This effect was similar to the “small island effect” but occurred for arbitrarily delineated areas within continuous habitats. Parameters of the SAR curves depended on organism size. The upper limit of the SAE increased from microorganisms to meiofauna to macrofauna. Also, SAR curves for unicellular groups had significantly lower slopes. SAE is supposed to indicate a spatial range of statistical homogeneity in species composition. Its upper limit corresponds to the characteristic size of a local community (a single habitat occupied by a common species pool). Interpretations of SAR and SSER parameters in terms of α‐ and β‐diversity are proposed. Both SAR and SSER slopes obtained from univariate regressions are overestimated. This upward bias depends on sampling design, decreasing for SAR but increasing for SSER with more unequally spaced samples. Both spatial extent and sampling effort should be taken into account to disentangle properly their effects on diversity.     

A test of the metapopulation model of the species–area relationship

Abstract Aim The species–area relationship is a ubiquitous pattern. Previous methods describing the relationship have done little to elucidate mechanisms producing the pattern. Hanski & Gyllenberg (Science, 1997, 275 , 397) have shown that a model of metapopulation dynamics yields predictable species–area relationships. We elaborate on the biological interpretation of this mechanistic model and test the prediction that communities of species with a higher risk of extinction caused by environmental stochasticity should have lower species–area slopes than communities experiencing less impact of environmental stochasticity. Methods We develop the mainland–island version of the metapopulation model and show that the slope of the species–area relationship resulting from this model is related to the ratio of population growth rate to variability in population growth of individual species. We fit the metapopulation model to five data sets, and compared the fit with the power function model and Williams's (Ecology, 1995, 76 , 2607) extreme value function model. To test that communities consisting of species with a high risk of extinction should have lower slopes, we used the observation that small‐bodied species of vertebrates are more susceptible to environmental stochasticity than large‐bodied species. The data sets were divided into small and large bodied species and the model fit to both. Results and main conclusions The metapopulation model showed a good fit for all five data sets, and was comparable with the fits of the extreme value function and power function models. The slope of the metapopulation model of the species–area relationship was greater for larger than for smaller‐bodied species for each of five data sets. The slope of the metapopulation model of the species–area relationship has a clear biological interpretation, and allows for interpretation that is rooted in ecology, rather than ad hoc explanation.     

Biodiversity on urban roundabouts—Hemiptera, management and the species–area relationship

The biodiversity of insects within urban areas has been relatively little studied. Given the large and ever increasing extent of urban areas, and that the insect species richness there can be high, it is important to know the factors determining that aspect of biodiversity. In this study two of these factors, namely habitat management and area, were considered. Arboreal and grassland Hemiptera, and grassland plants, were sampled on 18 roundabouts and other road enclosed sites in the town of Bracknell. Hemiptera were sampled using suction sampling and tree beating. A significant species–area relationship was found for arboreal Hemiptera, which was strongly related to habitat diversity. For both grassland plants and Hemiptera, grassland management, by mowing, had a significant effect on species richness. Despite the management grassland plants showed a significant species–area relationship. However the effect of management on Hemiptera was great enough to outweigh any area effect. As the size of open spaces is often constrained in urban areas, altering habitat management has a greater potential for enhancing biodiversity. For arboreal Hemiptera choice of trees for planting is of particular importance, while for grassland Hemiptera diversity would be increased with a reduction in the intensity of management, such a reduction in the frequency of mowing.

Zusammenfassung

Die Biodiversität der Insekten auf urbanen Flächen ist relativ wenig untersucht. Angesichts der großen und der immer größer werdenden Ausdehnung urbaner Gebiete und angesichts dessen, dass der Artenreichtum der Insekten dort groß sein kann, ist es wichtig die Faktoren zu kennen, die diesen Aspekt der Biodiversität bestimmen. In dieser Untersuchung wurden zwei dieser Faktoren, nämlich Habitatmanagement und Fläche, betrachtet. Baum- und wiesenbewohnende Hemiptera sowie Wiesenpflanzen wurden in 18 Kreisverkehren und anderen straßenumschlossenen Orten innerhalb der Stadt Bracknell gesammelt. Die Hemiptera wurden mit Saugproben und Klopfproben an den Bäumen gesammelt. Für die baumbewohnenden Hemiptera wurde eine signifikante Art-Areal-Beziehung gefunden, die in enger Beziehung zur Habitatdiversität stand. Sowohl für die Wiesenpflanzen als auch für die Hemiptera hatte das Wiesenmanagement in Form von Mahd einen signifikanten Einfluss auf den Artenreichtum. Trotz des Managements zeigten die Wiesenpflanzen eine signifikante Art-Areal-Beziehung. Die Auswirkungen des Managements auf die Hemiptera waren jedoch groß genug, um den Arealeffekt zu überwiegen. Da die Größe offener Flächen in städtischen Gebieten oft beschränkt ist, hat die Änderung des Habitatmanagements ein größeres Potenzial die Biodiversität zu erhöhen. Für baumbewohnende Hemiptera ist die Auswahl der Bäume für die Bepflanzung von besonderer Wichtigkeit, während für die wiesenbewohnenden Hemiptera die Diversität durch eine Verringerung der Managementintensität erhöht würde, wie z. B. durch die Verringerung der Mahdfrequenz.     

The congener; A neglected area in the study of behaviour

This paper seeks a deeper understanding of the congener as a factor in animal and human behaviour. It does so, not by concentrating on analyses of stimulus exchanges - largely specific to the species - by which a congener is recognized, but on the more general questions of why a notion of congener exists at all and why it plays such an extraordinary important role in animal and human behaviour.Three separate approaches, by way of anthropomorphic psychology, a paraphysical energy model and the physical theory of the implicate order, lead to the recognition of a certain inseparability of self and congener; and to an interpretation of the content of the notion of congener and of the behaviour in relation to it, in terms of the fundamental concept of energy and the even more fundamental one of order.Dedicated to Professor Gerard P. Baerends, one of my valued guides in ethology.     

Insect–fern interactions: macrolepidopteran utilization and species–area association

Abstract.

• 1 The generalization that ferns are under-utilized by phytophagous insects in comparison to angiosperms may be invalid because of biases involving plant growth form, plant range, and unequal sampling efforts.
• 2 Comparison of nineteen fern species with 652 herb species, the angjosperm growth form most similar to the ferns, indicates no significant difference in the mean number of supported macrolepidopteran species. When the herbs are subdivided into annuals, biennials and perennials, only the annual herbs are significantly different than the ferns.
• 3 Comparisons of the occurrence distributions for ferns and the herb categories also demonstrate that only the annual herbs support more macrolepidopteran species than the ferns. The same results are obtained when random assemblages of herbs are created that are the same size as the fern assemblage.
• 4 Both the occurrence distributions and the species–area relationship for the ferns indicate that host records for insects feeding on ferns may be grossly incomplete.
• 5 The similarity of exploitation of ferns and perennial herbs by the Macro-lepidoptera suggests that other foliage feeding insects may also use ferns at levels equivalent to angiosperms.