High tropical net diversification drives the New World latitudinal gradient in palm (Arecaceae) species richness

Aim Species richness exhibits striking geographical variation, but the processes that drive this variation are unresolved. We investigated the relative importance of two hypothesized evolutionary causes for the variation in palm species richness across the New World: time for diversification and evolutionary (net diversification) rate. Palms have a long history in the region, with the major clades diversifying during the Tertiary (65–2 Ma). Location Tropical and subtropical America (34° N–34° S; 33–120° W). Methods Using range maps, palm species richness was estimated in a 1° × 1° grid. Mean lineage net diversification was estimated by the mean phylogenetic root distance (MRD), the average number of nodes separating a species from the base of the palm phylogeny for the species in each grid cell. If evolutionary rate limits richness, then richness should increase with MRD. If time limits richness, then old, relict species (with low root distance) should predominantly occur in long‐inhabited and therefore species‐rich areas. Hence, richness should decrease with MRD. To determine the influence of net diversification across different time frames, MRD was computed for subtribe, genus and species levels within the phylogeny, and supplemented with the purely tip‐level measure, mean number of species per genus (MS/G). Correlations and regressions, in combination with eigenvector‐based spatial filtering, were used to assess the relationship between species richness, the net diversification measures, and potential environmental and geographical drivers. Results Species richness increased with all net diversification measures. The regression models showed that richness and the net diversification measures increased with decreasing (absolute) latitude and, less strongly, with increasing energy/temperature and water availability. These patterns therefore reflect net diversification at both deep and shallow levels in the phylogeny. Richness also increased with range in elevation, but this was only reflected in the MS/G pattern and therefore reflects recent diversification. Main conclusions The geographical patterns in palm species richness appear to be predominantly the result of elevated net diversification rates towards the equator and in warm, wet climates, sustained throughout most of the Tertiary. Late‐Tertiary orogeny has caused localized increases in net diversification rates that have also made a mark on the richness pattern.     

Latitudinal patterns of mammalian species richness in the New World: the effects of sampling method and faunal group

Abstract. Although the latitudinal gradient of species richness for mammals in North America is well documented, few investigators have quantified the relationship in South America. We examined the pattern in North and South America, at two spatial scales (2.5° and 5°) for each of two sampling methods (quadrats and latitudinal bands). A scale effect was evident for quadrats but not for bands. Significant linear relationships between species richness and latitude were found for three faunal groups: all mammals, nonvolant species, and bats. Effects of area confound the latitudinal relationship. By statistically removing such effects, we found that the latitudinal gradient is not an artifact of the species-area relationship, and that the latitudinal gradients for North and South America were statistically indistinguishable. Our data suggest that both faunal subgroups, nonvolant species and bats, contributed substantially to the overall mammalian pattern. Further, multiple regression analyses showed that only latitude is a necessary variable to explain bat richness; for nonvolant species, in addition to latitude, area and longitude may be important.     

The relationship between geographic area and the latitudinal gradient in species richness in New World birds

One hypothesis for the latitudinal gradient in species richness observed in most animal taxa is that the richness of a region is determined by its geographic area. However, the relationship between geographic area and species richness across regions is generally weak. It has been suggested that this is because species from the tropics spill out of this region of high richness, artificially inflating the richness of other regions. This generates the interesting prediction that the area and richness of extra-tropical regions should be more strongly correlated if tropical species are excluded. We test this prediction using the avifauna of the New World. We find that there is indeed a relationship between the land area and species richness of a region once tropical species are excluded. This relationship is independent of the latitude and productivity of regions. Both latitude and productivity can explain variance in richness unexplained by land area. There is no relationship between land area and species richness if tropical species are not excluded from the analysis, suggesting that tropical species do indeed mask the relationship between richness and area. We conclude that our results generally support the geographic area hypothesis, although tests of its other predictions and on other land masses are required.     

Latitudinal gradients of parasite species richness in primates

Infectious disease risk is thought to increase in the tropics, but little is known about latitudinal gradients of parasite diversity. We used a comparative data set encompassing 330 parasite species reported from 119 primate hosts to examine latitudinal gradients in the diversity of micro and macroparasites per primate host species. Analyses conducted with and without controlling for host phylogeny showed that parasite species richness increased closer to the equator for protozoan parasites, but not for viruses or helminths. Relative to other major parasite groups, protozoa reported from wild primates were transmitted disproportionately by arthropod vectors. Within the protozoa, our results revealed that vector‐borne parasites showed a highly significant latitudinal gradient in species richness. This higher diversity of vector‐borne protozoa near the tropics could be influenced by a greater abundance or diversity of biting arthropods in the tropics, or by climatic effects on vector behaviour and parasite development. Many vector‐borne diseases, such as leishmaniasis, trypanosomiasis, and malaria pose risks to both humans and wildlife, and nearly one‐third of the protozoan parasites from free‐living primates in our data set have been reported to infect humans. Because the geographical distribution and prevalence of many vector‐borne parasites are expected to increase because of global warming, these results are important for predicting future parasite‐mediated threats to biodiversity and human health.     

People, energy and avian species richness

Aim To investigate the inter‐relationships between energy availability, species richness and human population density, particularly whether human population density influences the manner in which species richness responds to energy availability. Location British 10‐km grid cells. Methods Using regressions, we investigate how human population density varies with energy availability and the nature of relationships between the numbers of species, classified by abundance and threat categories, and human population density. We then assess whether the relationships between these species richness measures and energy availability are altered when accounting for human population density. We conduct analyses using both independent error models and ones that control for spatial autocorrelation. Results Human population density was strongly and positively correlated with energy availability. Total species richness, and that of unthreatened, threatened, common and moderately common species, increases in a positive decelerating manner with human density. When human population density was taken into account, these species groups exhibited similar species–energy relationships, but the slopes of these relationships were significantly reduced in independent error models and, in the case of total richness, in spatial models. Main conclusions Positive correlations between human density and species richness probably arise as both increase with energy availability. Our data are compatible with the suggestion that high human population densities reduce the rate at which species richness increases with energy availability, but additional research is required before causality can be confirmed.     

Latitudinal effects on treehopper species richness (Hornoptera: Membracidae)

Abstract. 1. Membracid species richness declines with increasing latitude in the Western hemisphere but begins to increase again in temperate regions. In northern latitudes this transition occurs in the highlands of Mexico and is the result of the emergence of a new tribe and greater host specialization.
2. The relationship between number of species per genus and latitude is parabolic.
3. We suggest the low number of species per genus in tropical regions may be due to the lack of host specialization and reduced coordination of life history with host phenology.     

Variation in plant species richness of different life forms along a subtropical elevation gradient in the Himalayas, east Nepal

Aim To explore the variation in species richness along a subtropical elevation gradient, and evaluate how climatic variables explain the richness of the different life forms such as trees, shrubs, climbers, herbs and ferns. Location The study was made in a subtropical to warm temperate region in the south‐eastern part of Nepal, between 100 and 1500 m above sea level (a.s.l.). Methods The number of species was counted in six plots (50 × 20 m) in each of the 15 100 m elevation bands covering the main physiognomic structures along an imaginary transect. Each species recorded was assigned to a life form. Potential evapotranspiration (PET, i.e. energy), mean annual rainfall (MAR), and their ratio (MI = moisture index) were evaluated as explanatory variables by means of generalized linear models (GLM). Each variable was tested individually, and in addition MAR and PET were used to test the water‐energy dynamics model for each life form. Results The richness of herbaceous species, including herbaceous climbers, was unrelated to any of the climate variables. PET was strongly negatively correlated with elevation, and the following relationships were found between increasing PET and richness: (i) shrubs, trees and total species (sum of all life forms) showed unimodal responses (ii) ferns decreased monotonically, and (iii) woody climbers increased monotonically. Richness of all woody groups increased monotonically with MAR and MI. The water‐energy dynamics model explained 63% of the variation in shrubs, 67% for trees and 70% for woody species combined. Main conclusions For the various herbaceous life forms (forbs, grasses, and herbaceous climbers) we found no significant statistical trends, whereas for woody life forms (trees, shrubs, and woody climbers) significant relationships were found with climate. E.M. O’Brien's macro‐scale model based on water‐energy dynamics was found to explain woody species richness at a finer scale along this elevational‐climatic gradient.     

Ambient, productive and wind energy, and ocean extent predict global species richness of procellariiform seabirds

Aims Tests of the energy hypothesis for the large‐scale distribution of species richness have largely been concerned with the influence of two alternative forms of environmental energy, temperature and energy from primary productivity, both of which (at least in terrestrial systems) peak within the tropics. Taxa showing extra‐tropical diversity peaks present a potential challenge to the generality of species–energy theory. One such group are pelagic seabirds of the order Procellariiformes that show not only an extra‐tropical diversity peak but one confined to the Southern Ocean, hence a highly asymmetric one. They are distinct in being exceptionally adapted to take advantage of wind energy, which they may rely on for long‐distance ocean foraging for the patchy resources needed to meet their energetic needs. Wind represents a readily available source of kinetic energy, shows a strong latitudinal gradient, and has been largely omitted from species–energy theory. Moreover, maximal benefits of wind are likely to be afforded in areas of greatest available contiguous ocean extent. We compare the relative importance of wind speed, ocean productivity (chlorophyll concentration), air temperature and available ocean extent (distance) in explaining large‐scale global distribution of procellariiform species richness across the world's oceans. Location Global, oceanic. Methods Hierarchical partitioning, model selection, ordinary least squares (OLS) and spatial generalized least squares (GLS) regression. Results Hierarchical partitioning of non‐spatial regression models indicates that ocean distance is the most important predictor of procellariiform species richness followed by wind speed and then temperature. In contrast, that of spatial regression models indicates the roughly equal importance of ocean distance and temperature, followed by wind speed. Although contributing additional model fit, ocean productivity is consistently the weakest predictor. Best‐fit models include all four predictors and explain 67% of observed variation. The species–productivity relationship is negative overall, while the species–temperature relationship is hump‐shaped. In contrast, ocean distance and wind speed are positively associated with species richness. Conclusions Large‐scale procellariiform species richness distribution may represent a trade‐off in the use of different energy forms, being highest in Southern Ocean areas where productive energy and temperature are relatively low, but where available ocean foraging extent and wind energy required to utilize it are near‐maximal.     

Elevational gradient and human effects on butterfly species richness in the French Alps

We examined how butterfly species richness is affected by human impact and elevation, and how species ranges are distributed along the elevational gradient (200–2700 m) in the Isère Department (French Alps). A total of 35,724 butterfly observations gathered in summer (May–September) between 1995 and 2015 were analyzed. The number of estimated species per 100‐m elevational band was fitted to the elevational gradient using a generalized additive model. Estimations were also performed on a 500 m × 500 m grid at low altitude (200–500 m) to test for the human impact on species richness using generalized least squares regression models. Each species elevational range was plotted against the elevational gradient. Butterfly richness along the elevational gradient first increased (200–500 m) to reach a maximum of 150 species at 700 m and then remained nearly constant till a sharp decrease after 1900 m, suggesting that after some temperature threshold, only few specialized species can survive. At low elevation, urbanization and arable lands had a strongly negative impact on butterfly diversity, which was buffered by a positive effect of permanent crops. Butterfly diversity is exceptionally high (185 species) in this alpine department that represents less than 5% of the French territory and yet holds more than 70% of all the Rhopalocera species recorded in France. Both climate and habitat shape the distribution of species, with a negative effect of anthropization at low altitude and strong climatic constraints at high altitude.     

Triatominae species of Suriname (Heteroptera: Reduviidae) and their role as vectors of Chagas disease

Nine species of Triatominae, representing three tribes and five genera, are currently known in Suriname. An annotated list of the species based on the collections of the Bureau of Public Health (Suriname), the National Zoological Collection Suriname and the National History Museum Leiden (the Netherlands) is provided. Additionally, the results of several years of opportunistic collection in two domestic environments are presented. The most common species are Rhodnius pictipes Stål, 1972, Rhodnius robustus Larrouse, 1972 and Panstrongylus geniculatus (Latreille, 1811). The significance of the species as vectors of Chagas disease in Suriname is discussed.     

External female genitalia of six species of the genus Meccus (Hemiptera: Reduviidae: Triatominae)

Triatomine classification is based on morphological characteristics. Studies have been conducted to improve their identification by observing many characteristics. However, there are problems of differentiating among highly interrelated species and new criteria are required. The purpose of this study was to determine the morphological differences in the external female genitalia of M. pallidipennis, M. longipennis, M. picturatus, M. bassolsae, M. mazzottii, and M. phyllosomus in order to distinguish among species using scanning electron microscopy. Observations were made of the dorsal, posterior, lateral, and ventral views of the female external genitalia for each species. In the six species we studied, relevant differences were observed in the dorsal view of the X segment, as well as the IX, VIII, and VII tergites. In the posterior and lateral view, the most visible differences were registered in the gonocoxite size of the segments VIII, IX, and X. Finally, in the ventral view of the VII sternite, differences among species were observed in the size of the inflection in the top and upper corner. Our results show that it was possible to differentiate among the triatominae species for each of the four views by using a scanning electron microscope to analyze morphological characteristics of the VII, VIII, IX, and X abdomen segments.     

Energy flux, body size and density in relation to bird species richness along an elevational gradient in Taiwan

Aim To examine the species richness of breeding birds along a local elevational gradient and to test the following assumptions of the energy limitation hypothesis: (1) the energy flux through birds is positively correlated with above‐ground net primary productivity, (2) bird density is positively correlated with total energy flux, and (3) bird species richness is positively correlated with bird density. Location An elevational gradient from 1400 to 3700 m on Mt. Yushan, the highest mountain in Taiwan (23°28′30″ N, 120°54′00″ E), with a peak of 3952 m a.s.l. Methods We established 50 sampling stations along the elevational gradient. From March to July 1992, we estimated the density of each bird species using the variable circular‐plot method. Above‐ground net primary productivity was modelled using monthly averages from weather data for the years 1961–90. Results Bird species richness had a hump‐shaped relationship with elevation and with net primary productivity. Bird energy flux was positively correlated with net primary productivity and bird species richness was positively correlated with bird density. The relationship between bird density and energy flux was hump‐shaped, which does not support one assumption of the energy limitation hypothesis. Main conclusions The results supported two essential assumptions of the energy limitation hypothesis. However, when energy availability exceeded a certain level, it could decrease species richness by increasing individual energy consumption, which reduced bird density. Thus, energy availability is a primary factor influencing bird species richness at this scale, but other factors, such as body size, could also play important roles.     

Description of a new species of Nesotriatoma Usinger, 1944 from Cuba and revalidation of synonymy between Nesotriatoma bruneri (Usinger, 1944) and N. flavida (Neiva, 1911) (Hemiptera,Reduviidae, Triatominae)

Nesotriatoma confusa sp. nov. (Hemiptera, Reduviidae, Triatominae) is described based on specimens from Cuba. From one male, one female, and eleven nymphs of a then‐undescribed species of Nesotriatoma collected in Cuba, a colony was formed and its specimens were used to describe N. confusa sp. nov. Characters were observed on the head, thorax, abdomen, female external genitalia, and male genitalia with optical microscopy and scanning electron microscopy. We concluded that N. bruneri (Usinger, 1944 ) was indeed a synonym of N. flavida (Neiva, 1911 ) as previously proposed.     

The environmental basis of North American species richness patterns among Epicauta (Coleoptera: Meloidae)

Understanding regional variability in species richness is necessary for conservation efforts to succeed in the face of large-scale environmental deterioration. Several analyses of North American vertebrates have shown that climatic energy provides the best explanation of contemporary species richness patterns. The paucity of analyses of insect diversity patterns, however, remains a serious obstacle to a general hypothesis of spatial variation in diversity. We collected species distribution data on a North American beetle genus, Epicauta (Coleoptera: Meloidae) and tested several major diversity hypotheses. These beetles are generally grasshopper egg predators as larvae, and angiosperm herbivores as adults. Epicauta richness is highest in the hot, dry American southwest, and decreases north and east, consistent with the species richness-energy hypothesis. Potential evapotranspiration, which is also the best predictor of richness patterns among North American vertebrates, explains 80.2% of the variability in Epicauta species richness. Net primary productivity and variables measuring climatic heat energy only (such as PET) are not generally comparable, though they are sometimes treated as if they were equivalent. We conclude that the species richness-energy hypothesis currently provides a better overall explanation for Epicauta species richness patterns in North America than other major diversity hypotheses. The observed relationship between climatic energy and regional species richness may provide significant insight into the response of ecological communities to climate change.     

Elevational species richness patterns for vascular plants on Mount Kinabalu, Borneo

Aim  We quantify the elevational patterns of species richness for all vascular plants and some functional and taxonomic groups on a regional scale on a tropical mountain and discuss some possible causes for the observed patterns.
Location  Mount Kinabalu, Sabah, Borneo.
Methods  A data base containing elevational information on more than 28,000 specimens was analysed for vascular plant distribution, taking into account sampling effort. The total species richness pattern was estimated per 300-m elevational interval by rarefaction analyses. The same methods were also applied to quantify species richness patterns of trees, epiphytes, and ferns.
Results  Total species richness has a humped relationship with elevation, and a maximum species richness in the interval between 900 and 1200 m. For ferns and epiphytes the maximum species richness is found at slightly higher elevations, whereas tree species did not have a statistically significant peak in richness above the lowest interval analysed.
Main conclusions  For the first time a rigorous estimate of an elevational pattern in species richness of the whole vascular plant flora of a tropical mountain has been quantified. The pattern observed depends on the group studied. We discuss the differences between the groups and compare the results with previous studies of elevational patterns of species richness from other tropical areas. We also discuss the methods used to quantify the richness pattern and conclude that rarefaction gives an appropriate estimate of the species richness pattern.     

高黎贡山种子植物物种丰富度沿海拔梯度的变化

物种丰富度沿海拔梯度的分布格局成为生物多样性研究的热点。为探讨中尺度区域物种丰富度沿海拔梯度的分布,本文以高黎贡山为研究对象,利用该地区的地方植物志资料,结合通过GIS生成的区域数字高程模型(DEM)数据,分析了该区域全部种子植物和乔木、灌木、草本三种生活型种子植物物种丰富度的垂直分布格局以及物种密度沿海拔梯度的变化特征。结果表明：(1)全部种子植物和不同生活型植物物种丰富度随着海拔的升高呈现先增加后减小的趋势,最大值出现在海拔1500—2000m的范围;(2)物种密度与海拔也呈现单峰曲线关系;(3)物种丰富度和物种密度分布格局的形成主要受海拔所反映的水、热状况组合以及物种分布的边界影响。