Determinants of diversity in afrotropical herbivorous insects (Lepidoptera: Geometridae): plant diversity,vegetation structure or abiotic factors?

Aim This study was conducted to investigate the potential of predicting alpha diversity and turnover rates of a highly diverse herbivorous insect family (Geometridae) based on vascular plant species richness and vegetation structure. Location The study was carried out on the south‐western slopes of Mount Kilimanjaro within a wide range of habitats between 1200 and 3150 m elevation. Methods The floristic and structural composition of the vegetation was recorded at 48 plots of 400 m². Geometrid moths were sampled manually at light sources located at the plot centres. Principal components analysis, redundancy analysis and multiple linear regression were used to explore how alpha diversity and species turnover of geometrid moths are related to vegetation structure and plant species richness. Results Alpha diversity of geometrid moths was significantly correlated with species diversity patterns in the most common vascular plant families (R² = 0.49) and with plant structural parameters (R² = 0.22), but not with overall floristic diversity. Species turnover of geometrid moths was strongly linked to diversity changes in a range of plant families (40% explained variance), less strongly to changes in vegetation physiognomy (25%), and only weakly to overall floristic diversity (5%). Changes in elevation were a better predictor of both alpha diversity and species turnover of geometrid moths than any principal component extracted from the vegetation data. Main conclusions Vegetation composition, diversity and structure all showed significant correlations with the diversity and species composition of geometrid moth assemblages. Nevertheless, in most cases relationships were indirect, via environmental parameters such as temperature and humidity, which influenced both vegetation and moth fauna. Possible direct links between geometrid diversity and potential food plants were much weaker. The lack of a significant correlation between overall plant species richness and geometrid diversity indicates that tropical geometrid moths may not be very selective in their food plant choice. Accordingly, a clear correlation between floral diversity and herbivore species richness must be regarded as overly simplistic, and the diversity of vascular plants cannot universally be used as a suitable biodiversity indicator for diverse insect taxa at higher trophic levels.     

Ecology of the Pteridophytes on the Southern Slopes of Mt. Kilimanjaro. Part II: Habitat Selection

Abstract: Based on the evalutation of 957 vegetation plots on the southern slope of Mt. Kilimanjaro, habitat preferences for 140 species of pteridophytes were evaluated. Using the average percentage cover value, and taking into account the pteridophyte flora's composition, life form spectra and its spectra of seasonal growth pattern, eight vegetation formations were recognized. Ferns contributed less than 1 % of the vegetation cover of salt marshes, ruderal vegetation, grasslands and (sub-)alpine heathlands. In contrast, pteridophytes constituted the most important vascular plant group on rocks, where 64 species were found, forming about two-thirds of the vegetation cover. With respect to alpha and beta diversity and fern biomass, luxuriant montane forest was the main habitat for pteridophytes on Mt. Kilimanjaro. Here 130 pteridophyte species (93 % of the whole pteridophyte flora of the study area), on average, contributed 16 % of the total vegetation cover. Epiphytic ferns, tree ferns and filmy ferns had their main distribution between 1900 and 2400 m, in a zone coinciding with the maximum rainfall on Mt. Kilimanjaro's southern slope.
Poikilohydrous species were typical of dry habitats, such as on rocks, in meadows or along roadsides, but they also occurred in the often sun-exposed epiphyte layer in moist montane forests. Deciduous species, which were in many cases fire resistant, had a similar distribution; however, inside the forest belt they were restricted to the lower and upper parts, where fires are a common phenomenon. Evergreen species were the dominant group in swamps, forests and forest clearings.
Compared to other volcanoes in East Africa, Mt. Kilimanjaro is distinctly richer in fern species in general and in filmy ferns, tree ferns and epiphytic ferns in particular, suggesting that the forest belt of the southern slope of Mt. Kilimanjaro is wetter than those of other high mountains in East Africa.     

Disturbance effects on diversity of epiphytes and moths in a montane forest in Ecuador

We sampled the diversity of epiphytes (lichens, bryophytes, vascular plants) and moths (Geometridae, Arctiidae) in mature and recovering forest and in open vegetation in the montane belt in Ecuador. No uniform pattern of change in species richness was detected among the different taxonomic groups with increasing disturbance. Species richness of epiphytic bryophytes and vascular plants declined significantly from mature forest towards open vegetation. In contrast, species richness of epiphytic lichens did not change with increasing forest alteration, while that of geometrid moths was significantly higher in recovering forest compared with mature forest and open habitats. Arctiidae were significantly more species-rich in recovering forest and open vegetation than mature forest. Hence, for some organisms, modified habitats may play an important role for biodiversity conservation in the Andes, whereas others suffer from habitat disturbance. However, trends of changes in species composition following deforestation were surprisingly concordant across most studied epiphyte and moth taxa.     

Diversity of geometrid moths (Lepidoptera: Geometridae) along an Afrotropical elevational rainforest transect

Geometrid moths were investigated at 26 sites on 9 elevational levels along an elevational transect at Mt. Kilimanjaro (Tanzania), stretching from the fine‐grained mosaic of small agroforest plots with combined cultivation of trees, shrubs and crops at 1650 m through mountain rainforest to heathland at 3300 m. We sampled moths manually at light between 19 : 00 and 22 : 00 in the rainy seasons of March to May and October to January in the years 2000, 2001 and 2002. Along the transect, the composition of moth communities changed from a domination by Sterrhinae and Ennominae to a dominance of Larentiinae with increasing elevation. Overall, alpha diversity was very low compared to other tropical mountain regions. Fisher's alpha showed a maximum of 30 in the agroforest mosaic at 1650 m and decreased to values around 12 in the mountain rainforest. Communities of geometrid moths within the forest belt were significantly dissimilar from communities outside the forest. The diversity patterns on Mt. Kilimanjaro can be related to the young age, island‐like position and history of the mountain. These factors have led to the formation of a homogeneous upper mountain rainforest habitat which in turn houses homogeneous moth communities with a low diversity compared to habitats at lower elevations. Here, a heterogeneous habitat mosaic allowing the intrusion of savannah species into this former forest habitat may account for an increased diversity. In the heath zone above the forest, climatic conditions are very harsh, permitting only few specialists to thrive in this ericaceous woodland. Edge effects were discernible at the forest–heathland boundary where some moth species from heathland invaded the closed forest. At the boundary between agroforest and a forest mosaic of exotic Acacia and Eucalyptus forest plantations and natural mountain forest, diversity values remained low as the dominant species Chiasmia fuscataria accounted for far higher proportions than other dominant species in any of the other habitats.     

TOC, TON, TOS and TOP in Rainfall, Throughfall, Litter Percolate and Soil Solution of a Montane Rainforest Succession at Mt. Kilimanjaro, Tanzania

Organic nutrients have proven to contribute significantly to nutrient cycling in temperate forest ecosystems. Still, little is known about their relevance in the tropics. In the present study, organic C, N, S and P were analysed in rainfall, throughfall, litter percolate and soil solution of a montane rainforest at Mt. Kilimanjaro, Tanzania. The aim was to determine the amounts of organic nutrients in different water pathways and to assess the influence of forest disturbance on organic nutrients by comparing mature forests, secondary forests and shrub vegetation in clearings. Concentrations of all studied elements increased from rainfall to throughfall and litter percolate and then exhibited a rapid decrease in the mineral soil. Concentrations of organic P were above the detection limit only in the litter percolate. Organic N (ON) as a fraction of total N increased from 50% in rainfall (0.19 mg l⁻¹) to 66% (0.45 mg l⁻¹) in throughfall followed by a decline to 39% in the litter percolate (0.77 mg l⁻¹) of the mature forest. Similarly, proportions of organic S and P amounted to 43 and 34%, respectively, in the litter percolate in mature forest. For ON, this proportion further decreased to less than 10% in the soil solution. The latter was probably attributable to a high sorption capacity of the studied Andisols, which led to overall low organic element concentrations in the soil solution (OC: 1.2 mg l⁻¹, ON: 0.05 mg l⁻¹ at 1 m soil depth) as compared to other temperate and tropical forest ecosystems. Organic element concentrations were higher in litter percolate and soil solution under the clearings, but there were no differences in the relative contribution of these elements. Organic nutrient forms at Mt. Kilimanjaro appeared to be much less susceptible to leaching than their inorganic forms.     

Montane Andean rain forests are a global diversity hotspot of geometrid moths

Aim Andean forests are known to be a major diversity hotspot for vascular plants and vertebrates, but virtually nothing is known about the diversity of arthropods. We examined whether montane rain forests in southern Ecuador are also a diversity hotspot for arthropods, and chose geometrid moths as a model group. Location The study area in southern Ecuador (Province Zamora‐Chinchipe, 79° W, 04° S) covers c. 40 km², with 39 collecting sites within an elevational range of 1040–2677 m a.s.l. Thirty‐five of the sites were situated in an area c. 2.5 km². Additional qualitative sampling was carried out in the same area and up to an elevation of 3100 m. Methods Nocturnal moths were collected quantitatively and qualitatively using portable light towers consisting of two 15 W fluorescent tubes, and diurnal moths were collected qualitatively using an insect net. Insects were sampled in six fieldwork periods in the years 1999–2003. As diversity measures, Fisher's alpha of the log‐series distribution as well as eight estimators of total species richness were applied. Results A total of 1266 species were recorded, 63% of which were identified to named species, whereas the remainder are likely to include many undescribed species. Quantitative samples at light towers collected 35,238 specimens representing 1223 species. The extrapolated species number for these data is 1420 (incidence coverage estimator). Twenty‐one additional nocturnal species and 22 exclusively diurnal species were sampled qualitatively at elevations between 1040 and 3100 m. The pooled value of Fisher's alpha for all quantitative samples is 246 ± 3. Main conclusions The diversity of Geometridae documented here is much higher than anywhere else in the world, even without the inclusion of additional species from adjacent lowland rain forests. The number of recorded species in this small area corresponds to more than 6% of the known world fauna of geometrid moths. Our study emphasizes the importance of protecting the remaining montane Andean rain forests. For setting priorities in conservation, more studies on insect diversity are urgently required in other regions of the Andes, since montane forests are being destroyed at an alarming rate.     

Contrasting patterns of vertical stratification in two moth families in a Costa Rican lowland rain forest

The vertical stratification of two species-rich moth families (Lepidoptera: Arctiidae, Geometridae) was analysed in a lowland rain forest in Costa Rica. Moths were trapped with automatic 8 W ultraviolet light traps at three sites in the understorey and canopy of a primary forest at the La Selva Biological Station (10.4° N, 84.0° W). A total of 846 arctiid moths (148 species) and 946 geometrid moths (140 species) were analysed. Species richness and diversity of arctiid moths was significantly lower in the understorey than in the canopy (Fisher's alpha: 24 vs. 49) whereas geometrid moths showed an inverse pattern (Fisher's alpha: 44 vs. 27). This resulted in an overall increase in the proportion of conspicuously coloured species towards the canopy. Moth ensembles were clearly separated in multidimensional scaling ordinations, and differed significantly in their faunal composition and dominance between the strata. The available host plant data suggest that the flight height of moths was determined by larval resource availability. Examples include understorey flyers such as the geometrid genus Eois feeding on Piper, and canopy flyers such as the arctiid moth genera Aclytia, Macrocneme and Poliopastea which feed on lianas.     

Woody plant communities of isolated Afromontane cloud forests in Taita Hills,Kenya

In the Taita Hills in southern Kenya, remnants of the original Afromontane forest vegetation are restricted to isolated mountain peaks. To assess the level of degradation and the need for forest restoration, we examined how forest plant communities and their indicator species vary between and within remnant patches of cloud forest. We used ordinal abundance data to compare plant communities in eight forest fragments. We also analyzed data on the diversity and abundance of trees in 57 0.1 ha plots to compare tree communities within and between the largest two of these fragments, Ngangao (120 ha) and Mbololo (220 ha). The extant vegetation of the Taita Hills at landscape scale consists of secondary moist montane to intermediate montane forest. There was a high species dissimilarity between fragments (69%). Variation in species composition coincided with an abiotic gradient related to elevation. At plot level, secondary successional species and species of forest edges were most abundant and most frequent. Inferred clusters of plots almost entirely coincided with the two forest fragments. Indicator species associated with forest margins and gaps were more frequent in the smaller of the two forest fragments, while indicators for the larger fragment were more typical for less disturbed moist forest. Abiotic site variability but also different levels of disturbance determine site-specific variants of the montane forest. Conservation efforts should not only focus on maintaining forest quantity (size), but also on forest quality (species composition). Late-successional rainforest species are underrepresented in the woody plant communities of the Taita Hills and assisting restoration of viable populations of cloud forest climax tree species is urgently needed.     

Broken bridges: The isolation of Kilimanjaro's ecosystem

Biodiversity studies of global change mainly focus on direct impacts such as losses in species numbers or ecosystem functions. In this study, we focus on the long‐term effects of recent land‐cover conversion and subsequent ecological isolation of Kilimanjaro on biodiversity in a paleobiogeographical context, linking our findings with the long‐standing question whether colonization of African mountains mainly depended on long‐distance dispersal, or whether gradual migration has been possible through habitat bridges under colder climates. For this, we used Orthoptera as bioindicators, whose patterns of endemism and habitat demands we studied on about 500 vegetation plots on Kilimanjaro and Mt. Meru (Tanzania) since 1996. Land‐cover changes in the same area were revealed using a supervised classification of Landsat images from 1976 to 2000. In 1976, there was a corridor of submontane forest vegetation linking Kilimanjaro with Mt. Meru, replaced by human settlements and agriculture after 2000. Until recently, this submontane forest bridge facilitated the dispersal of forest animals, illustrated by the large number of endemic submontane forest Orthoptera shared by both mountains. Furthermore, the occurrence of common montane endemics suggests the existence of a former forest corridor with montane vegetation during much earlier times under climatic conditions 2–7°C cooler and 400–1,700 mm wetter than today. Based on the endemicity patterns of forest Orthoptera, negative consequences are predicted due to the effects of isolation, in particular for larger forest animals. Kilimanjaro is becoming an increasingly isolated ecosystem with far reaching consequences for diversity and endemism. Forest bridges between East African mountains acted as important migratory corridors and are not only a prehistoric phenomenon during periods with other climatic conditions but also disappeared in some places recently due to increasing and direct anthropogenic impact.