Ants, altitude and change in the northern Cape Floristic Region

Aim Climate‐modelling exercises have demonstrated that the Cape Floristic Region is highly sensitive to climate change and will apparently lose much of its northern limits over the next few decades. Because there is little monitoring of diversity in this area, ant assemblage structure was investigated within the main vegetation types in the Greater Cederberg Biodiversity Corridor. In particular, we sought to determine how ant assemblage structure differs between the main vegetation types, how restricted ants – and in particular the major myrmecochores – are to the major vegetation types, and which environmental variables might underlie differences in the ant assemblages and in the specificity of species to particular areas. Location Northern Cape Floristic Region, Western Cape, South Africa. Methods Sampling was undertaken during October 2002 and March 2003 across an altitudinal gradient ranging from sea level (Lambert's Bay) to c. 2000 m a.s.l. (Sneeukop, Cederberg) and down again to 500 m a.s.l. (Wupperthal) in the Western Cape, South Africa. Pitfall traps were used to sample ants at 17 altitudinal bands, stretching over three vegetation types (Strandveld, Mountain Fynbos and Succulent Karoo). Biotic and abiotic environmental variables were collected at each sampling site. Generalized linear models were used to determine the relationships between species richness, density, abundance and the abundance of the major myrmecochores, and the environmental variables. Redundancy analysis was used to determine the relationship between ant assemblage structure and the environmental variables. The Indicator Value Method was used to identify characteristic ant species for each vegetation type and altitudinal site. Results Temperature explained significant proportions of the variation in species density and abundance, and, together with area and several vegetation variables, contributed significantly to the separation of the assemblages in the major vegetation types and biomes. Four major myrmecochores were identified [Anoplolepis sp. (cf. custodiens), Anoplolepis sp. (cf. steinergroeveri), Camponotus niveosetosus, Tetramorium quadrispinosum]. The abundances of the two Anoplolepis species were related to vegetation variables, while the abundance of the other two species showed opposite relationships with temperature variables. Fourteen ant species were characteristic of certain vegetation types and altitudes. Several of these species contributed to the differences between the assemblages. Main conclusions There are likely to be substantial and complex changes to ant assemblages as climates change in the northern Cape Floristic Region. Moreover, the importance of ants for ecosystem functioning suggests that these responses are not only likely to be a response solely to vegetation changes, but might also precipitate vegetation changes. The changes that are predicted to take place in the next 50 years in the Cape Floristic Region could be substantially exacerbated by such synergistic effects, which have major implications for long‐term conservation plans. Ongoing monitoring of this transect will reveal the nature and pace of the change as it unfolds.     

Consumer-specific responses to riverine subsidy pulses in a riparian arthropod assemblage

1. Aquatic resource fluxes from streams can provide significant subsidies for riparian consumers. Because aquatic resource fluxes can be highly variable in space and time, the subsidy efficiency (i.e. transfer to the recipient food web) is controlled by the short‐term aggregative response of riparian consumers. 2. Field manipulations of stream‐derived invertebrate prey subsidies were used to examine specific aggregative responses of ground‐dwelling arthropods to riverine subsidy pulses in a braided‐river (Tagliamento River, NE Italy). Subsidy manipulation comprised short‐term reductions of natural stream‐derived subsidies and increased subsidies of stream‐derived invertebrate prey during four seasons. 3. We hypothesised that specific aggregative responses of riparian arthropods depend on their specialisation on aquatic insects which was inferred from stable isotope analysis. Natural riverine subsidy sources including aquatic insect emergence and surface‐drifting organisms were quantified. 4. Arthropods responded significantly with a reduction in abundance by 51%, at reduced subsidies and an increase by 110% at increased subsidies, when averaged over all seasons. Different arthropod taxa responded differently to subsidy manipulations in relation to their specialisation on aquatic subsidies: ground beetles with a diet consisting predominantly of aquatic insects responded only to subsidy reductions, indicating that their local abundance was not limited by natural stream‐derived subsidies; lycosid spiders with a partly aquatic diet showed no significant response; and ants, although relying on a terrestrial diet, responded positively to added stream‐derived invertebrate prey, indicating that stranding of surface‐drifting terrestrial invertebrates represented an important subsidy pathway. 5. Ground beetles and lycosid spiders were seasonally separated in their use of aquatic subsidies. Results indicate that the life‐history characteristics of riparian consumers can control the subsidy efficiency for the recipient community. By the effective uptake of pulsed riverine‐derived subsidies, riparian arthropods can enhance the transfer of riverine food sources to the riparian food web.     

Developmental Plasticity in Protea as an Evolutionary Response to Environmental Clines in the Cape Floristic Region

Local adaptation along steep environmental gradients likely contributes to plant diversity in the Cape Region of South Africa, yet existing analyses of trait divergence are limited to static measurements of functional traits rather than trajectories of individual development. We explore whether five taxa of evergreen shrubs (Protea section Exsertae) differ in their developmental trajectories and capacity for plasticity using two environmentally-distinct common gardens in South Africa. We measured seedlings in the summer-dry season and winter-wet season of each of two consecutive years to characterize ontogeny and plasticity within years, as same-age leaf cohorts mature, and between years, i.e., from leaf one cohort to the next. We compared patterns of development between gardens to assess whether trait trajectories are programmed versus plastic and examined whether developmental differences covaried with characteristics of a seedling’s home environment. We detected plasticity in developmental trajectories for leaf area, stomatal size, stomatal pore index, and to a limited extent specific leaf area, but not for stomatal density. We showed that the species growing in the harshest environments exhibits both the smallest increase in leaf area between years and the least change in SLA and photosynthetic rates as leaves age within years. These results show that within this clade, species have diverged in developmental trajectories and plasticity as well as in mean trait values. Some of these differences may be associated with adaptation to cold and drought stress within an environmentally-complex region.     

Dated Plant Phylogenies Resolve Neogene Climate and Landscape Evolution in the Cape Floristic Region

In the context of molecularly-dated phylogenies, inferences informed by ancestral habitat reconstruction can yield valuable insights into the origins of biomes, palaeoenvironments and landforms. In this paper, we use dated phylogenies of 12 plant clades from the Cape Floristic Region (CFR) in southern Africa to test hypotheses of Neogene climatic and geomorphic evolution. Our combined dataset for the CFR strengthens and refines previous palaeoenvironmental reconstructions based on a sparse, mostly offshore fossil record. Our reconstructions show remarkable consistency across all 12 clades with regard to both the types of environments identified as ancestral, and the timing of shifts to alternative conditions. They reveal that Early Miocene land surfaces of the CFR were wetter than at present and were dominated by quartzitic substrata. These conditions continue to characterize the higher-elevation settings of the Cape Fold Belt, where they have fostered the persistence of ancient fynbos lineages. The Middle Miocene (13–17 Ma) saw the development of perennial to weakly-seasonal arid conditions, with the strongly seasonal rainfall regime of the west coast arising ~6.5–8 Ma. Although the Late Miocene may have seen some exposure of the underlying shale substrata, the present-day substrate diversity of the CFR lowlands was shaped by Pliocene-Pleistocene events. Particularly important was renewed erosion, following the post-African II uplift episode, and the reworking of sediments on the coastal platform as a consequence of marine transgressions and tectonic uplift. These changes facilitated adaptive radiations in some, but not all, lineages studied.     

Influence of fire on critically endangered Swartland Shale Renosterveld in the Cape Floristic Region

Questions

The degree to which renosterveld shrublands are fire‐dependent is currently unclear. To address this issue, the following questions were asked: (1) does smoke stimulate germination of soil‐stored seeds in renosterveld; (2) does recently‐burned renosterveld display changed composition and higher diversity than unburned vegetation; and (3) how do the species compositions of renosterveld soil seed banks and standing vegetation compare?

Location

Swartland, Cape Floristic Region, South Africa.

Methods

Soil seed bank samples from a north‐ and south‐facing slope were smoke‐treated and germinated to test for smoke‐stimulated germination. Burned standing vegetation was surveyed 16 months post‐fire, as was unburned vegetation on the same slopes. Seed bank species richness and density were compared between smoke‐treated and untreated samples within and between slopes. Burned and unburned standing vegetation were compared within and between slopes in terms of species richness, abundance and aerial cover. Compositional similarity of the seed banks and standing vegetation was assessed.

Results

Seed banks were dominated by annuals and graminoids. Smoke treatment had no effect, except for driving significantly higher species richness and seedling density in south‐facing slope perennial shrubs. Species richness and seedling density were significantly higher in seed banks on the south‐facing slope compared to the north‐facing slope. Burned standing vegetation exhibited significantly higher diversity than unburned vegetation. Annuals and graminoids displayed significantly higher species richness and aerial cover in burned renosterveld. The north‐facing slope contained less than half the number of species/m² compared to the south‐facing slope. The seed banks and standing vegetation showed low to intermediate similarity (Sørensen = 31%–53%), but grouped close together on an NMDS plot, suggesting intermediate similarity overall.

Conclusions

Elevated germination of perennial shrubs in smoke‐treated seed bank samples and increased diversity of post‐fire standing vegetation suggest the renosterveld in this study shows elements of a fire‐driven system. Certain species only recruited in burned sites, suggesting fire‐stimulated germination. Aspect had a major influence on plant community composition, with the mesic south‐facing slope being more diverse than the xeric north‐facing slope. The similarity between the seed banks and standing vegetation was higher than previously shown for renosterveld, and appears to be higher than for fynbos.     

Montane Refugia for Endemic and Red Listed Dragonflies in the Cape Floristic Region Biodiversity Hotspot

One of the features of many endemic organisms is that they are highly spatially restricted, and habitat specialists. The Kogelberg Biosphere Reserve (KBR) is a major centre of plant endemism within a global hotspot, the Cape Floristic Region (CFR). Dragonflies in this botanical hotspot have a range of habitat specialization from narrow-range specialists to widespread generalists, with an unusually strong bias towards the specialists. A huge 53% of dragonfly individuals and 26% of taxa recorded are national endemics, and three species are Red Listed. Thus, a group of predatory insects, which are largely not dependent on plant composition, mirrors the level of habitat specialization and restricted distributions of the plants at the spatial scale of the whole reserve. Although some studies caution the use of one taxon as a surrogate for another, the results here show that at the reserve scale in this global hotspot there can be remarkable concordance, suggesting further studies on other taxa should be carried out to determine the full extent of taxonomic concordance in this irreplaceable area.     

Fungal radiation in the Cape Floristic Region: an analysis based on Gondwanamyces and Ophiostoma

The Cape Floristic Region (CFR) displays high levels of plant diversity and endemism, and has received focused botanical systematic attention. In contrast, fungal diversity patterns and co-evolutionary processes in this region have barely been investigated. Here we reconstruct molecular phylogenies using the ITS and beta-tubulin gene regions of the ophiostomatoid fungi Gondwanamyces and Ophiostoma associated with southern African Protea species. Results indicate that they evolved in close association with Protea. In contrast to Protea, Ophiostoma species migrated to the CFR from tropical and subtropical Africa, where they underwent subsequent radiation. In both Gondwanamyces and Ophiostoma vector arthropods probably facilitated long-distance migration and shorter-distance dispersal. Although ecological parameters shaped most associations between ophiostomatoid fungi and Protea, there is congruence between fungal-host-associations and the systematic classification of Protea. These results confirm that the entire biotic environment must be considered in order to understand diversity and evolution in the CFR as a whole.     

Surface-active arthropods in organic vineyards,integrated vineyards and natural habitat in the Cape Floristic Region

The Cape Floristic Region (CFR), South Africa, is a biodiversity hotspot challenged by intensive wine production. Innovative approaches are being explored to optimize wine production without compromising biodiversity. As organic farming enhances biodiversity conservation in many other regions, the aim here was to assess the potential of organic vineyard management for conserving CFR soil surface arthropod diversity. Pitfall traps were used to sample arthropods in three study areas, each of which included an organic vineyard, an integrated vineyard and a natural vegetation reference habitat. Overall arthropod morphospecies richness was highest in natural sites, followed by organic vineyards and then integrated vineyards. The same trend was seen for predators, saprophages and phytophages. The ability of organic vineyards to sustain more morphospecies than integrated vineyards were partially due to higher non-crop vegetation complexity and less intense management in the organic vineyards. Arthropod assemblages were similar in organic and integrated vineyards, while both land-uses differed greatly from natural sites. Variation among natural vegetation assemblages in different study areas was also much greater than among assemblages of cultivated sites. Organic vineyard management has the potential to make an important contribution to arthropod conservation in the CFR at the field scale. However, at the landscape scale, natural habitat supports a much wider variety of morphospecies, and the preservation of natural fragments in the vineyard landscape may be the most effective measure to increase biodiversity in the winelands.     

Potential impact of viticulture expansion on habitat types in the Cape Floristic Region, South Africa

The wine industry in the Western Cape, South Africa has expanded over the past decade, particularly since the lifting of trade sanctions in 1992. Wine grapes are cultivated on fertile soils upon which threatened biodiversity habitat units of the Cape Floristic Region occur naturally. There is a concern as to whether further expansion of the wine industry, which would benefit the economy through increased foreign exchange, would encroach on the little remaining vegetation in vineyard-producing areas. Predictive land use modeling using logistic regression techniques was applied to determine suitable areas for vineyard cultivation according to climatic, topographic, and soil/geology variables. Of the most threatened habitats, 14849 hectares are particularly suitable for vineyards. Breede fynbos/renosterveld mosaic was the habitat most likely to be converted, and was considered 89.3% irreplaceable to current conservation goals. Also vulnerable are Ashton inland renosterveld and Boland coast renosterveld, the latter being 100% irreplaceable. Although the high rate in vine replanting suggests that the need for untransformed land will not be great immediately, an economic analysis showed that protection of these areas against future ploughing will be vital if targets of adequately representing each habitat in the Cape Floristic Region are to be met. Land use change modeling, especially if done in a spatially explicit and integrated manner with expert input, was shown to be an important technique for the extrapolation of historical patterns to understand the forces that shape landscapes, allowing for the assessment of management alternatives, and testing our understanding of key processes in land use changes that effect conservation planning.     

High ploidy diversity and distinct patterns of cytotype distribution in a widespread species of Oxalis in the Greater Cape Floristic Region

Background and Aims

Genome duplication is widely acknowledged as a major force in the evolution of angiosperms, although the incidence of polyploidy in different floras may differ dramatically. The Greater Cape Floristic Region of southern Africa is one of the world''s biodiversity hotspots and is considered depauperate in polyploids. To test this assumption, ploidy variation was assessed in a widespread member of the largest geophytic genus in the Cape flora: Oxalis obtusa.

Methods

DNA flow cytometry complemented by confirmatory chromosome counts was used to determine ploidy levels in 355 populations of O. obtusa (1014 individuals) across its entire distribution range. Ecological differentiation among cytotypes was tested by comparing sets of vegetation and climatic variables extracted for each locality.

Key Results

Three majority (2x, 4x, 6x) and three minority (3x, 5x, 8x) cytotypes were detected in situ, in addition to a heptaploid individual originating from a botanical garden. While single-cytotype populations predominate, 12 mixed-ploidy populations were also found. The overall pattern of ploidy level distribution is quite complex, but some ecological segregation was observed. Hexaploids are the most common cytotype and prevail in the Fynbos biome. In contrast, tetraploids dominate in the Succulent Karoo biome. Precipitation parameters were identified as the most important climatic variables associated with cytotype distribution.

Conclusions

Although it would be premature to make generalizations regarding the role of genome duplication in the genesis of hyperdiversity of the Cape flora, the substantial and unexpected ploidy diversity in Oxalis obtusa is unparalleled in comparison with any other cytologically known native Cape plant species. The results suggest that ploidy variation in the Greater Cape Floristic Region may be much greater than currently assumed, which, given the documented role of polyploidy in speciation, has direct implications for radiation hypotheses in this biodiversity hotspot.     

Impact of a dam on benthic macroinvertebrates in a small river in a biodiversity hotspot: Cape Floristic Region, South Africa

Suitable reservoirs and monitoring methods are needed to manage scarce water supplies in dry countries. We assessed here the impact on aquatic macroinvertebrates of the only dam on the Eerste River, which runs through the heart of a biodiversity hotspot, the Cape Floristic Region, South Africa. The dam and associated activities, were the only forms of disturbance in this otherwise pristine area. We sampled over 20,000 macroinvertebrate individuals and illustrated some categorical effects of the impoundment and its effects on macroinvertebrate assemblages. Macroinvertebrate species diversity below the dam was only half of that in the pristine catchment area above the dam. Furthermore, Ephemeroptera, Plecoptera and Trichoptera diversity and abundance dropped to almost zero as a result of the impoundment. In contrast, the abundance of the Diptera family Chironomidae increased substantially below the dam. These changes in macroinvertebrate diversity mirrored those recorded in biologically less diverse areas, but are of major concern in this biodiversity hotspot with its rich endemic fauna. We conclude that such an impoundment, while important for human welfare, results in a high price being paid in terms of loss of local biodiversity.     

Coexistence theory in the Cape Floristic Region: revisiting an example of leaf niches in the Proteaceae

Competition in plant communities is often a contentious issue because the mechanisms of competitive interactions are not obvious. We sought evidence that Proteaceae communities are competing along two leaf niche axes as observed in a previous study. Two functional characters, leaf size and leaf shape were measured on numerous individuals per species per plot of six communities from two different regions. Patterns of overdispersion along these leaf trait axes between species were observed, similar to the earlier study. The observed results were compared with the patterns expected under a null model using standard and novel indices to test the significance of trait dispersion between species within a plot. Competition and niche differentiation in the observed plots were not supported as the observed trait overlaps were not significantly different from the null expectation. Our results do not support the theory that Proteaceae communities compete along the two proposed functional leaf traits.     

N and P colimitation of N2-fixing and N-supplied fynbos legumes from the Cape Floristic Region

Background and aims

Legume species in the fynbos vegetation of the Cape Floristic Region, that fix N₂ in soils with low P, may have evolved for enhanced acquisition and efficient use of P. It was hypothesized that N₂-fixing and combined-N supplied (N-supplied) A. linearis, P. calyptrata and C. genistoides are adapted to low P and would be relatively unresponsive to increased P of 100 μM.

Methods

18 legume species were evaluated for their nodulation response to low P availability. The N X P interaction was then examined in A. linearis, P. calyptrata and C. genistoides reliant on either N₂-fixation or 300 μM N (NH₄NO₃), and receiving 0.1, 1.0, 10 and 100 μM P (NaH₂PO₄).

Results

In the species selection experiment, A. linearis, P. calyptrata and C. genistoides, with the greatest nodule fresh weight (FW) and nodule FW to root FW ratio, were the most prolific nodulating species. In the N X P experiment, with low P supply, the biomass of N₂-fixing P. calyptrata and C. genistoides was consistently greater than that of N-supplied plants. In contrast, with high P supply of 100 μM P, all N-supplied plants accumulated more biomass than the corresponding N₂-fixing plants. High P-use efficiency, poor down-regulation of P uptake and P storage was evident in A. linearis and P. calyptrata.

Conclusion

The growth response to P and the significant N X P interactions indicate that N₂-fixing and N-supplied plants were not adapted to low P, but rather colimited by both N and P.     

Immediate impact of piscicide operations on a Cape Floristic Region aquatic insect assemblage: a lesser of two evils?

The piscicide rotenone is used as a conservation tool to remove alien fishes from rivers, though there is controversy over its effects on aquatic insects. An alien fish removal operation in the Rondegat River, Cape Floristic Region, South Africa, allowed the immediate impact of rotenone on an aquatic insect community in a region with high conservation values to be quantified. The insect community within the treated river was sampled in February 2011 (1 year before rotenone operations), February 2012 (1 week before) and March 2012 (1 week after). Insects were collected using kick sampling across multiple biotopes, together with samples from individual stones. We considered rotenone-precipitated losses to be those taxa captured a week before treatment but absent after, and assessed the endemism of lost species to determine the conservation impact of the rotenone. Species richness decreased significantly following treatment, even though many rare taxa were not recorded immediately prior to treatment. Of the 85 taxa identified, 18 were lost including five endemic to the mountain range which the river drains. Ephemeroptera were most severely affected, with a significant loss of density on stones post-rotenone and six out of 20 species missing. Since half the missing taxa were recorded upstream of the treatment area, recovery of diversity is likely to be relatively rapid. Given that alien invasive fish negatively affect both fish and aquatic insect communities in South Africa, the long-term positive conservation impact of removing these fish is likely to outweigh the short-term negative effects of the piscicide.     

Specialised host-use and phenophase tracking in restio leafhoppers (Cicadellidae: Cephalelini) in the Cape Floristic Region

The Cape Floristic Region (CFR) has exceptionally high plant diversity, but because there are so few studies on insect diversity and diet breadth, little is known about the relationship between plant and insect diversity. One possibility is that plant and insect diversity in the CFR are linked through host specialisation. Alternatively, the nutrient-poor soils of the CFR may favour generalist feeding strategies with insects tracking the favourable phenophases of a variety of host plants. We studied Cephalelus, a genus of leafhoppers apparently specialised on the Restionaceae, a diverse and dominant plant family in the CFR. We examined patterns of Cephalelus host association at a single site during a 24 month field survey to determine whether Cephalelus diversity is related to the partitioning of host plant or temporal niches; or whether Cephalelus tracks the most nutritive phenophases of restios by temporal host-switching. Seven Cephalelus species were recorded which varied in their seasonal abundance patterns. The majority of these species exhibited specialised host use on different Restionaceae species, with the exception of C. pickeri. This species specialised on two host plants. The populations of two dominant species, C. pickeri and C.uncinatus, tracked the phenology of their primary host plants but not of the Restionaceae in general. To conclude; we find no evidence for host-switching or generalism in Cephalelus. Instead, they appear to be host-specialised, suggesting coupling between their diversity and that of their host plants; the Restionaceae in the CFR.