Seed storage and germination in Kumara plicatilis,a tree aloe endemic to mountain fynbos in the Boland,south-western Cape,South Africa

Seed storage under appropriate conditions is a relatively inexpensive means of safeguarding plant genetic material for ex situ conservation. Post-storage germination trials are used to determine the viability of stored seeds, and hence the efficacy of the particular storage treatment. Kumara plicatilis (= Aloe plicatilis) is a tree aloe endemic to mountain fynbos in the Boland, south-western Cape. The viability and germination behaviour of K. plicatilis seeds were assessed for seeds stored for four and nine months at − 80 °C, 4 °C, 25 °C and under ambient conditions in a laboratory. Seeds were germinated under controlled conditions and germination rates and percentages determined. Ungerminated seeds were tested for viability using tetrazolium salt. Seed viability was not significantly reduced during storage. Seeds stored at − 80 °C for four and nine months exhibited the fastest germination rate overall (both 5.9 ± 0.3 weeks, mean ± S.E.), and slowest was for seeds stored under ambient conditions for four and nine months (both 7.8 ± 0.4 weeks). All seed lots showed similar percentage germination after four months of storage (78.0–90.4%). The highest percentage germination overall was for seeds stored at − 80 °C for four months (90.4%) and the lowest was for seeds kept at 4 °C and − 80 °C for nine months (39.2 and 39.6%, respectively). Respective percentage viability for ungerminated seeds in these two treatments was 82% and 87%, respectively, indicating the induction of secondary dormancy. Induced dormancy triggered by protracted cold temperatures may be an adaptation that enables seeds to survive prolonged extreme conditions that are unfavourable for germination. Further research on the long-term storage of aloe seeds would be beneficial for developing long-term seed storage and germination testing protocols for ex situ conservation.     

Reproductive ecology of Aloe plicatilis,a fynbos tree aloe endemic to the Cape Winelands,South Africa

While the pollination ecology of many Aloe species is well-documented, knowledge on aloe seed ecology, and hence aloe reproductive ecology in its entirety is limited. The aim of this study was to investigate the reproductive ecology of Aloe plicatilis, a Cape fynbos tree aloe endemic to the Cape Winelands, South Africa. Results from a pollinator exclusion experiment conducted at an A. plicatilis population on Paarl mountain suggests pollination primarily by insects, although bird visitation significantly increased seed set/fruit indicating possible co-pollination with insects. The species' long-tubed flowers and production of concentrated nectar, with observations of malachite sunbirds as the most common avian visitors to A. plicatilis flowers indicate the importance of long-billed specialist avian nectarivores as floral visitors. Analysis of the relationship between plant size and inflorescence production for five populations combined revealed a significant, positive linear relationship between plant size and the logarithm of the number of inflorescences/plant. Natural fruit and seed set determined for three populations (1325, 27,930 and 251,616 seeds/population) suggests low reproductive output compared to several other Aloe species. The smallest (31 individuals) and least dense (75 plants/ha) A. plicatilis population produced the lowest seed set/plant (128 seeds) and per population (1325 seeds), suggesting an Allee effect. Evaluation of seed dispersal potential showed that potential dispersal distances were approximately three times the canopy height; however, the occurrence of A. plicatilis on mountains isolated from more continuous mountain ranges on which the species also occurs suggests the possibility of long-distance dispersal by strong, gusty, summer winds. Soil seed bank samples collected from 13 populations yielded close to zero seedling emergence, indicating the absence of persistent seed banks. A. plicatilis seeds stored under ambient laboratory conditions for 3, 18 and 24 months were germinated in an environmental control chamber and a laboratory. High percentage germination was recorded for 18- and 24-month-seed (86 and 80%, respectively), while germination of 3-month-old seeds was three times lower, suggesting the need for after-ripening. Germination of fresh and one-year-old seed under ambient nursery conditions at the Karoo Desert National Botanical Garden in Worcester yielded emergence percentages of 67 and 44%, respectively, and were therefore less successful than germination under more controlled conditions. This is the first known study to investigate the reproductive ecology of a tree aloe species and that of a Cape fynbos aloe. The study highlights the importance of further studies on aloe seed ecology, particularly for rare and threatened species.     

Species turnover of monkey beetles (Scarabaeidae: Hopliini) along environmental and disturbance gradients in the Namaqualand region of the succulent Karoo, South Africa

Species turnover of monkey beetle (Scarabaeidae: Hopliini) assemblages along disturbance and environmental gradients was examined at three sites within the arid, winter rainfall Namaqualand region of the succulent Karoo, South Africa. At each site two study plots with comparable vegetation and soils but contrasting management (grazing) histories were chosen, the disturbed sites having fewer perennial shrubs and generally more annuals and bare ground. Beetles collected using coloured pan-traps showed a consistently higher abundance in disturbed sites. Lepithrix, Denticnema and Heterochelus had higher numbers in disturbed plots, while Peritrichia numbers were lower in disturbed areas. Measures of species richness and diversity were consistently higher in the undisturbed sites. Distinctive assemblages of monkey beetles and plants occurred at each site. A high compositional turnover ( diversity) was recorded for both monkey beetles and plants along a rainfall gradient; between-site diversity values ranged from 0.7 to 0.8 (out of a maximum of 1.0). Species turnover of beetles was higher between the disturbed sites along the environmental gradient than the corresponding undisturbed sites. The high monkey beetle species turnover is probably linked to the high plant species turnover, a distinctive feature of succulent Karoo landscapes. Monkey beetles are useful indicators of overgrazing disturbance in Namaqualand, as their pollinator guilds are apparently disrupted by overgrazing. A shift away from perennial and bulb pollinator guilds towards those favouring weedy annuals was observed in disturbed areas. The consequences to ecosystem processes due to the effects of disturbance on monkey beetle communities and the role of monkey beetles as indicators of disturbance is discussed, as well as the implications of disturbance on monkey beetle pollination guilds.     

Relevance of elephant herbivory as a threat to Important Plants in the Addo Elephant National Park, South Africa

Although elephants are recognized as keystone species, the mechanisms of their impacts on biodiversity and community structure are rarely identified. In the Addo Elephant National Park (AENP), South Africa, elephant Loxodonta africana herbivory is apparently responsible for a significant reduction in plant richness, especially among the regionally rare and endemic small succulent shrubs and geophytes (Important Plants). We used faecal analysis to investigate the utilization of Important Plants in elephant diet in the AENP. Ninety plant species were identified in the diet. Only 14 of the 77 ( c . 18%) Important Plants previously thought particularly vulnerable to elephant browsing occurred in the diet, while at least 6% of species for which there are data were avoided. This refutes the generally held belief that elephant herbivory is the major driver of decline among Important Plants, and emphasizes the likely contribution of other mechanisms (e.g. knock-on effects, trampling, zoochory, etc.) to this phenomenon. The accurate prediction of impacts caused by elephants in the AENP and elsewhere, therefore requires an understanding of these previously marginalized mechanisms. By demonstrating appropriate cause-and-effect relationships between elephants and ecosystem change, we will be able to move beyond assuming that all the observed changes are due to elephant herbivory.     

Annesorhiza calcicola (Apiaceae), a new limestone endemic species from the Western Cape Province of South Africa

The new species, Annesorhiza calcicola Magee and J.C. Manning, is described. It is known only from a few limestone outcrops around Jacobsbaai on the Saldanha Peninsula on the West Coast of South Africa. The species differs from all others in the genus in its small stature (inflorescence less than 0.3 m and leaves less than 150 mm long), the hysteranthous, pilose leaves which are completely absent at flowering (in the other species dead or dying at flowering and usually glabrous), the greyish-black, pilose inflorescence (yellowish-brown and glabrous in the other species) and the smaller homomericarpic fruit less than 5 mm long.     

Profiling a besieged flora: endemic and threatened plants of the Cape Peninsula, South Africa

The Cape Peninsula (area: 471 km²), situated at the south-western extremity of the Cape Floristic Region, has exceptionally high plant species richness (2285 species and infraspecific taxa) and numbers of endemic (90; 88 species and two infraspecific) and threatened (141; 138 species and three infraspecific) taxa (termed species from here on). This biodiversity is threatened by urban development and the spread of invasive alien plants. Peninsula endemics are concentrated in a few, predominantly species-rich families and these correspond well with endemic-rich families in other areas of the Cape Floristic Region. A high level of similarity exists between families with threatened and families with endemic species. A frequency analysis of the biological traits of both endemic and threatened species shows that low growing, ant-dispersed shrubs are over-represented in both groups. Endemics are most likely to be non-sprouters, but threatened plants do not have a specific post-fire regeneration strategy. Threatened species have higher frequencies of geophytes, sprouters and wind-dispersed species compared to endemic species. Numbers of endemic and threatened species are not randomly distributed with regard to occurrence in vegetation types and patterns are similar for both groups. The habitat and biological profiles of both endemic and threatened species suggest that they are highly vulnerable to extinction as a result of increasing rates of alien plant infestation, urbanization and inappropriate fire regimes.     

Physiological responses to folivory and phytopathogens in a riparian tree,Brabejum stellatifolium,native to the fynbos biome of South Africa

The canopies of many tree species sustain a large diversity of folivorous arthropods and phytopathogenic fungi. These organisms are thought to influence overall tree and stand productivity. Leaf diseases caused by Phyllosticta owaniana and Periconiella velutina, phytopathogenic fungi commonly found on the native riparian tree Brabejum stellatifolium (wild almond), like any other leaf disease, can potentially reduce a plant's photosynthetic efficiency. In addition to these two phytopathogens, the weevils Setapion provinciale and Setapion quantillum are abundant in wild almond canopies. Despite their pervasive occurrence, the impacts of these phytopathogens and arthropods on host tree leaf physiology have not been examined. The gas exchange response of wild almond leaves to phytopathogens and folivore damage was assessed. Leaf nitrogen, phosphorus and water content were also determined. Declines in photosynthetic rates and other physiological parameters were associated with increasing damage severity by weevils and phytopathogens in leaves of B. stellatifolium. Nitrogen and phosphorus contents were negatively associated with disease severity. Water and phosphorus contents were also negatively correlated with increased weevil damage, while nitrogen content was positively correlated with it. The observed responses of B. stellatifolium metabolic functioning to fungal pathogen and folivory indicate a possibility of suppressed wild populations of wild almond.     

Between an ocean and a high place: coastal drainage isolation generates endemic cryptic species in the Cape kurper Sandelia capensis (Anabantiformes: Anabantidae), Cape Region,South Africa

This study investigated the range-wide phylogenetics and biogeography of the Cape kurper Sandelia capensis, a primary freshwater fish endemic to and widespread within the Cape Floristic Region (CFR) of South Africa. Maximum likelihood, Bayesian phylogenetic and haplotype network analyses, based on two mitochondrial and two nuclear genes, revealed the existence of three reciprocally monophyletic, deeply divergent and allopatric clades that probably represent cryptic species. The West Coast Clade is largely confined to the Langvlei, Verlorenvlei, Berg and Diep Rivers, the Klein River Clade is endemic to the Klein River and the South Coast Clade is found everywhere else in the range of S. capensis sensu lato. It was hypothesised that divergences within S. capensis sensu lato probably occurred because of isolation of coastal drainages by persistent drainage divides or vicariance of current tributaries by the drowning of their confluences by high sea levels. The current distribution of lineages could be due to historical range expansion and gene flow via river capture or some other mode of transdivide dispersal or dispersal during periods of low sea level via palaeoriver confluences of currently isolated coastal rivers. Comparison of BEAST2 estimated divergence times with the timing of climatic, geological and geomorphological events supported long-term coastal drainage isolation, punctuated by rare transdivide dispersal events and limited palaeoriver dispersal, as the best explanation of current phylogeographic and divergence patterns in S. capensis. Hydrological barriers that block upstream passage in palaeotributaries could hypothetically explain why S. capensis failed to disperse through certain palaeoriver confluences. There were several sites where biogeographic patterns have likely been confounded by human translocation of S. capensis. Alien fish predators and water extraction may threaten the three cryptic species more severely than previously realised, due to their smaller population sizes and inhabitation of only a portion of the range previously ascribed to S. capensis sensu lato. The preponderance of cryptic diversity and endemism in the CFR suggests that additional undescribed cryptic species of obligate freshwater fishes may be found in short coastal river systems around the world, especially in regions with a history of geological stability and a narrow continental shelf.     

Palystes kreutzmanni sp. n. - a new huntsman spider species from fynbos vegetation in Western Cape Province, South Africa (Araneae, Sparassidae, Palystinae

Palystes kreutzmannisp. n. is described from habitats close to Kleinmond, in the Western Cape Province, South Africa. Spiders of this new species live in the typical fynbos vegetation of the Western Cape region. They build retreats between apical leaves of Leucadendron bushes. The systematic position of Palystes kreutzmannisp. n. is discussed. Male and female show characters of different species groups, especially the female copulatory organ seems to be unique within the genus Palystes L. Koch, 1875.     

<Emphasis Type="Italic">Isoetes eludens</Emphasis> (Isoetaceae), a new endemic species from the Kamiesberg,Northern Cape,South Africa

Summary   Isoetes eludens, a new, highly localised, endemic species from the Kamiesberg, Northern Cape is described. The aquatic species grows in a shallow temporary gnamma or !gau (rock pool) in Kamiesberg gneiss at an altitude of 1,284 m. It is characterised by a short, strongly 3-lobed stem, relatively short, and broad estomate sporophylls, a complete velum, the presence of an outer sporangium sack (a feature not previously reported for the genus), graphite-black, laevigate megaspores with a narrow, but tall laesura, microspores with near smooth proximal surfaces but with a papillate distal surface, the absence of a ligule and labium, and simple roots with few root-hairs. The systematic relationship of the species is difficult to determine as a result of convergent evolution in the genus. The sporangium sack does not support an affinity with any of the known species. Further surveys are needed to establish the geographical extent and conservation status of this remarkable local endemic.     

Fire season effects on the recruitment of non‐sprouting serotinous Proteaceae in the eastern (bimodal rainfall) fynbos biome,South Africa

Abstract Research in Mediterranean‐climate shrublands in both South Africa and Australia shows that recruitment of proteoid shrubs (non‐sprouting, serotinous Proteaceae) is best after warm‐season (summer and autumn) fires and worst after cool‐season (winter and spring) ones. This pattern has been attributed to post‐dispersal seed attrition as well as size of pre‐dispersal seed reserves. Here we investigate patterns of post‐fire recruitment for four proteoid species in the eastern part of South Africa's fynbos biome, which has a bimodal (spring and autumn) rainfall regime. Despite the lack of significant differences in recruitment between cool‐ and warm‐season burns, we find some evidence for favourable recruitment periods following fires in spring and autumn, immediately before, and coinciding with, the bimodal rainfall peaks. This suggests that enhanced recruitment is associated with conditions of high soil moisture immediately after the fire, and that rapid germination may minimize post‐dispersal seed attrition. In two of the species, we also find a shift from peak flowering in winter and spring in the Mediterranean‐climate part of the fynbos biome, to summer and autumn flowering in the eastern part. Because these two species are only weakly serotinous, warm‐season flowering would result in maximal seed banks in spring, which could explain the spring recruitment peak, but not the autumn one. We conclude that eastern recruitment patterns differ significantly from those observed in the western and central parts of the biome, and that fire management protocols for the east, which are currently based on data and experience from the winter‐rainfall fynbos biome, need to be adjusted accordingly. Fire managers in the eastern fynbos biome should be less constrained by requirements to burn within a narrow seasonal range, and should therefore be in a better position to apply the required management burns.     

Persisting in a windy habitat: population ecology and behavioral adaptations of two endemic grasshopper species in the Cape region (South Africa)

Global biodiversity hotspots are rich in endemic insect species, many of which are threatened by the ongoing anthropogenic pressures on their habitats. The Cape region (South Africa) is one of these biodiversity hotspots, maintaining a high number of endemics. However, the ecology of most insect species in this region remains poorly understood. The two Orthoptera species Betiscoides meridionalis and Betiscoides parva are endemic to the Cape region and specialized on restio vegetation. They are threatened by increasing wildfire frequencies and invasions of non-native plant species. However, this information has been inferred from habitat changes, whereas no ecological study on these species has been conducted since they have been described. In order to facilitate conservation management, information on the ecology of these species is urgently required. The aim of our study was (1) to obtain data on the population ecology (particularly population sizes and mobility), and (2) to study the behavior of both species in response to environmental factors. For this purpose a mark-recapture-study and an observational behavior study were conducted. Both species had small population sizes and a low mobility with males moving greater distances than females. Wind had a strong influence on the behavior of Betiscoides, particularly on the small males of B. parva. Future studies might thus focus on the question whether wind-exposure is a critical factor for habitat choice of this species. We strongly recommend enhancing the connectivity of restio habitats and restoring these habitats to prevent extinction of specialized insect species.     

Pelargonium elandsmontanum (Geraniaceae: section Hoarea), a new species from the Western Cape Province, South Africa

The new species Pelargonium elandsmontanum is a local endemic from Elandsberg Nature Reserve near Malmesbury in the Western Cape Province, South Africa. One of six species of sect. Hoarea with just the posterior two petals developed, it resembles P. ternifolium in its trifoliolate leaves and pink petals but is distinguished from that species by the short, stout petioles, rhombic, acute leaflets with the upper surface glabrous or with spreading hairs (vs cuneate, apically incised leaflets with both surfaces adpressed-hairy), and five (vs four) fertile stamens.     

Fossil wood charcoal assemblages from Elands Bay Cave, South Africa: implications for Late Quaternary vegetation and climates in the winter-rainfall fynbos biome

Aim The aim of this paper is to analyse fossil charcoal deposits, largely identified to the species level and spanning a sequence from the late Holocene to < 40,000 BP , in order to reconstruct Late Quaternary vegetation and climatic patterns in the western (winter-rainfall) fynbos biome of South Africa. Location The charcoals were excavated from the Elands Bay Cave (32°19S, 18°20E) on the semiarid (200–250 mmyr^?1), winter-rainfall coastline of the western fynbos biome. Methods Patterns in the charcoal data set over time were sought by manual sorting of the charcoal×sample matrix, as well as by subjecting the data to multivariate analysis. Palaeoclimatic reconstruction was attempted by comparing the climatic controls on contemporary vegetation communities that resembled the fossil assemblages. Charcoal diversity was modelled using sample age and number of charcoal fragments as explanatory variables. Results The fossil assemblages ranged from xeric communities (similar to those presently occurring at the site) during the Holocene, to more mesic thicket and fynbos vegetation in the terminal Pleistocene, to Afromontane forest and riverine woodland communities after about 18,000 BP . Diversity of the charcoal samples increased monotonically with increasing sample age. Main conclusions The results suggest that, unlike the eastern fynbos biome, which is under fundamentally different climatic controls, soil moisture conditions in the western part of the biome were higher in the Last Glacial than during the Holocene. This scenario may help to explain the higher regional richness and associated diversification in the western than eastern part of the biome.     

Disa albomagentea (Orchidaceae), a new species from the Hottentots Holland Mountains in the Cape Floristic Region, South Africa

A new orchid, Disa albomagentea, is described from the Hottentots Holland Mountains in the Western Cape Province of South Africa. It is an obligately fire-dependent species closely related to Disa obtusa (sect. Disella), from which it can be easily distinguished by the larger flowers, i.e. galea 7-9 mm long versus 4-7 mm long, which are patent and bicoloured, i.e. magenta, white and not spotted versus purple brown, white to cream to greenish, and variously purple spotted. It appears to be endemic to seeps and marshes on the high peaks.     

Variable wood formation and adaptation to the alpine environment of Ephedra pachyclada (Gnetales: Ephedraceae) in the Mustang district, western Nepal

BACKGROUND AND AIMS: Plants of Ephedra normally have vessels, but are known to become nearly vessel-less in some alpine localities. Previous studies implied that wood formation in Ephedra differs fundamentally from that in dicotyledons in which vessel-bearing and vessel-less taxa are systematically distinct. Using E. pachyclada in the Mustang district of Nepal, growing in an altitudinal range of over 2000 m, variation in wood formation and adaptation to alpine environment was studied in this normally vessel-bearing species. METHODS: Variation in wood anatomy and wood formation was observed with conventional optical microscopy. The lengths of three kinds of tracheary elements were measured and statistically analysed against habitat altitude and plant size of the individuals studied. KEY RESULTS: In E. pachyclada three kinds of tracheary elements, vessel elements, tracheids and fibre-tracheids, were nearly equal in length within individuals showing no elongation after differentiation from cambial initials. Tracheary element lengths among individuals had a negative correlation with altitude and a positive correlation with plant size. Multivariate analyses showed that altitude has a stronger correlation with tracheary element lengths than plant height or stem diameter. Moreover, several individuals from high elevations completely lacked vessels, and vessel formation fluctuated even in individuals from lower elevations. CONCLUSIONS: Wood anatomical trends in E. pachyclada are considered as an adaptation to extremely dry conditions in high mountains. Fluctuation in vessel formation in individuals from low elevations indicated that vessels differentiate only when their lateral expansion is allowed. These results showed that E. pachyclada has a different system of wood formation from dicotyledons and supported the opinion that the wood structure of Gnetales is fundamentally different from that of angiosperms.     

Isolepis namaquana (Cypereae, Cyperaceae), a new endemic species from the winter rainfall area of South Africa

A new species, Isolepis namaquana Muasya & J.Viljoen, is described and illustrated. It is known from two localities in Namaqualand: near Kamieskroon and at the foot of the Matsikammaberge, where it grows in ephemeral wetlands. The tufted annual species is diagnosed by floral and fruit characters including glume size, bifid stigma and colliculate nutlet surface ornamentation.     

Pelargonium saxatile (Geraniaceae: Section Hoarea), a new species from the southwestern Cape,South Africa,and a key to the species of the P. dipetalum group

The new species Pelargonium saxatile is a local endemic of sandstone rock pavement in the Elandskloofberge near Wellington in Western Cape Province, South Africa. One of just seven species of sect. Hoarea that have only the posterior two petals developed, it most closely resembles Pelargonium dipetalum from the southern Cape coastal lowlands but is distinguised by its rosette of numerous, small, mainly simple leaves, glabrous above and strigose beneath only along the midrib and sometimes also the veins. In addition, the pink flowers have the dorsal filament shorter, 4–5 mm long. We include a key to the species of the P. dipetalum group.     

Fire‐induced tree mortality in a neotropical forest: the roles of bark traits,tree size,wood density and fire behavior

Large‐scale wildfires are expected to accelerate forest dieback in Amazônia, but the fire vulnerability of tree species remains uncertain, in part due to the lack of studies relating fire‐induced mortality to both fire behavior and plant traits. To address this gap, we established two sets of experiments in southern Amazonia. First, we tested which bark traits best predict heat transfer rates (R) through bark during experimental bole heating. Second, using data from a large‐scale fire experiment, we tested the effects of tree wood density (WD), size, and estimated R (inverse of cambium insulation) on tree mortality after one to five fires. In the first experiment, bark thickness explained 82% of the variance in R, while the presence of water in the bark reduced the difference in temperature between the heat source and the vascular cambium, perhaps because of high latent heat of vaporization. This novel finding provides an important insight for improving mechanistic models of fire‐induced cambium damage from tropical to temperate regions. In the second experiment, tree mortality increased with increasing fire intensity (i.e. as indicated by bark char height on tree boles), which was higher along the forest edge, during the 2007 drought, and when the fire return interval was 3 years instead of one. Contrary to other tropical studies, the relationship between mortality and fire intensity was strongest in the year following the fires, but continued for 3 years afterwards. Tree mortality was low (≤20%) for thick‐barked individuals (≥18 mm) subjected to medium‐intensity fires, and significantly decreased as a function of increasing tree diameter, height and wood density. Hence, fire‐induced tree mortality was influenced not only by cambium insulation but also by other traits that reduce the indirect effects of fire. These results can be used to improve assessments of fire vulnerability of tropical forests.     

Exploring patterns of population subdivision in the net-winged midge, Elporia barnardi (Diptera: Blephariceridae), in mountain streams of the south-western Cape, South Africa

1. The net-winged midges (Diptera: Blephariceridae) are a highly specialized group whose morphological characteristics and specific habitat requirements suggest a limited potential for dispersal. Levels of genetic variation were examined within streams, between streams in the same range and between mountain ranges in larval populations of Elporia barnardi in the south-western Cape of South Africa. The aim was to examine the hypothesis that population structure would reflect the poor potential for dispersal.
2. Significant deviations from Hardy–Weinberg equilibrium in 17 of the 57 individual comparisons indicate a non-random mating population. Given the swarming behaviour and life history traits, larvae sampled may reflect the product of limited matings.
3. Analysis of population substructuring revealed significant levels of differentiation among geographically proximate populations. Large differences between streams within Table Mountain, similar in magnitude to those between mountain ranges, suggest that movement of individuals out of the stream catchment is rare. Observed F _ST values are more similar to those of fully aquatic species than other lotic insects with winged adult stages.
4. Results suggest that mountain ridges provided effective physical barriers to the dispersal of E. barnardi , with the catchment representing the effective population unit.