Confirmation of pleisiomorphic daily torpor in mammals: the round-eared elephant shrew Macroscelides proboscideus (Macroscelidea)

The characteristics of daily torpor were measured in the round-eared elephant shrew Macroscelides proboscideus (Macroscelidea) in response to ambient temperature and food deprivation. Elephant shrews are an ancient mammal order within a superordinal African clade including hyraxes, elephants, dugongs and the aardvark. M. proboscideus only employed torpor when deprived of food; torpor did not occur under an ad libitum diet at ambient temperatures of 10, 15 and 25?°C. Torpor bout duration ranged from <1?h to ca. 18?h. The times of entry into torpor were restricted to the scotophase, despite normothermic body temperature patterns indicating a rest phase coincident with the photophase. Full arousal was always achieved within the first 3?h of the photophase. When food deprived, the onset of the rest phase, and hence torpor, advanced with respect to the experimental photoperiod. The lowest torpor body temperature measured was 9.41?°C. Daily torpor in M. proboscideus confirms a pleisiomorphic origin of daily heterothermy. Torpor facilitates risk-averse foraging behaviour in these small omnivores by overcoming long-term energy shortfalls generated by the inherent variability of food availability in their semi-arid, El Niño-afflicted habitats.     

Dental Ontogeny in Macroscelides proboscideus (Afrotheria) and Erinaceus europaeus (Lipotyphla)

We investigated the state of dental eruption in specimens of Macroscelides proboscideus and Erinaceus europaeus of known age. When M. proboscideus reaches adult size and sexual maturity, few or none of its replaced permanent cheek teeth have erupted. The approximate sequence of upper tooth eruption is P1, [I3, C, M1], [I1–2], M2, P4, [P2, P3]. Chronologically, E. europaeus erupts its molars and most premolars prior to M. proboscideus; but its first two upper incisors erupt after those of M. proboscideus, and its canines erupt around the same time. The approximate sequence of upper tooth eruption in E. europaeus is [M1, M2, P2, I3], C, M3, P4, P3, I2, I1. Unlike M. proboscideus, E. europaeus does not reach adult size until all permanent teeth except for the anterior incisors have erupted. While not unique among mammals, the attainment of adult body size prior to complete eruption of the permanent cheek teeth is particularly common among macroscelidids and other afrotherians.     

Phylogeny and taxonomy of the round-eared sengis or elephant-shrews, genus Macroscelides (Mammalia, Afrotheria, Macroscelidea)

The round-eared sengis or elephant-shrews (genus Macroscelides) exhibit striking pelage variation throughout their ranges. Over ten taxonomic names have been proposed to describe this variation, but currently only two taxa are recognized (M. proboscideus proboscideus and M. p. flavicaudatus). Here, we review the taxonomic history of Macroscelides, and we use data on the geographic distribution, morphology, and mitochondrial DNA sequence to evaluate the current taxonomy. Our data support only two taxa that correspond to the currently recognized subspecies M. p. proboscideus and M. p. flavicaudatus. Mitochondrial haplotypes of these two taxa are reciprocally monophyletic with over 13% uncorrected sequence divergence between them. PCA analysis of 14 morphological characters (mostly cranial) grouped the two taxa into non-overlapping clusters, and body mass alone is a relatively reliable distinguishing character throughout much of Macroscelides range. Although fieldworkers were unable to find sympatric populations, the two taxa were found within 50 km of each other, and genetic analysis showed no evidence of gene flow. Based upon corroborating genetic data, morphological data, near sympatry with no evidence of gene flow, and differences in habitat use, we elevate these two forms to full species.     

Fungal abundance spike and the Permian-Triassic boundary in the Karoo Supergroup (South Africa)

The most severe mass extinction of marine species and terrestrial vertebrates and plants is associated with the Permian-Triassic boundary (∼251 Ma). The extinction interval is also marked by the disappearance of most Late Permian gymnosperm palynomorphs at a layer containing solely the abundant remains of fungi. This ‘fungal spike’ apparently represents widespread devastation of arboreous vegetation. Stratigraphic and palynological study of the Carlton Heights section in the southern Karoo Basin of South Africa revealed a 1-m-thick fungal spike zone that occurs simultaneously with the last appearance of typically Late Permian gymnosperm pollen. The plant extinction and fungal spike zone are found above the last occurrence of Late Permian mammal-like reptiles of the Dicynodont Zone at other Karoo sections. Using the fungal event as a time line in marine and non-marine sections allows placement of the marine extinctions and the extinction of terrestrial plants and reptiles within a brief crisis interval of less than about 40?000 years at the end of the Permian.     

Identification and quantification of edible plant foods in the Upper (Nama) Karoo,South Africa

More than 18 000 archaeological sites have been identified, and a dozen rock shelter sites investigated in the Zeekoe Valley, Upper Karoo, South Africa, but no work has been performed to date regarding edible plant remains. A baseline for understanding potential contribution of plants to the Zeekoe Valley diet is crucial for developing models of land use and mobility patterns for Late Stone Age (LSA) inhabitants, particularly since ethnographically known foragers in Botswana, some 800 km to the north, are as much as 80% dependent on plant foods for their survival. Since native foraging groups are long extinct in this semi-arid region with abundant natural springs, a rigorous investigation of botanical and ethnohistoric literature forms the groundwork for field investigations where direct ethnographic observation is no longer an option. First, edible species in the study area are identified in the literature. They are then sought on the ground with the aid of local informants whose families have resided in the valley for generations. It appears as though some knowledge of local plants gained from native foragers 150 years ago or more has remained in Afrikaans families passed down from landowning parent to child. On the ground, collection and middle range experimentation was followed by basic nutritional analysis. These measures were then used to compare foraging efficiency to measures from extant foraging groups. While dozens of edible species were identified, five plant species formed the focus of this study: Slymstock Uintjie (Albuca canadensis); Boesman Uintjie (Cyperus usitatus, Cyperus fulgens); Jakkalsbosberry (Diospyros austro-africana); Rooi wortel (Pelargorium sidoides); and Osbossie (Talinum caffrum). Results suggest native foraging groups, now extinct in the vicinity, may have been 60% or more dependent on plant food resources, despite abundance of game in the area and in the archaeological record.     

Environmental History and the Desertification of the Karoo,South Africa

Abstract

This paper reviews the desertification debate in South Africa and emphasizes the methods that have been used to understand the environmental history of the semi–arid rangelands in the eastern Karoo during the last 500 years. These mixed grass/dwarf shrub rangelands, with mean annual rainfall totals typically between 300–400 mm, are described and the main driving variables discussed. A brief historical account of the European settlement of the region is also presented. The desertification debate has focussed on three main issues. Firstly, because arid and semi–arid lands appear to be heavily influenced by climate, the changing climatic regime during the Holocene and particularly during the last two hundred years has been the subject of intensive investigation. While all studies conclude that there is no evidence in the historical record to support the popular perception that rainfall totals have decreased this century, the question of changing rainfall seasonality has not been adequately explored. Although mean annual temperatures over the last 50 years have not changed there have been significant increases in mean monthly maximum temperatures and significant decreases in mean monthly minimum temperatures for some of the stations investigated. The second issue of great interest in the South African desertification literature is that of the nature of pre–colonial environments. A wide variety of archaeological, historical and ecological techniques have been used, including analyses of fossil mammal bones in owl pellets, fossil pollen in hyrax middens, notes from traveller's accounts of the region and stable carbon isotopes in soil as well as fossil mammal bones. While all authors agree that the eastern Karoo was more grassy at some stage in the past there is disagreement as to both the timing and cause of the changes to a more shrubby vegetation. The final issue of great concern to the desertification debate in South Africa concerns the rate of change during the last 100 years. Satellite imagery, matched ground and aerial photography, survey data and an analysis of historical stock records cannot agree as to whether the Karoo is degrading or not. Certainly, the classic view of an annually expanding desert margin has been replaced in recent years by a more realistic understanding of the seasonal dynamics of the vegetation. The recent trend to detailed modelling of the demographic process in key species holds much promise for our understanding of the degradation process. The vibrant community of researchers, employing a range of archaeological, historical and ecological techniques, will make important contributions to South Africa's National Action Plan to Combat Desertification.     

The density and stability of birds in shrubland and drainage line woodland in the southern Karoo,South Africa

Comparative surveys of bird species richness and abundance showed that a total of 69 bird species was recorded on 1 km transects (n = 996) through shrubland, with a total estimated density of birds of 32.7 ± 32.8 (S.D.) birds/km², of which resident birds made up 24.4 ± 21.0 birds/km², nomads made up 1.9 ± 11.7 birds/km² and local nomads made up 6.3 ± 20.1 birds/km². There was no correlation in general between total bird numbers or total bird biomass and rain, but numbers and biomass of nomads showed the strongest correlations with rainfall two months previously. A total of 86 species was recorded on 276 surveys along a 1 km stretch of drainage line woodland. Species richness varied from an average of about nine species to 24 species each month, and only showed a marked change during the 'dry' period of 1990 to early 1991. Density of birds in this woodland was 59.2 ± 20.8 birds/km. Biomass of birds was relatively constant throughout the study period, with marked increases in certain months when large species were present.     

Trophic status of small mammals in the semi-arid Karoo, South Africa

The feeding habits of a community of small mammals from the semi-arid Karoo, South Africa were analysed by microscopic examination of the stomach contents of animals caught during a 13-month snap-trapping study. The community comprised eight rodent species ( Gerbillurus paeba, Mus minutoides, Rhabdomys pumilio, Otomys unisulcatus, Saccostomus campestris, Desmodillus auricularis, Malacothrix typica, Mastomys natalensis ) and a single macroscelid elephant shrew ( Macroscelides proboscideus. ) The rodents were all predominantly herbivorous, while the elephant shrew ate mainly insects. No granivores or omnivores were represented in this community. The results presented here (including the first analysis of the diet of M. typica ) indicate a greater emphasis on herbivory than previously described for these species. Limited dietary overlap was recorded between these species, although some instances of potential dietary competition that deserve further study were identified. The paucity of granivorous small mammals in the semi-arid Karoo contrasts the patterns of trophic specialization for North American and Israeli semi-arid communities, but is similar to that of South American, Australian and other southern African semi-arid communities.     

Evaluation of Restoration Techniques for the Succulent Karoo, South Africa

Abstract Possible constraints on the passive recovery of bare areas in the Karoo, a semiarid region in South Africa, include inadequate supply of seed, availability of suitable microsites for plant establishment, altered soil properties, and the truncation of key soil biotic processes. Here we investigate the possibility of initiating the restoration of bare areas by soil surface treatments with gypsum (CaSO₄) and/or organic mulch. We also apply an exogenous seed source to test the hypothesis that seed availability limits autogenic recovery. Both gypsum and mulch improved rain water infiltration, gypsum more so than mulch, and both treatments resulted in significantly higher numbers of reseeded seedlings compared with controls. Gypsum also improved the survival of the cohorts of seedlings of the larger seeded Tripteris sinuata. Tripteris showed the highest number of seedlings (maximum count of 150 seedlings/1,000 viable seeds sown) and surviving plants of the three reseeded species, which included two small‐seeded species, Ruschia spinosa and Chaetobromus dregeanus. Throughout the study period significantly higher plant volumes of naturally seeded annuals and perennials were recorded in the gypsum and/or mulch treatments compared with the controls. Germination and emergence of reseeded and naturally seeded plants appears to be determined by the availability of cool season (autumn to spring ) soil moisture, whereas follow‐up rainfall during this time is important for plant survival. Mulching of bare areas in the Succulent Karoo has the potential to re‐create vegetated areas that will further capture and conserve water, soil, and nutrients. Gypsum also showed positive results but might not be a cost‐effective option because of transport costs to these remote arid areas.     

The effect of feeding immature Karoo paralysis ticksIxodes rubicundus (Acari: Ixodidae) on the metabolic rate of the rock elephant shrew

The effect of feeding immature Karoo paralysis ticks (Ixodes rubicundus) on the resting metabolic rate (RMR) of their principal natural host, the rock elephant shrew (Elephantulus myurus), was investigated under laboratory conditions. The elephant shrews were artificially infested with numbers of ticks simulating natural burdens. The RMR of the elephant shrews was quantified by measuring the oxygen consumption in an open through-flow system. The RMR of hosts infested only with larvae did not differ from that of the control group. Those infested with nymphs, or nymphs and larvae, had significantly lower RMR's compared to the control animals. There were no signs of paralysis in any of the infested hosts.     

The distribution and biology of nomadic birds in the Karoo, South Africa

Dryland nomadic bird species, as a proportion of all bird species in a biome in southern Africa, are highest in the arid grassland and arid and semi-arid Karoo in South Africa. Nomadic birds, of which the most widespread species is the greybacked finchlark Eremopterix verticalis (Smith), are most frequently observed in the north-central and north western Nama Karoo. The species richness of nomadic species is inversely correlated with species richness of all bird species in the Karoo. Since the distribution of nomadic birds is in areas where rainfall is patchy, low (<250 mm per year) and aseasonal, this supports the idea that fewer species are able to cope with resources that are patchy in time and space, and that there has been selection for nomadism in the species that are able to use patchy environments. Species richness and abundance of nomadic birds is negatively correlated with rainfall amount but positively correlated with the coefficient of variation of the rainfall and with rainfall in autumn. The frequency of nomadic birds is inversely correlated with altitude range; nomadic species are most often recorded in structurally simple habitats (shrubland and grassland) on open plains. Most nomadic bird species in the Karoo are granivorous. Perennial desert grasses are important components of the habitat and diet of small nomadic granivores, and also provide nest sites and nest material. Nomadic birds can breed throughout the year, without a clearly defined ‘season’ in both the Succulent and Nama Karoo. Average clutch sizes do not differ significantly between resident and all nomadic species in the arid and semi-arid Karoo. Nomadism is an evolutionary stable strategy for individual species only when extremes in environmental conditions are frequent enough, and unpredictable enough, to maintain movements to high resource patches or to maintain dispersal away from low resource patches. If high rainfall years are too regular or infrequent, or peaks in fluctuations of resources in the environment too low, or rainfall patches are randomly distributed, nomadism would not be maintained as part of the individual behaviour pattern.     

Relationship between soil-stored seed banks and degradation in eastern Nama Karoo rangelands (South Africa)

Landscape, local scale and seasonal composition of soil-stored seed banks of a southern African semi-arid rangeland (Nama Karoo) were investigated through assessment of seed germinated from field-collected soil. Variables included percentage above-ground canopy cover, numbers of seeds germinated, and selected physical and chemical soil characteristics. Organic matter content of soil sampled from under plant canopies (closed-canopy) is significantly higher than that of inter-canopy samples (open-canopy). Significantly more seeds germinated from closed-canopy soil samples are compared with those from open-canopy microsites. It is argued that reduction in rangeland canopy cover due to overgrazing (and resultant erosion through wind and water movement) leads to reduced seed retention by the leaf litter layer and thus by the soil. This degraded resource appears to have gradually lost an inherent buffer capacity, undermining attempts at ecological restoration and necessitating concerted and directed efforts to restore these fragile systems.     

Changes in Permo-Triassic terrestrial tetrapod ecological representation in the Beaufort Group (Karoo Supergroup) of South Africa

Nicolas, M. & Rubidge, B.S. 2010: Changes in Permo-Triassic terrestrial tetrapod ecological representation in the Beaufort Group (Karoo Supergroup) of South Africa. Lethaia, Vol. 43, pp. 45–59.
For more than a century, large collections of fossils from the Beaufort Group have been built up at various museums in South Africa and have been handled as separate databases in the individual museums. Because of the unique time-extensive record of continental vertebrate biodiversity represented by the fossils of the Beaufort Group, a single standardized database has been compiled for the fossils collected from the Beaufort Group housed in South African museums. This unique data set has enabled the determination of terrestrial tetrapod ecological representation from the Middle Permian to Middle Triassic Beaufort Group of South Africa. □ Beaufort Group , biodiversity , ecology , Permo-Triassic .     

Pollination of sevenPlectranthus spp. (Lamiaceae) in southern Natal, South Africa

The genusPlectranthus (Lamiaceae) shows remarkable radiation on the sandstones of southern Natal and northern Transkei in South Africa, where six endemic species occur. Two of these endemic species,P. hilliardiae andP. oribiensis, are included in this study, as well asP. reflexus, for which only limited data are available. The other species that were studied areP. ambiguus, P. ciliatus, P. ecklonii, P. madagascariensis andP. zuluensis. Four of these taxa,P. ambiguus, P. hilliardiae, P. reflexus andP. saccatus var.longitubus, have uniquely long corolla-tubes (20–30mm) and this is related to pollination by nemestrinid flies of the genusStenobasipteron that have proboscides of similar length. Other nemestrinid species of the genusProsoeca have shorter proboscides and pollinate two species ofPlectranthus with shorter corolla-tube lengths (6–15mm). Acrocerid flies, tabanid flies and anthophorid bees are also important visitors to these species. This study on the pollination of seven species of varying corolla-tube lengths shows a correlation between floral tube length and proboscis length of insect visitors, many of which are recorded for the first time as pollinators ofPlectranthus.