Alien phytogeographic regions of southern Africa: numerical classification, possible drivers, and regional threats

The distributions of naturalised alien plant species that have invaded natural or semi-natural habitat are often geographically restricted by the environmental conditions in their new range, implying that alien species with similar environmental requirements and tolerances may form assemblages and characterise particular areas. The aim of this study was to use objective numerical techniques to reveal any possible alien phytogeographic regions (i.e. geographic areas with characteristic alien plant assemblages) in southern Africa. Quarter degree resolution presence records of naturalised alien plant species of South Africa, Lesotho, Swaziland, Namibia and Botswana were analysed through a divisive hierarchical classification technique, and the output was plotted on maps for further interpretation. The analyses revealed two main alien phytogeographic regions that could be subdivided into eight lower level phytogeographic regions. Along with knowledge of the environmental requirements of the characteristic species and supported by further statistical analyses, we hypothesised on the main drivers of alien phytogeographic regions, and suggest that environmental features such as climate and associated biomes were most important, followed by human activities that modify climatic and vegetation features, such as irrigation and agriculture. Most of the characteristic species are not currently well-known as invasive plant species, but many may have potential to become troublesome in the future. Considering the possibility of biotic homogenization, these findings have implications for predicting the characteristics of the plant assemblages of the future. However, the relatively low quality of the dataset necessitates further more in-depth studies with improved data before the findings could be directly beneficial for management.     

Developing wetland inventories in southern Africa: A review

The status of wetland inventory effort and availability of maps and other data sources is reviewed for the ten countries of southern Africa: Angola, Bostwana, Lesotho, Malawi, Mozambique, Namibia, South Africa, Swaziland, Zambia and Zimbabwe. The aims and strategies for inventory are discussed and the main survey methods compared. Prior to commissioning new inventory work careful collation of existing maps and imagery is recommended together with targeting of strategic inventory at Province level, reserving high resolution effort only for certain important sites.     

A revision of the genus Dolichos (Fabaceae,Papilionoideae, Phaseoleae), including Lesotho and Swaziland

A revision of the genus Dolichos in South Africa (Lesotho and Swaziland included) is presented. This legume genus, belonging to the bean tribe Phaseoleae, mainly has an African distribution, extending into Asia. In South Africa it is represented by nine species, two (D. sericeus and D. trilobus) of which extend into Tropical Africa. Dolichos is closely related to the genus Macrotyloma from which it can be distinguished by the short standard appendages, reticulate pollen and the generally purple flowers (standard appendages long, pollen tuberculate or spinulose and flowers yellow or orange in Macrotyloma). It also has affinities with the genera Dipogon and Lablab. The correct nomenclature, as well as complete synonymy, typification and distribution maps of all the species are provided.     

<Emphasis Type="Italic">Athyrium crassicaule</Emphasis> (<Emphasis Type="Italic">Woodsiaceae</Emphasis>), a new fern species from the Drakensberg,southern Africa

Summary  A new Athyrium species, A. crassicaule J. P. Roux (Woodsiaceae), is described from the Drakensberg in Lesotho and South Africa. A key to the Athyrium species in Lesotho, South Africa and Swaziland is provided.     

Geographic distribution patterns of small mammals in Swaziland in relation to abiotic factors and human land-use activity

Swaziland is a small, topographically diverse southern Africa country whose mammalian fauna has been poorly studied. The distributions of small mammals in Swaziland were surveyed at 39 localities. A total of 15 species of rodents and ten species of insectivores were captured during the survey. The number of species of insectivore captured at a locality was positively correlated with mean annual rainfall, while the number of rodent species captured was negatively correlated with altitude. The number of rodent species captured was lower on Swazi Nation Land compared with privately owned land or protected land (reserves). This was probably due to the radical habitat alteration that had taken place on Swazi Nation Land, inter alia overgrazing by livestock, cultivation of maize and deforestation. The effect of this habitat alteration on the number of rodent species was more pronounced in high-lying areas of Swaziland. This may have been due to the fact that a large number of the rodent species inhabiting high-lying areas require thick, tall grassland habitats, whereas many of the low-lying species prefer more open habitats with less grass cover. Since grazing acts to reduce grass cover, it is suggested that the species inhabiting high-lying areas would be more affected by overgrazing, than low-lying species.     

Environmental factors determining the phylogenetic structure of C4 grass communities

Aim To determine how the distribution of species richness is associated with environmental factors for the four major C₄ grass lineages in South Africa, as a means to explore the mechanisms responsible. Location South Africa, Lesotho and Swaziland. Methods The geographical distributions of species richness for four major C₄ grass lineages (Aristidoideae, Chloridoideae, Andropogoneae and Paniceae) were sourced from a recently published flora that divided the study region into different vegetation types. Mean values of potential environmental correlates were calculated for each vegetation type, and the relative importances of these were determined using single‐ and multiple‐predictor generalized linear models, with and without control for spatial autocorrelation. Model selection of the multiple‐predictor generalized linear models was conducted using an Akaike’s information criterion–information theoretic approach. Association with wet, intermediate or dry, shady or open, and disturbed or undisturbed habitats was also determined for each C₄ grass clade using habitat data for all the grass species, and analysed using chi‐square tests of independence. Results Andropogoneae and Paniceae are most species‐rich in areas of high precipitation and in mesic habitats. Andropogoneae are associated with high fire frequencies. Species richness in Andropogoneae decreases and in Paniceae increases in relation to livestock density. Chloridoideae species richness is relatively constant across South Africa, but is highest where there are infrequent fires, high temperatures and basic soils, and in mesic and disturbed habitats. Aristidoideae are most species‐rich in arid regions and in habitats with high temperatures, and are associated with disturbed habitats. Main conclusions Environmental variables other than precipitation, including temperature, fire frequency and grazing pressure, are strongly associated with the contrasting distributions of species richness for the various C₄ grass clades in South Africa. Our results suggest that ecological sorting is an important determinant of phylogenetic patterns in the species richness of these C₄ grass lineages.     

Commercially Important Trees as Invasive Aliens – Towards Spatially Explicit Risk Assessment at a National Scale

Alien species that are desirable and commercially important in parts of the landscape, but damaging invaders in other parts, present a special challenge for managers, planners, and policy-makers. Objective methods are needed for identifying areas where control measures should be focussed. We analysed the distribution of forestry plantations and invasive (self-sown) stands of Acacia mearnsii and Pinus spp. in South Africa; these two taxa account for 60% of the area under commercial plantations and 54% of the area invaded by alien trees and shrubs. The distribution of commercial forestry plantations and invasive stands of these taxa were mapped and the data was digitised and stored as Geographic Information System (GIS) (Arc/Info) layers. A series of environmental parameters were derived from GIS layers of climate, topography, geology, land use, and natural vegetation. The current distribution of the two taxa was subdivided into three groups according to the degree of invasion, planting history and the precision of the data collection. We used regression-tree analysis to relate, for each taxon, the distribution of invasive stands with environmental variables, and to derive habitat suitability maps for future invasion. The current distribution of invasive stands in South Africa was largely influenced by climatic factors. At a national scale, the distribution of large commercial plantations was a poor predictor of areas invaded by both taxa. Using environmental factors identified by the regression trees, we found that 6.6% and 9.8% of natural habitats currently not invaded and untransformed by urbanisation or agriculture are suitable for invasion by Pinus spp. and A. mearnsii, respectively. We then derived guidelines for policy on alien plant management based on vegetation type, degree of transformation, extent of invasion, and the risk of future alien spread. These factors were used to identify demarcated areas where these alien species can be grown with little risk of invasions, and areas where special measures are needed to manage spread from plantations.     

A conservation assessment of the freshwater crabs of southern Africa (Brachyura: Potamonautidae)

Recent taxonomic revisions of the freshwater crabs of southern Africa (Angola, Botswana, Lesotho, Mozambique, Namibia, South Africa, Swaziland, Zambia and Zimbabwe) allow accurate depictions of their diversity, distribution patterns and conservation status. The southern African region is home to nineteen species of freshwater crabs all belonging to the genus Potamonautes (family Potamonautidae). These crabs show high levels of species endemism (84%) to the southern African region and to the country of South Africa (74%). The conservation status of each species is assessed using the IUCN (2003) Red List criteria, based on detailed compilations of the majority of known specimens. The results indicate that one species should be considered vulnerable, fifteen species least concern and three species data deficient. The results have been utilized by the IUCN for Red Lists, and may prove useful when developing a conservation strategy for southern Africa’s endemic freshwater crab fauna.     

Echinoderms (Echinodermata) from the Gulf of California, Mexico

A systematic list of the echinoderms of the Gulf of California, based on museum specimens of the Colección Nacional de Equinodermos, Instituto de Ciencias del Mar y Limnologia, Universidad Nacional Autónoma de México and the National Museum of Natural History, Smithsonian Institution, Washington, D.C. is presented. A total of 193 echinoderm species is recorded, distributed in 108 genera, 51 families and 19 orders. 12 new records for the Gulf of California are presented: Asteroidea (four), Ophiuroidea (three) and Holothuroidea (five).     

Predicting the distribution of <Emphasis Type="Italic">Encephalartos latifrons</Emphasis>, a critically endangered cycad in South Africa

This study evaluates how a modelling approach to determine areas of suitable habitat for the Critically Endangered Albany cycad Encephalartos latifrons can assist in systematic conservation planning for this and other rare and threatened cycads. A map distinguishing suitable from unsuitable habitat for E. latifrons was produced and important environmental predictors (climate, geology, topography and vegetation) influencing the suitable habitat were estimated. The maximum entropy (MaxEnt) modelling technique was chosen for this study as it has consistently performed well compared with alternative modelling methods and is also an appropriate model choice when the sample size is small and locality records are relatively few. Predicted habitat suitability showed that some locations chosen for translocation and restoration of E. latifrons specimens are not suitable. This revealed that modelling suitable habitat can guide relocation and regeneration of E. latifrons and perhaps other threatened cycads with restricted distributions and few locality records. The species distribution model constructed for E. latifrons is the first reported habitat model for a Critically Endangered cycad in South Africa. The results may be incorporated into conservation planning and structured decision-making about translocations and restoration programmes involving vulnerable cycads, which are among the most threatened organisms globally.     

Neogene paleobiogeography and East African paleoenvironments: contributions from the Tugen Hills rodents and lagomorphs.

A minimum of 28 genera of rodents and one genus of lagomorph were recovered from the Tugen Hills, Baringo District, Kenya, from localities dating from over 15.5 to about 4.4 Ma. The middle Miocene (sites dated between 15.8 and 15.3 Ma) rodent fauna recovered primarily from the Kipsaramon site complex, Muruyur Formation, includes a mixture of characteristically early Miocene taxa, and more derived forms. Composition of the African rodent fauna changes dramatically with the introduction of myocricetodontines, democricetodontines, and dendromurines, immigrants primarily from southern Asia. In the Tugen Hills, these taxa are first found in the Kabasero localities, Ngorora Formation, at sites dating from 12.5-12.33 Ma. A second major change in the African rodent fauna reflects the introduction of murines, immigrants from southern Asia. In the Tugen Hills murines are first encountered at Kapcheberek, Lukeino Formation, dated to 5.9-5.7 Ma. One rodent genus from the Lukeino Formation (Arvicanthis), and two from the Tabarin locality, Chemeron Formation (Heliosciurus, Paraxerus; 4.5-4.4 Ma), represent the earliest records of these extant African genera. A cricetomyine from the Ngorora Formation (12.5 Ma) is likely the earliest report of this exclusively African group. One of the earliest African records of porcupines (Hystricide) is from the Lukeino Formation. Lagomorphs are poorly represented, but include one of the earliest African occurrences of the family Leporidae from the Mpesida Beds (bracketed by dates of 7-6.2 Ma), and possibly a new genus of leporid from the Kapcheberek locality. Analysis of the Tugen Hills small mammals in association with other African records suggests several episodes of dispersal between Africa and Eurasia during the middle and late Miocene. Rodents from Kipsaramon are indicative of forests in conjunction with more open habitats. Those from the Kapcheberek locality are suggestive of a savanna habitat. The rodents from the Tabarin locality suggest a woodland environment.     

The large Branchiopoda (Crustacea) from temporary habitats of the Drakensberg region,South Africa

The Drakensberg forms part of the Escarpment which separates the coastal plain from the inland plateau in southern Africa. This mountain range runs for about 400 km along the KwaZulu-Natal/Lesotho border and into the Eastern Cape province of South Africa. Altitude ranges from 1500 to 3000 masl. Rock pools and tarns are the main type of temporary pool habitat in the Drakensberg. 90 different habitats were sampled over a four-year period and various physical and chemical characteristics of these pools are presented. Large branchiopods occurred in 26 of these habitats. Five Branchipodopsis and two Streptocephalus species composed the anostracan fauna, while Triops granarius was collected only from the summit of the Escarpment at one locality. Four genera of Spinicaudata are represented in the Drakensberg but the current state of the taxonomy of this group does not allow identification to species level. Absence of large branchiopods from a high percentage of temporary habitats could be attributed to predation or many pools may not be sufficiently ephemeral. Most pools were inhabited by a single species of large branchiopod. This could be a result of low levels of food resources in pools, as indicated by extremely low conductivities. Restricted food resources could also be the limiting factor in the distribution of Streptocephalus and Triops which are only found in pools with higher conductivities. Four of the five Branchipodopsis species are endemic to the Drakensberg pools. These habitats are also inhabited by a high diversity of ostracods and other micro-crustaceans. Most of the KwaZulu-Natal Drakensberg falls within conservation areas but the Lesotho and Free State localities could be threatened by future development.     

THE DISTRIBUTION OF THE PIED STARLING,AND SOUTHERN AFRICAN BIOGEOGRAPHY

Craig, A. J. F. K. 1985. The distribution of the Pied Starling, and southern African biogeography. Ostrich 56:123-131.

The Pied Starling is restricted to South Africa, Lesotho and Swaziland, but within this region it occupies a variety of habitats. Its modern distribution is certainly influenced by its association with man. However, a comparison with other vertebrate species, and with patterns of plant distribution, suggests that its geographical distribution has been determined primarily by its evolutionary history. A hypothesis for the origin of this species leads to several predictions concerning biogeography in southern Africa.     

Distribution of lichen flora on South Korea

After an overview on the temporary situation of the lichenology in South Korea, localities of 95 macrolichen taxa are reported for South Korea. In this revised lichen flora of South Korea, 16 species are apparently new to the territory. Voucher specimens have been deposited in the Korean Lichen Research Institute (KoLRI) at Sunchon National University in Korea, and duplicates have also been donated to the National History Museum and Institute, in Chiba, (CBM) Japan.     

The taxonomy, distribution and origins of two species of Phaulacridium (Orthoptera: Acrididae) in the South Island of New Zealand

Extensive collecting of grasshoppers of the genus Phaulacridium in New Zealand has indicated the presence of a new species, P. otagoense Ritchie & Westerman, distributed parapatrically with respect to the common and widespread species, P. marginale (Walker) in the Mackenzie Plains and Central Otago regions of South Island. Phaulacridium otagoense occurs on exposed hillsides with bare patches caused by rabbit grazing while P. marginale is found in adjoining areas with lusher vegetation and less severe erosion. A key to the two species and a differential diagnosis are provided. Statistically significant differences in morphology and in nuclear DNA content and chiasma frequency confirm the separate species status of the two taxa. The present-day distribution and probable allopatric origin of P. otagoense from P. marginale is discussed in the light of past and present vegetational and climatic factors and their recent disruption by human agencies. The holotype and some paratypes of P. otagoense are deposited in the Canterbury Museum, Christchurch, New Zealand and further paratypes are deposited in the Australian National Insect Collection, Canberra and the British Museum (Natural History), London     

藜科植物的起源、分化和地理分布

全球藜科植物共约130属1500余种,广泛分布于欧亚大陆、南北美洲、非洲和大洋洲的半干旱及盐碱地区。它基本上是一个温带科,对亚热带和寒温带也有一定的适应性。本文分析了该科包含的1l族的系统位置和分布式样,以及各个属的分布区,提出中亚区是现存藜科植物的分布中心,原始的藜科植物在古地中海的东岸即华夏陆台(或中国的西南部)发生,然后向干旱的古地中海沿岸迁移、分化,产生了环胚亚科主要族的原始类群;起源的时间可能在白垩纪初,冈瓦纳古陆和劳亚古陆进一步解体的时期。文章对其迁移途径及现代分布式样形成的原因进行了讨论。     

Origin,Differentiation, and Geographic Distribution of the Chenopodiaceae

Numbers of species and genera,endemic genera,extant primitive genera,relationship and distribution patterns of presently living Chenopodiaceae(two subfamilies,12 tribes,and 118 genera)are analyzed and compared for eight distributional areas,namely central Asia,Europe,the Mediterranean region,Africa,North America,South America, Australia and East Asia． The Central Asia,where the number of genera and diversity of taxa are greater than in other areas,appears to be the center of distribution of extant Chenopodiaceae．North America and Australia are two secondary centers of distribution． Eurasia has 11 tribes out of the 12,a total of 70 genera of extant chenopodiaceous plants,and it contains the most primitive genera of every tribe． Archiatriplex of Atripliceae,Hablitzia of Hablitzeae,Corispermum of Corispermeae,Camphorosma of Camphorosmaea,Kalidium of Salicornieae,Polecnemum of Polycnemeae,Alexandra of Suaedeae,and Nanophyton of Salsoleae,are all found in Eurasia,The Beteae is an Eurasian endemic tribe,demonstrating the antiquity of the Chenopodiaceae flora of Eurasia．Hence,Eurasia is likely the place of origin of chenopodiaceous plants． The presence of chenopodiaceous plants is correlated with an arid climate．During the Cretaceous Period,most places of the continent of Eurasia were occupied by the ancient precursor to the Mediterranean,the Tethys Sea．At that time the area of the Tethys Sea had a dry and warm climate．Therefore,primitive Chenopodiaceae were likely present on the beaches of this ancient land．This arid climatic condition resulted in differentiation of the tribes Chenopodieae,Atripliceae,Comphorosmeae,Salicornieae,etc．,the main primitive tribes of the subfamily Cyclolobeae. Then following continental drift and the Laurasian and Gondwanan disintegration, the Chenopodiaceae were brought to every continent to propagate and develop, and experience the vicissitudes of climates, forming the main characteristics and distribution patterns of recent continental floras. The tribes Atripliceae, Chenopodieae, Camphorosmeae, and Salicornieae of recent Chenopodiaceae in Eurasia, North America, South America, southern Africa, and Australia all became strongly differentiated. However, Australia and South America, have no genera of Spirolobeae except for a few maritime Suaeda species. The Salsoleae and Suaedeae have not arrived in Australia and South America, which indicates that the subfamily Spirolobeae developed in Eurasia after Australia separated from the ancient South America-Africa continent, and South America had left Africa. The endemic tribe of North America, the tribe Sarcobateae, has a origin different from the tribes Salsoleae and Suaedeae of the subfamily Spirolobeae. Sarcobateae flowers diverged into unisexuality and absence of bractlets. Clearly they originated in North America after North America had left the Eurasian continent. North America and southern Africa have a few species of Salsola, but none of them have become very much differentiated or developed, so they must have arrived through overland migration across ancient continental connections. India has no southern African Chenopodiaceae floristic components except for a few maritime taxa, which shows that when the Indian subcontinent left Africa in the Triassic period, the Chenopodiaceae had not yet developed in Africa. Therefore, the early Cretaceous Period about 120 million years ago, when the ancient Gondwanan and Laurasian continents disintegrated, could have been the time of origin of Chenopodiaceae plants.The Chinese flora of Chenopodiaceae is a part of Chenopodiaceae flora of central Asia. Cornulaca alaschnica was discovered from Gansu, China, showing that the Chinese Chenopodiaceae flora certainly has contact with the Mediterranean Chenopodiaceae flora. The contact of southeastern China with the Australia Chenopodiaceae flora, however, is very weak.     

内蒙古贺兰山啮齿动物群落多样性及其与环境因子关系

啮齿动物群落结构可以反映生态环境特征。本研究对处于阿拉善荒漠区呈“孤岛”状态的内蒙古贺兰山国家级自然保护区内啮齿动物的种类、分布型及群落多样性进行系统研究。将内蒙古贺兰山划分为五种生境类型,于2013年春、夏、秋3季共布设有效铗日18748个,捕获啮齿动物235只,分属2目5科11属13种。在整体研究区域,阿拉善黄鼠（Spermophilus alashanicus）为优势种。明确了喜湿型是本研究区域的主要分布型,占捕获啮齿动物的54%,并且在中低海拔区域,随着海拔的升高,喜湿型所占比例增加。受“边缘效应”的影响,山地荒漠和荒漠草原生境以及山地草原灌丛生境的啮齿动物群落多样性指数高于其他生境,除环境因素外,啮齿动物群落的多样性受两种因素的影响,既可随群落内物种数量的增加而增大,同时又受制于群落内部物种分布的均匀程度。冗余分析结果表明,植被高度、植被盖度、灌木（乔木）高度和海拔4个环境因子是决定啮齿动物群落结构最主要的环境因子,其中植被盖度与群落多样性呈负相关,随着植被盖度的增加,多样性指数随之减低。     

Evidence of environmental niche differentiation in the striped mouse (Rhabdomys sp.): inference from its current distribution in southern Africa

The aim of this study was to characterize environmental differentiation of lineages within Rhabdomys and provide hypotheses regarding potential areas of contact between them in the Southern African subregion, including the Republic of South Africa, Lesotho, and Namibia. Records of Rhabdomys taxa across the study region were compiled and georeferenced from the literature, museum records, and field expeditions. Presence records were summarized within a 10 × 10 km grid covering the study area. Environmental information regarding climate, topography, land use, and vegetation productivity was gathered at the same resolution. Multivariate statistics were used to characterize the current environmental niche and distribution of the whole genus as well as of three mitochondrial lineages known to occur in southern Africa. Distribution modeling was carried out using MAXENT in order to generate hypotheses regarding current distribution of each taxa and their potential contact zones. Results indicate that the two species within Rhabdomys appear to have differentiated across the precipitation/temperature gradient present in the region from east to west. R. dilectus occupies the wettest areas in eastern southern Africa, while R. pumilio occupies the warmer and drier regions in the west, but also penetrates in the more mesic central part of the region. We provide further evidence of environmental differentiation within two lineages of R. dilectus. Contact zones between lineages appear to occur in areas of strong environmental gradients and topographic complexity, such as the transition zones between major biomes and the escarpment area where a sharp altitudinal gradient separates coastal and plateau areas, but also within more homogeneous areas such as within grassland and savannah biomes. Our results indicate that Rhabdomys may be more specialized than previously thought when considering current knowledge regarding mitochondrial lineages. The genus appears to have differentiated along two major environmental axes in the study region, but results also suggest dispersal limitations and biological interactions having a role in limiting current distribution boundaries. Furthermore, the projection of the potential geographic distribution of the different lineages suggests several contact zones that may be interesting study fields for understanding the interplay between ecological and evolutionary processes during speciation.     

A PCA-based modelling technique for predicting environmental suitability for organisms from presence records

We present a correlative modelling technique that uses locality records (associated with species presence) and a set of predictor variables to produce a statistically justifiable probability response surface for a target species. The probability response surface indicates the suitability of each grid cell in a map for the target species in terms of the suite of predictor variables. The technique constructs a hyperspace for the target species using principal component axes derived from a principal components analysis performed on a training dataset. The training dataset comprises the values of the predictor variables associated with the localities where the species has been recorded as present. The origin of this hyperspace is taken to characterize the centre of the niche of the organism. All the localities (grid-cells) in the map region are then fitted into this hyperspace using the values of the predictor variables at these localities (the prediction dataset). The Euclidean distance from any locality to the origin of the hyperspace gives a measure of the 'centrality' of that locality in the hyperspace. These distances are used to derive probability values for each grid cell in the map region. The modelling technique was applied to bioclimatic data to predict bioclimatic suitability for three alien invasive plant species ( Lantana camara L., Ricinus communis L. and Solanum mauritianum Scop.) in South Africa, Lesotho and Swaziland. The models were tested against independent test records by calculating area under the curve (AUC) values of receiver operator characteristic (ROC) curves and kappa statistics. There was good agreement between the models and the independent test records. The pre-processing of climatic variable data to reduce the deleterious effects of multicollinearity, and the use of stopping rules to prevent overfitting of the models are important aspects of the modelling process.