The taxonomic status and relationships of bracken ferns (Pteridium: Dennstaedtiaceae) from sub-Saharan Africa

Two morphotypes of bracken fern in the genus Pteridium Gled. ex Scop. occur in sub-Saharan Africa. The more common and widespread of these is assigned in contemporary systematic treatments to the European subspecies, P. aquilinum (L.) Kuhn ssp. aquilinum . We show that this African form is separable morphometrically and genomically from the European subspecies, and reinstate its earlier name P. aquilinum ssp. capense (Thunb.) C.Chr. The second African bracken, with a more localized tropical distribution mainly in the drainage basins of the Congo and Zambezi River systems, is confirmed as P. aquilinum ssp. centrali-africanum Hieron. ex R.E. Fr. We reject suggestions that this taxon be treated as a full species. Phenetic cluster analysis based on use of Arbitrarily Primed Polymerase Chain Reaction (A-P PCR) and Inter Simple Sequence Repeat (ISSR) fingerprinting of the nuclear genome provides evidence that the two African brackens are more closely related to each other than to other taxa, and sister to a grouping of the European sspp. aquilinum and pinetorum . The two African subspecies share solely with ssp. aquilinum a distinctive chloroplast haplotype carrying a 5-base direct repeat in the rps 4 –trn S region, confirming the close phyletic relationship between sspp. aquilinum , capense and centrali-africanum .  © 2005 The Linnean Society of London, Botanical Journal of the Linnean Society , 2005, 148 , 311–321.     

Ecology and systematics of mangrove crabs of the genus Perisesarma (Crustacea: Brachyura: Sesarmidae) from East Africa

East Africa has a reduced mangrove crab species richness when compared to Asian mangroves. To date, only one species of Perisesarma de Man, 1895 has been reported in East Africa, despite more than 30 years of mangrove research in this region. Based on morphology, colour, mtDNA and behaviour, we describe a new species of Perisesarma from Kenya, P. samawati sp. nov. Surprisingly, when comparing molecular data from other species within this genus, P. samawati sp. nov. and the sympatric P. guttatum (A. Milne Edwards, 1869) are not sister species. Some aspects of the ecology of P. guttatum and P. samawati sp. nov. are compared and the differences discussed. Additionally, we compare P. samawati sp. nov. with the ecological literature of a possible sister species P. eumolpe de Man, 1895 from Malaysian mangroves. Our findings suggest that the new species is an ecologically important species in East African mangroves.  © 2004 The Linnean Society of London, Zoological Journal of the Linnean Society , 2004, 141 , 435–445.     

High species diversity in one of the dominant groups of spiders in East African montane forests (Araneae: Pholcidae: Buitinga n. gen., Spermophora Hentz)

The six-eyed pholcid spiders of East Africa are revised. A new genus is recognized, Buitinga n. gen ., with 17 new species and three species transferred from Spermophora Hentz. The new genus is characterized by the presence of a scape on the epigynum. This scape may be straight or tightly curled up at rest, and is usually highly expandable. Seven additional African and Comoran species are newly described and tentatively assigned to Spermophora . A data matrix with 60 characters and 77 taxa (including 20 East African species and 25 additional Spermophora and ' Spermophora -like' species) is analysed cladistically. Buitinga is closer to the genera Paramicromerys Millot (endemic in Madagascar) and Spermophorides Wunderlich (Mediterranean and Canary Islands) as well as to several African and Comoran species tentatively assigned to Spermophora than to the type species of Spermophora . It is argued that current estimates of species numbers may be inaccurate and that pholcids may turn out to be one of the most diverse spider families.  © 2003 The Linnean Society of London. Zoological Journal of the Linnean Society, 2003, 137, 555−619     

Studies on Thapsia (Apiaceae) from north-western Africa: a forgotten and a new species

Following a revision of Thapsia (Apiaceae) in north-western Africa, the name Thapsia platycarpa is resurrected and lectotypified for a species that grows between Algeria and Morocco, and a new species Thapsia cinerea is described from the Rif region of north-eastern Morocco. Morphological features that differentiate between these and other species ( T. villosa , T. garganica , T. transtagana and T. gymnesica ) are discussed. An identification key for the plants of the area is presented.  © 2003 The Linnean Society of London, Botanical Journal of the Linnean Society , 2003, 143 , 433–442.     

A synopsis of Goniothalamus species (Annonaceae) in Peninsular Malaysia, with a description of a new species

The species of Goniothalamus (Annonaceae) occurring in Peninsular Malaysia and Singapore are revised, updating the previous taxonomic treatment by James Sinclair, published in 1955. A total of 18 species is recognized, including a new species, G. tomentosus . Collections referrable to G. tomentosus were previously determined as ' G. marcanii '; examination of the types of G. marcanii reveals that the two taxa are not conspecific, however, and a new name is accordingly validated here. Goniothalamus tomentosus has closest affinities with the Javan/Sumatran species G. costulatus . Other important nomenclatural changes include the reduction of G. umbrosus to synonymy with G. tapis .  © 2003 The Linnean Society of London, Botanical Journal of the Linnean Society, 2003, 142 , 321−339.     

Phylogeny and biogeography of Crinum L. (Amaryllidaceae) inferred from nuclear and limited plastid non-coding DNA sequences

The genus Crinum L. is the only pantropical genus of the Amaryllidaceae. Phylogenetic and biogeographical analyses of nrDNA ITS and plastid trnL-F sequences for all continental groups of the genus Crinum and related African genera are presented, with the genus Amaryllis used as outgroup. ITS indicates that C. baumii is more closely related to Ammocharis and Cybistetes than to Crinum sensu stricto . Three clades are resolved in Crinum s.s. One unites a monophyletic American group with tropical and North African species. The second includes all southern African species and the Australian endemic C. flaccidum . The third includes monophyletic Madagascar, Australasian and Sino-Himalayan clades, with southern African species. The trnL-F phylogeny resolves an American and an Asian/Madagscar clade, and confirms the relationship of C. flaccidum with species endemic to southern Africa. The salverform, actinomorphic perianths of subg. Crinum appear to have evolved several times in the genus from ancestors with zygomorphic perianths (subg. Codonocrinum ), thus neither subgenus is monophyletic. Biogeographical analyses place the origin of Crinum in southern Africa, as the region is optimized at all ancestral nodes in the tree topology, and in basal interior nodes of all but one of the major clades. The genus underwent three major waves of radiation corresponding to the three main clades resolved in our trees. Two entries into Australia for the genus are indicated, as are separate Sino-Himalayan and Australasian dispersal events.  © 2003 The Linnean Society of London, Botanical Journal of the Linnean Society , 2003, 141 , 349–363.     

Isoetes fluitans sp. nov.: the identity of Spanish plants of 'I. longissimum'

Recent revision of North African specimens of Isoetes velata A. Braun and the closely related taxon I. longissimum Bory, together with Spanish material conventionally designated I. longissimum , suggests that the Spanish specimens constitute a new species, I. fluitans . This is described and illustrated. The North African taxon I. longissimum is probably not specifically distinct from I. velata .  © 2004 The Linnean Society of London, Botanical Journal of the Linnean Society , 2004, 146 , 231–236.     

A synopsis of Goniothalamus species (Annonaceae) in Thailand, with descriptions of three new species

A comprehensive taxonomic revision of Goniothalamus species (Annonaceae) occurring in Thailand is presented for the first time. Twenty-five species are recognized, including three that are described as new to science ( Goniothalamus aurantiacus from South-Western Thailand, Goniothalamus maewongensis from Northern Thailand, and Goniothalamus rongklanus from Northern and North-Eastern Thailand). Several taxonomic and nomenclatural misunderstandings are corrected. The name G. griffithii is shown to be widely misapplied for populations in Northern Thailand, for which the name G. calvicarpus should be applied; 'true' G. griffithii is restricted to South-Western Thailand and Myanmar. In addition, the widely used name G. marcanii is shown to be a synonym of G. tamirensis , and the name G. latestigma , previously regarded as a synonym of G. undulatus , is reinstated. Goniothalamus cheliensis is furthermore newly recorded from Thailand. Most species are restricted to Peninsular Thailand and represent a Malesian floristic component.  © 2008 The Linnean Society of London, Botanical Journal of the Linnean Society , 2008, 156 , 355–384.     

Patterns of diversification on old volcanic islands as revealed by the woodlouse-hunter spider genus Dysdera (Araneae, Dysderidae) in the eastern Canary Islands

Long-term erosion and subsidence cause dramatic alterations in the physical and ecological features of oceanic islands. Although oceanic islands have been extensively used as models for the study of speciation, little attention has been given to investigating evolutionary patterns in old volcanic islands that have suffered severe climatic degradation. The spider genus Dysdera has diversified across the Canary Islands and has evolved endemisms in the low-elevation, xeric eastern islands, which sharply contrast with the younger, higher, and more humid western islands. A combined phylogenetic analysis of seven mitochondrial and nuclear genes reveals that the eastern Canaries were colonized twice, although only one lineage underwent in situ diversification. Origins of the speciose lineage remain obscure, but probably preceded diversification of present-day Iberian and North African species. A second colonization of the eastern Canaries from North Africa has occurred in more recent times. Molecular analyses reveal several instances of geographically coherent cryptic lineages further supported by morphometric evidence. Analyses of diversification rates suggest deceleration of diversification over the course of time, and this is compatible with increasing extinction rates due to drastic yet continuous ecological changes. Extinction may also explain incongruent patterns of morphological differentiation and species coexistence. Despite a general trend towards community impoverishment, there is also evidence for recent speciation events linked to ecological shifts, which may illustrate the origins of nonspeciose relic lineages on islands.  © 2008 The Linnean Society of London, Biological Journal of the Linnean Society , 2008, 94 , 589–615.     

Monophyly and phylogenetic placement of the spider genus Labulla Simon, 1884 (Araneae, Linyphiidae) and description of the new genus Pecado

The genus Labulla Simon is circumscribed in phylogenetic terms to include the species Labulla thoracica (Wider), L. flahaulti Simon and L. machadoi sp. nov. The genital anatomy of the genus is described in detail and the taxonomy of the genus is reviewed. The monophyly of Labulla is supported by numerous morphological apomorphies of the male palp and female epigynum. Based on a cladistic analysis, a new genus, Pecado gen. nov. , is erected to place Labulla impudica Denis, from Northern Africa. Lepthyphantes insularis Saito and ' Labulla ' nepula Tikader, both formerly included in Labulla , are not congeneric with the type species of Labulla .  © 2005 The Linnean Society of London, Zoological Journal of the Linnean Society , 2005, 143 , 359–404.     

Southern African pholcid spiders: revision and cladistic analysis of Quamtana gen. nov. and Spermophora Hentz (Araneae: Pholcidae), with notes on male−female covariation

The southern African pholcid spiders previously assigned to Pholcus and Spermophora are revised and their phylogenetic position within Pholcidae is analysed cladistically. Two distinct groups are recognized: the first is restricted to the Cape region and probably correctly placed in Spermophora ; it includes Spermophora peninsulae Lawrence and four new species. The second is more widely distributed in southern and eastern South Africa, and reaches as far north as Cameroon, Congo (Democratic Republic) and Uganda; it is assigned generic status ( Quamtana gen. nov .), and includes two species transferred from Pholcus [ Q. leptopholcica (Strand), Q. ciliata (Lawrence)] as well as 23 new species. A key to the pholcid spiders of South Africa is presented. A close correlation between a male character (distance between cheliceral apophyses) and a female character (distance between pockets on epigynum) is documented in Quamtana .  © 2003 The Linnean Society of London, Zoological Journal of the Linnean Society, 2003, 139 , 477−527.     

Can we predict centres of plant species richness and rarity from environmental variables in sub‐Saharan Africa?

In order to investigate continental-scale patterns of plant species richness and rarity, distribution maps of 3661 plant species were digitized into a one degree grid of sub-Saharan Africa using the WORLDMAP computer programme. Cells with high species richness were also likely to be those containing the greatest number of species of restricted range, but areas such as the South African Cape and the Eastern Arc mountains were found to have more restricted-range species than predicted from their richness scores. The two environmental predictors which had the strongest individual relationships with both species richness and range-size rarity were absolute maximum annual temperature and mean monthly potential evapotranspiration. However, correlative predictive powers of these variables were low, with R =−0.58 and R =−0.54, respectively ( P  < 0.01). Multiple regression also failed to produce a strong explanatory model for observed continental-scale patterns of diversity. Spatial variability analysis showed that this was likely to be because different environmental parameters predicted different centres of richness and rarity. West African species richness was better predicted by absolute maximum annual temperature, whereas East African species richness was better predicted by mean monthly potential evapotranspiration.  © 2003 The Linnean Society of London, Botanical Journal of the Linnean Society , 2003, 142 , 187–197.     

The fossil history of palms (Arecaceae) in Africa and new records from the Late Oligocene (28–27 Mya) of north-western Ethiopia

The African palm fossil record is limited but the data provide an outline of palm evolution from the Late Cretaceous through the Neogene. Pollen attributed to palms is reported from the Aptian (125–112 Mya), but the earliest unequivocal record in Africa is Campanian (83.5–70.6 Mya). Palms diversified 83.5–65.5 Mya and became widespread, although most records are from the west and north African coasts. Many taxa were shared between Africa and northern South America at that time, but a few were pantropical. Extirpations occurred throughout the Palaeogene, including a notable species turnover and decline at the Eocene–Oligocene boundary (33.9 Mya), a change that resulted in the elimination of nypoid palms from Africa. The Neogene plant macrofossil record is better sampled than the Palaeogene, although few palms are documented. Thus, the low diversity of African palms today is more likely the result of Palaeogene, rather than Neogene extinctions. Newly discovered palm fossils of leaves, petioles and flowers from the Late Oligocene (27–28 Mya) of north-western Ethiopia document the abundance and dominance of palms in some communities at that time. The fossils represent the earliest records of the extant genera Hyphaene (Coryphoideae) and Eremospatha (Calamoideae).  © 2006 The Linnean Society of London, Botanical Journal of the Linnean Society , 2006, 151 , 69–81.     

Revision of the Micromeria (Labiatae) in tropical to southern Africa and on the Arabian Peninsula

The material of Micromeria s.s. ( M. sect. Micromeria ) from tropical to southern Africa and the Arabian Peninsula is revised. The endemic Ethiopian M. unguentaria Schweinf. is retained as a species, while the other African and Arabian material of Micromeria is included in M. imbricata (Forssk.) C. Chr. M. imbricata is divided into the following three varieties: var. imbricata , which is very polymorphic, widespread in the African highlands, and also occurs in Arabia; var. rhodesiaca (Elly Walther & K.H. Walther) Ryding, which occurs in the highlands of Malawi, Mozambique and Zimbabwe; and var. villosa (Elly Walther & K.H. Walther) Ryding, which occurs in montane to alpine areas of Ethiopia and East Africa. Discontinuities in the variation within var. imbricata , observed at some localities, are suggested to be a result of partial autogamy. Problems with delimitation between M. imbricata , the East Asiatic M. biflora (Buch.-Ham. ex D.Don) Benth. and some Mediterranean species are discussed.  © 2007 The Linnean Society of London, Botanical Journal of the Linnean Society , 2007, 155 , 427–446.     

Chromosome studies in Hippeastrum (Amaryllidaceae): variation in genome size

This paper presents the karyotype and DNA content of 12 diploid species of Hippeastrum from South America. The variation in genome size is compared with the karyotype and DNA content of Amaryllis belladonna from South Africa. The Hippeastrum species present a uniform and bimodal basic karyotype formula, but significant differences are found in the total chromosome volume (TCV) and nuclear DNA content. A positive correlation between the DNA content and TCV is also observed. The karyotype's constancy is a product of changes in DNA content occurring in the whole chromosome complement. The DNA addition to the long and short sets of chromosomes varies independently. In species with higher DNA contents, the short chromosomes add equal DNA amounts to both arms, maintaining their metacentric morphology, whereas the long chromosomes add DNA only to the short arm, increasing the chromosome symmetry. These data show that the evolutionary changes in DNA amount are proportional to chromosome length, maintaining the karyotypic uniformity. A. belladonna has a larger DNA content and possesses a karyotype different from that of Hippeastrum spp., supporting the distinction between the two genera and upholding the name Amaryllis for the South African entity against Hippeastrum for the South American genus.  © 2007 The Linnean Society of London, Botanical Journal of the Linnean Society , 2007, 155 , 171–178.     

Systematics and phylogeny of the cane rats (Rodentia: Thryonomyidae)

The family Thryonomyidae is known from the Eocene up to the present. Today, this group comprises just two closely related species, which are restricted to sub-Saharan Africa. However, various thryonomyids have been recorded in strata of Miocene age, when the group spread out of Africa eastward to southern Asia (Pakistan). A systematic revision and a cladistic analysis shows that 20 species can be referred to this family: Thryonomys swinderianus (Temninck), T. gregorianus (Thomas), Paraphiomys pigotti Andrews, P. occidentalis Lavocat, P. simonsi Wood, P. hopwoodi Lavocat, P. shipmani Denys et Jaeger, P. australis Mein, Pickford et Senut, P. roessneri Mein, Pickford et Senut, P. afarensis Geraads, Paraphiomys sp. nov. from Saudi Arabia López-Antoñanzas et Sen , P. renelavocati sp. nov. , Neosciuromys africanus Stromer, Apodecter stromeri Hopwood, Paraulacodus indicus Hinton, Paraulacodus johanesi Jaeger, Michaux et Sabatier, Gaudeamus aegyptius Wood, Epiphiomys coryndoni Lavocat, Kochalia geespei (de Bruijn et Hussain), Paraphiomys sp. nov. from Saudi Arabia, and Paraphiomys orangeus Mein et Pickford. The unresolved basal position of Sacaresia moyaeponsi with respect to Metaphiomys schaubi and the clade comprising the above-cited species, suggest that this taxon should not be allocated to the family Thryonomyidae. A phylogenetic definition of the family Thryonomyidae is proposed as an outcome of the phylogenetic analysis: Epiphiomys corindoni , Thryonomys swinderianus , their most recent common ancestor and all its descendants (node-based taxon).  © 2004 The Linnean Society of London, Zoological Journal of the Linnean Society , 2004, 142 , 423–444.