A phylogeny of Delphinieae (Ranunculaceae) shows that Aconitum is nested within Delphinium and that Late Miocene transitions to long life cycles in the Himalayas and Southwest China coincide with bursts in diversification

The tribe Delphinieae (Ranunculaceae) comprises two species-rich genera, Aconitum and Delphinium, the latter including Consolida and Aconitella. The 650-700 species are distributed in Eurasia and North America; three species occur on tropical African mountains. Maximum likelihood analyses of 2088 aligned nucleotides of plastid and nuclear sequences obtained from up to 185 species of Delphinieae from throughout the geographic range (plus relevant outgroups) show that three short-lived (facultative annual or biennial) Mediterranean species belonging to Delphinium subgenus Staphisagria are the sister clade to all other Delphinieae, implying that Staphisagria needs to be raised to genus status if Delphinium and Aconitum are to become mutually monophyletic. Molecular clock dating suggests an origin of the sampled Delphinieae in the Early Oligocene (c. 32.3 Ma) and expansion to North America of Aconitum and Delphinium around 3.3 and 2.9 Ma ago, respectively; the East African Mts. were reached by long-distance dispersal some 2.4 Ma ago, coincident with the major uplift of the East African Rift system. The ancestral growth form of the Delphinieae could not be reconstructed, but Late Miocene bursts in diversification rates in the Himalayan and southwestern Chinese clades of Aconitum and Delphinium appear to be associated with transitions from short-lived to long-lived life histories.     

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.     

Time of diversification in the Cape fauna endemisms,inferred by phylogenetic studies of the genus Iselma (Coleoptera: Meloidae: Eleticinae)

The endemic diversification of the Cape zone fauna and the phylogenetic relationships among the 30 species of the blister beetle genus Iselma are investigated. We analysed morphological, molecular (mtDNA 16S) and combined datasets of characters using a number of approaches (maximum parsimony, maximum likelihood, Bayesian inference analyses). We propose hypotheses of diversification times among taxa from molecular clock analyses. Morphological and molecular analyses produced similar results. According to maximum likelihood molecular studies, radiation within the genus Iselma occurred during the Pliocene and Pleistocene, roughly contemporaneously with the shift in African climate, vegetation and faunal assemblages. Two main lineages, one Namibian and one South African, separated c. 4.9 Ma. Within the South African lineage we identify two groups of endemic species, one in Little Karoo and one that extend towards southern Namaqualand. The remaining South African groups of species are spread through Namaqualand and the southwestern Cape area. These include several endemisms, with different times of diversification during the Pleistocene, probably related in part to glacial cycles. These endemisms are distributed in small areas of the following ecosystems: coastal Strandveld, Succulent Karoo and particularly Fynbos.     

Historical biogeography of Androcymbium Willd. (Colchicaceae) in Africa: evidence from cpDNA RFLPs

The cpDNA restriction variation in 39 populations representing a geographical sampling of 18 species of Androcymbium in southwestern and northern Africa was examined to assess the historical biogeography of the genus. The cpDNA phylogeny indicates that the disjunction between South and North Africa is best explained by the dispersal of southern African ancestors into North Africa. Divergence time estimates suggest that the geographic range of the genus may have extended considerably north (perhaps to Tanzania and Kenya) prior to the global desiccation of Africa in the Miocene. Further expansion of the genus northward was probably stalled until climatic changes in the late Miocene brought about the gradual replacement of a subtropical woodland savanna with the arid landscape that gave rise to the Sahara. Aridification of the northern quarter of the continent provided the ecological conditions for fostering the expansion of Androcymbium along the Mediterranean fringe (probably east to west) and its introduction into the Canary Islands. Unlike their South African congeners, the northern species have experienced expansions, fragmentations, and local extinctions in response to the severe climatic shifts in this area during the Pliocene-Pleistocene. According to our divergence time estimates, the arid track may have already existed as a continuous area connecting southern and northern Africa in the late Miocene.     

Out of Africa? A dated molecular phylogeny of the cicada tribe Platypleurini Schmidt (Hemiptera: Cicadidae), with a focus on African genera and the genus Platypleura Amyot & Audinet‐Serville

The Platypleurini is a large group of charismatic cicadas distributed from Cape Agulhas in South Africa, through tropical Africa, Madagascar, India and eastern Asia to Japan, with generic diversity concentrated in equatorial and southern Africa. This distribution suggests the possibility of a Gondwanan origin and dispersal to eastern Asia from Africa or India. We used a four‐gene (three mitochondrial) molecular dataset, fossil calibrations and molecular clock information to explore the phylogenetic relationships of the platypleurine cicadas and the timing and geography of their diversification. The earliest splits in the tribe were found to separate forest genera in Madagascar and equatorial Africa from the main radiation, and all of the Asian/Indian species sampled formed a younger clade nested well within the African taxa. The tribe appears to have diversified during the Cenozoic, beginning c. 50–32 Ma, with most extant African lineages originating in the Miocene or later, well after the breakup of the Gondwanan landmass. Biogeographical analysis suggests an African origin for the tribe and a single dispersal event founding the Asian platypleurines, although additional taxon sampling and genetic data will be needed to confirm this pattern because key nodes in the tree are still weakly supported. Two Platypleurini genera from Madagascar (Pycna Amyot & Audinet‐Serville, Yanga Distant) are found to have originated by late Miocene dispersal of a single lineage from Africa. The genus Platypleura is recovered as polyphyletic, with Platypleura signifera Walker from South Africa and many Asian/Indian species apparently requiring assignment to different genera, and a new Platypleura concept is proposed with the synonymization of Azanicada Villet syn.n. The genera Orapa Distant and Hamza Distant, currently listed within separate tribes but suspected of platypleurine affinity, are nested deeply within the Platypleurini radiation. The tribe Orapini syn.n . is here synonymized while the tribe Hamzini is pending a decision of the ICZN to preserve nomenclatorial stability.     

Molecular phylogenetics of the clover genus (Trifolium--Leguminosae)

Trifolium, the clover genus, is one of the largest genera of the legume family. We conducted parsimony and Bayesian phylogenetic analyses based on nuclear ribosomal DNA internal transcribed spacer and chloroplast trnL intron sequences obtained from 218 of the ca. 255 species of Trifolium, representatives from 11 genera of the vicioid clade, and an outgroup Lotus. We confirm the monophyly of Trifolium, and propose a new infrageneric classification of the genus based on the phylogenetic results. Incongruence between the nrDNA and cpDNA results suggests five to six cases of apparent hybrid speciation, and identifies the putative progenitors of the allopolyploids T. dubium, a widespread weed, and T. repens, the most commonly cultivated clover species. Character state reconstructions confirm 2n=16 as the ancestral chromosome number in Trifolium, and infer a minimum of 19 instances of aneuploidy and 22 of polyploidy in the genus. The ancestral life history is hypothesized to be annual in subgenus Chronosemium and equivocal in subgenus Trifolium. Transitions between the annual and perennial habit are common. Our results are consistent with a Mediterranean origin of the genus, probably in the Early Miocene. A single origin of all North and South American species is hypothesized, while the species of sub-Saharan Africa may originate from three separate dispersal events.     

Radiation in the Cape flora and the phylogeny of peacock irises Moraea (Iridaceae) based on four plastid DNA regions

Phylogenetic analyses of four plastid DNA regions, the rbcL exon, trnL intron, trnL-trnF intergenic spacer, and rps16 intron from each of 73 species in the African genus Moraea (Iridaceae: Irideae) including accessions of all major species clusters in the genus, show Moraea to be paraphyletic when Barnardiella, Galaxia, Hexaglottis, Homeria (all southern African), and Gynandriris (Eurasian as well) were recognized as separate genera. There are several small, isolated species clusters at the basal nodes of the tree that are all restricted to the winter-rainfall zone of southern Africa (the Greater Cape floral kingdom) and a few, highly derived, large species groups that have radiated extensively within the winter-rainfall zone. Mapping of floral traits shows that an Iris-type flower is ancestral in Moraea. Floral changes are associated with shifts in pollination systems, either from passive pollen deposition on long-tongued bees foraging for nectar to active pollen collection by female bees foraging for pollen, fly, or hopliine scarab beetle pollination. Dating the nodes of the phylogenetic tree using non-parametric rate smoothing with a calibration point derived from broad dating of the angiosperms indicates that the divergence between Moraea and its sister genus Ferraria occurred about 25 mya in the early Miocene. The early radiation of Moraea took place against a background of aridification and the spread of open habitats, such as desert, shrubland, and fynbos.     

Phylogenetic and biogeographic diversification of Rhus (Anacardiaceae) in the Northern Hemisphere

Sequences of internal transcribed spacers (ITS) of nuclear ribosomal DNA, the chloroplast ndhF gene, and chloroplast trnL-F regions (trnL intron, and trnL [UAA] 3' exon-trnF [GAA] intergenic spacer) were used for phylogenetic analyses of Rhus, a genus disjunctly distributed in Asia, Europe, Hawaii, North America, and Northern Central America. Both ITS and cpDNA data sets support the monophyly of Rhus. The monophyly of subgenus Rhus was suggested by the combined cpDNA and ITS data, and largely supported in the cpDNA data except that Rhus microphylla of subgenus Lobadium was nested within it. The monophyly of subgenus Lobadium was strongly supported in the ITS data, whereas the cpDNA data revealed two main clades within the subgenus, which formed a trichotomy with the clade of subgenus Rhus plus R. microphylla. The ITS and cpDNA trees differ in the positions of Rhus michauxii, R. microphylla, and Rhus rubifolia, and hybridization may have caused this discordance. Fossil evidence indicates that Rhus dates back to the early Eocene. The penalized likelihood method was used to estimate divergence times, with fossils of Rhus subgenus Lobadium, Pistacia and Toxicodendron used for age constraints. Rhus diverged from its closest relative at 49.1+/-2.1 million years ago (Ma), the split of subgenus Lobadium and subgenus Rhus was at 38.1+/-3.0 Ma. Rhus most likely migrated from North America into Asia via the Bering Land Bridge during the Late Eocene (33.8+/-3.1 Ma). Rhus coriaria from southern Europe and western Asia diverged from its relatives in eastern Asia at 24.4+/-3.2 Ma. The Hawaiian Rhus sandwicensis diverged from the Asian Rhus chinensis at 13.5+/-3.0 Ma. Subgenus Lobadium was inferred to be of North American origin. Taxa of subgenus Lobadium then migrated southward to Central America. Furthermore, we herein make the following three nomenclatural combinations: (1) Searsia leptodictya (Diels) T. S. Yi, A. J. Miller and J. Wen, comb. nov., (2) Searsia pyroides (A. Rich.) T. S. Yi, A. J. Miller and J. Wen, comb. nov., and (3) Searsia undulata (Jacq.) T. S. Yi, A. J. Miller and J. Wen, because our analyses support the segregation of Searsia from Rhus.     

New platyrrhine monkeys from the Solimões Formation (late Miocene, Acre State, Brazil)

We report here a new fossil primate from the late Miocene of Brazil. The material consists of a lower first molar and a maxilla with P3-4. The fossils were collected in the Solim?es Formation at the locality of Patos, upper Acre River, Acre State, Brazil. The locality is assigned to the Huayquerian South American Land Mammal Age based on faunal content (late Miocene; dated to between 9 and 6 Ma). The new material is the oldest known occurrence of fossil primates in Brazil and is recognized as a new genus and species, Solimoea acrensis. Solimoea is the oldest known member of the ateline subfamily, which includes the living genera Ateles, Lagothrix, and Brachyteles. By analogy with the molar structures and diets of extant platyrrhines, Solimoea primarily had a diet of fruit, perhaps similar to that of the spider monkey, Ateles. Two other primate teeth described previously from the same formation in Bolivia document the occurrence of alouattines and cebines. One of those specimens is a late Miocene representative of the middle Miocene Colombian genus Stirtonia. The other represents one of the largest known platyrrhine primates, for which is erected a new primate genus, Acrecebus fraileyi.     

Pleistocene and pre-Pleistocene Begonia speciation in Africa

This paper presents a historical biogeographic analysis of African Begonia based on combined internal transcribed spacer (ITS) and trnL intron sequences. Age range estimates for Begonia in Africa ranged from only 1.5 Ma for some terminal nodes to 27 Ma for basal nodes when the ages of Réunion (2 Ma) andMayotte (5.4 Ma) were used to date the split between Begonia salaziensis and Begonia comorensis. Assuming a more recent origin age for Begonia salaziensis (2 Ma) provided age estimates in other parts of the phylogeny which agreed with patterns observed in other African organisms. A large proportion of the Begonia diversity seen today in Africa is of pre-Pleistocene origin. Species of Pleistocene origin are concentrated in species-rich groups such as sections Loasibegonia, Scutobegonia, and Tetraphila, which have their centre of diversity in western Central Africa. Phylogenetically isolated taxa such as Begonia longipetiolata, Begonia iucunda, and Begonia thomeana date to the late Miocene, a period of extended aridification on the African continent that had severe effects on African rain forest species. A general pattern is identified where phylogenetically isolated species occur outside the main identified rain forest refuges. Endemic species on the island of S?o Tomé such as Begonia baccata, Begonia molleri, and Begonia subalpestris appear to be palaeoendemics. Of these species, the most recent age estimate is for B. baccata, which is dated at ca. 3 Ma. Therefore, S?o Tomé appears to have functioned as an important (if previously unrecognised) pre-Pleistocene refuge. On the mainland, areas such as the Massif of Chaillu in Gabon, southern Congo (Brazzaville), and far western areas of Congo (Kinshasa) have played similar roles to S?o Tomé.     

Phylogenetics of the grass genus Ehrharta: evidence for radiation in the summer-arid zone of the South African Cape

Abstract Climatic and geological change may play a key role in stimulating biological radiations. Here, we use phylogenetic data to test whether the comparatively high diversity of ehrharteoid grasses in the Cape region of South Africa is the result of rapid radiation associated with the onset of a seasonally arid climate during the late Miocene. A phylogenetic hypothesis based on morphological and nucleotide sequence (nuclear ITS1 and plastid trn L-F) data confirms the monophyly of the African Ehrharta species and shows that the diversification of this lineage was centered in the Cape region. Sequence divergence data (ITS1 + trn L-F) indicate a pulse of rapid speciation, which may explain poor phylogenetic resolution within the African Ehrharta clade. Alternative calibrations yield a broad range of time estimates for the start and end of this radiation, most of which indicate a radiation inside the last 11 million years. A calibration based on the age of Ehrhartoideae suggests that radiation started 9.82 ± 0.20 million years ago and ended 8.74 ± 0.21 million years ago. Under alternative calibrations, estimated speciation rates during the period of radiation range between 0.87 and 4.18 species per million years. Parsimony optimization of habitat parameters reveals that radiation was correlated with the occupation of seasonally arid succulent karoo environments, wet heathy (fynbos) environments being ancestral. These data support earlier suggestions that late Miocene climatic change stimulated floristic radiation at the Cape, and highlight the potential importance of environmental change in powering diversification in continental floras.     

Molecular phylogeny of the antitropical subgenus Goniistius (Perciformes: Cheilodactylidae: Cheilodactylus): evidence for multiple transequatorial divergences and non-monophyly

The subgenus Goniistius comprises eight species of marine nearshore fishes which are antitropically distributed. The molecular phylogeny of these and other cheilodactylids was reconstructed from cytochrome oxidase I and cytochrome b mitochondrial DNA sequences. The placement within Goniistius of the morphologically divergent species Cheilodactylus (G.) nigripes was not supported. The remaining seven species are sufficiently divergent from other cheilodactylids to be designated as a separate genus. The antitropical distribution oi Goniistius is the result of three transequatorial divergences, which occurred during two periods. Based on molecular clock calibrations, these periods are suggested to be the mid Miocene, and late Miocene to early Pliocene. It is not known in which direction or by which mechanism these transequatorial divergences occurred, although biogeographic hypotheses of Mesozoic or Pleistocene separations can be discounted. The degree of genetic divergence between North and South Pacific populations of C. (G.) mttatus Garrett indicates that they have undergone cryptic speciation.     

A new species of the genus Microbunodon (Anthracotheriidae, Artiodactyla) from the Miocene of Pakistan: genus revision, phylogenetic relationships and palaeobiogeography

Lihoreau, F., Blondel, C., Barry, J. & Brunet, M. (2004). A new species of the genus Microbunodon (Anthracotheriidae, Artiodactyla) from the Miocene of Pakistan: genus revision, phylogenetic relationships and palaeobiogeography. — Zoologica Scripta , 33 , 97–115.
New unpublished remains of small Anthracotheriinae are described. First, materials from the upper Oligocene (MP 30) locality of La Milloque, southwest France, permit a review of the species Microbunodon minimum . Thereafter, fossils from the middle and late Miocene of the Potwar Plateau, Pakistan are attributed to the European genus Microbunodon . Microbunodon milaensis sp. n. from the Nagri Formation (between 10.3 and 9.2 Ma), Pakistan, is described and the species M. silistrensis from the Lower Manchar Formation (between 16 and 15 Ma) and from the Chinji Formation (between 12.7 and 11.5 Ma), Pakistan, is reviewed. The new species represents the last occurrence of the subfamily Anthracotheriinae, around 9.3 Ma. Similar materials from the Bugti and Siwalik Hills were previously considered as a small Anthracotherium . Comparisons with M. minimum from the European late Oligocene lead to a complete revision of the genus and permit definition of a new set of characters, which separate Microbunodon from Anthracotherium . A cladistic analysis reconsiders phylogenetic relationships among Anthracotheriinae, separating an Anthracothema–Anthracotherium clade and an Anthracokeryx–Microbunodon clade. Microbunodon appears to stem from the Asian late Eocene–lower Oligocene genus Anthracokeryx . These results imply a new distribution of the genus Microbunodon showing exchanges between Europe and Asia during the late Oligocene and probably the lower Miocene.     

Phylogeny of the Afro-Madagascan Aristea (Iridaceae) revisited in the light of new data on pollen morphology

Scanning and electron microscopic studies of the pollen grains of several species of the sub-Saharan African and Madagascan genus Aristea of c . 50 species, supplement data from an earlier study and enable us to include these species in a phylogenetic analysis. TEM examination of a range of pollen grains of representative pollen types in Aristea makes it possible to reinterpret past SEM results so that apertures covered with exine masses and previously called sulculate, are now seen to be either zonasulculate or dizonasulculate. Revised and expanded data are combined in a matrix together with new data for seed morphology in the genus, which like the capsules is remarkably diverse, including angular seeds with reticulate sculpturing, lamellate or triangular-columnar seeds with smooth or reticulate surfaces and with smooth or papillate or areolate margins. This new phylogenetic analysis suggests that Aristea should remain treated as comprising three subgenera. In subgenus Eucapsulares we add a new section Latifolieae for A. latifolia , which seems an isolated relic restricted to the mountains of the Cape Region of South Africa, an area of winter rainfall. Species with similar unspecialized capsule and seed morphology occur only in eastern south and tropical Africa and Madagascar, but they have derived pollen morphology whereas the pollen of A. latifolia is plesiomorphic. This species appears to be the closest living ancestor of the genus. In subgenus Aristea we continue to recognize three sections, sect. Racemosae , sect. Singulares and sect. Aristea. Nesting of subgenus Pseudaristea within section Racemosae of subgenus Aristea is probably an artefact of the analysis and does not receive bootstrap support, and hence we continue to recognize this group of species at subgeneric rank.  © 2004 The Linnean Society of London, Botanical Journal of the Linnean Society , 2004, 144 , 41–68.     

On the affinities of three fossil ocypodid crabs and their relevance to the time and place of origin of the genus Macrophthalmus (Crustacea: Brachyura)

All the species of Macrophthalmus described from fossil material belong to the still extant subgenus Venitus. M. vindobonensis and M. aquensis , the two earliest known species of the genus, are closely related to the extant M. latreillei , while M. guamensis is conspecific with M. leptophthalmus.
Doubt is cast on Remy's (1952) theory of the late Tertiary migration of Macrophthalmus from the Mediterranean, its suggested place of origin, to the Indo-West Pacific, its present distributional range. Instead it is proposed that Macrophthalmus arose in the Indo-West Pacific during the late Eocene-Oligocene, and extended its range into the Mediterranean during the early-middle Miocene. The Mediterranean species became isolated during the late Miocene by a land bridge, and probably became extinct following the elevation of their brackish habitat during the late Miocene-Pliocene.     

Phylogeny and biogeography of the hollies (Ilex L., Aquifoliaceae)

The holly genus, Ilex L., in the monogeneric Aquifoliaceae, is the largest woody dioecious genus (>664 spp.), with a near‐cosmopolitan distribution in mesic environments. We constructed a phylogeny based on two nuclear genes, representing 177 species spread across the geographical range, and dated using macrofossil records. The five main clades had a common ancestor in the early Eocene, much earlier than previously suggested. Ilex originated in subtropical Asia and extant clades colonized South America by 30 Ma, North America by 23 Ma, Australia by 8 Ma, Europe by 6 Ma, and Africa by 4 Ma. South and North America were colonized multiple times. Ilex also reached Hawaii (10 Ma) and other oceanic islands. Macrofossil and pollen records show the genus has tracked mesic climates through time and space, and had a wider distribution before late Miocene global cooling. Our phylogeny provides a framework for studies in comparative ecology and evolution.     

A new species of fossil Scutus Montfort, 1810 from New Zealand (Mollusca: Gastropoda: Fissurellidae)

ABSTRACT

A new fossil species of the genus Scutus (Scutus mirus n. sp.) is described from five Late Oligocene to Early Miocene (Waitakian to Altonian; 25.2–15.9?Ma) localities in the South Island, New Zealand. It is one of the oldest fossil species of Scutus known and probably inhabited very shallow, sub-tropical waters surrounding Zealandia during this time. The holotype of Scutus petrafixus Finlay, 1930 is re-examined; it is possibly from All Day Bay, Kakanui (Waitakian 25.2–21.7?Ma). The New Zealand species documented herein significantly expand our understanding of the fossil record of this shallow-marine molluscan lineage, and by proxy, also indicate the presence of very shallow coastal marine environments around the late Oligocene and early Miocene in southern Zealandia.     

Climate‐driven rampant speciation of the Cape flora

Aim To test whether the radiation of the extremely rich Cape flora is correlated with marine‐driven climate change. Location Middle to Late Miocene in the south‐east Atlantic and the Benguela Upwelling System (BUS) off the west coast of South Africa. Methods We studied the palynology of the thoroughly dated Middle to Late Miocene sediments of Ocean Drilling Program (ODP) Site 1085 retrieved from the Atlantic off the mouth of the Orange River. Both marine upwelling and terrestrial input are recorded at this site, which allows a direct correlation between changes in the terrestrial flora and the marine BUS in the south‐east Atlantic. Results Pollen types from plants of tropical affinity disappeared, and those from the Cape flora gradually increased, between 10 and 6 Ma. Our data corroborate the inferred dating of the diversification in Aizoaceae c. 8 Ma. Main conclusions Inferred vegetation changes for the Late Miocene south‐western African coast are the disappearance of Podocarpus‐dominated Afromontane forests, and a change in the vegetation of the coastal plain from tropical grassland and thicket to semi‐arid succulent vegetation. These changes are indicative of an increased summer drought, and are in step with the development of the southern BUS. They pre‐date the Pliocene uplift of the East African escarpment, suggesting that this did not play a role in stimulating vegetation change. Some Fynbos elements were present throughout the recorded period (from 11 Ma), suggesting that at least some elements of this vegetation were already in place during the onset of the BUS. This is consistent with a marine‐driven climate change in south‐western Africa triggering substantial radiation in the terrestrial flora, especially in the Aizoaceae.     

Orbital forcing and the spread of C4 grasses in the late Neogene: stable isotope evidence from South African speleothems

Reconstructing Plio-Pleistocene African paleoenvironments is important for models of early hominin evolution, but is often hampered by low-resolution or discontinuous climatic data. Here, we present high-resolution stable oxygen and carbon isotope time series data from two flowstones (secondary cave deposits) from the South African hominin-bearing Makapansgat Valley. The age of the older of the two flowstones (Collapsed Cone) is constrained by magnetostratigraphy to approximately 4-5 Ma; the younger flowstone (Buffalo Cave) grew between 2.0-1.5 Ma, as determined by magnetostratigraphy and orbital tuning of the isotopic data. The carbon isotope data is used as a proxy for the proportion of C(4) grasses in the local environment and the oxygen isotope data reflects monsoon rainfall intensity. The carbon isotope evidence indicates that in the late Miocene/early Pliocene, the local environment was dominated by C(3) vegetation, whereas, in the Plio-Pleistocene, it was composed of a mixture of C(3) and C(4) vegetation. This suggests that C(4) grasses became a significant part of the Makapansgat Valley ecosystem at approximately 4-5 Ma, towards the end of the late Neogene global expansion of C(4) grasses. After this initial expansion, South Africa experienced further fluctuations in the proportion of C(3) and C(4) vegetation during the Plio-Pleistocene, in response to regional and global climatic changes. Most notably, the Buffalo Cave flowstone provides evidence for C(4) grass expansion at ca. 1.7 Ma that we suggest was a response to African aridity caused by the onset of the Walker Circulation in the Pacific Ocean at this time.     

Phylogeny of the subgenus Culicoides and related species in Italy, inferred from internal transcribed spacer 2 ribosomal DNA sequences

Biting midges of the genus Culicoides (Diptera: Ceratopogonidae) include vectors for the economically important animal diseases, bluetongue (BT) and African horse sickness (AHS). In the Mediterranean Basin, these diseases are transmitted by four species of Culicoides: the first three belong in the subgenus Avaritia Fox and are Culicoides imicola Kieffer, Culicoides obsoletus (Meigen) and Culicoides scoticus Downes and Kettle; the fourth is Culicoides pulicaris (Linnaeus) in the subgenus Culicoides Latreille. In the Palaearctic Region, this subgenus (usually referred to as the C. pulicaris group) now includes a loose miscellany of some 50 taxa. The lack of clarity surrounding its taxonomy stimulated the present morphological and molecular study of 11 species collected in Italy. Phylogenetic analysis of nuclear ribosomal DNA internal transcribed spacer 2 (ITS2) sequence variation demonstrated a high degree of divergence. These results, combined with those from a parallel morphological study, disclosed: (1) that some previously described taxa should be resurrected from synonymy; (2) that there are new species to be described; (3) that the subgenus Culicoides (as currently employed) is a polyphyletic assemblage of four lineages - the subgenus Culicoides sensu stricto, the subgenus Silvicola Mirzaeva and Isaev, the subgenus Hoffmania Fox and the hitherto unrecognized Fagineus species complex. Each is discussed briefly (but not defined) and its constituent Palaearctic taxa listed. Strong congruence between morphological and molecular data holds promise for resolving many of the difficult taxonomic issues plaguing the accurate identification of vector Culicoides around the world.