On the systematic status of the desert plated lizard (Angolosaurus skoogi): phylogenetic inference from DNA sequence analysis of the African Gerrhosauridae

The desert plated lizard ( Angolosaurus skoogi ), a 'sand sea' endemic of the northern Namib Desert, exhibits remarkable morphological convergence with other dune-dwelling lizards worldwide. This distinct ecomorphic condition sets Angolosaurus apart from the remaining genera in the family Gerrhosauridae. Indeed, a morphological phylogeny addressing generic relationships within the Cordyliformes (Cordylidae + Gerrhosauridae) identified Angolosaurus as the earliest diverging taxon among African gerrhosaurids. We re-evaluated the basal status of Angolosaurus , conducting a molecular phylogenetic analysis of the African and Madagascan gerrhosaurid genera. Our survey involved a comprehensive species-level comparison among the four nominal genera of mainland Africa ( Angolosaurus , Cordylosaurus , Tetradactylus and Gerrhosaurus ). Mitochondrial DNA sequence data from the cytochrome b , ND2, 12S and 16S rRNA genes were combined for analysis using both parsimony and maximum likelihood procedures. In contrast to the morphological hypothesis, our results do not depict Angolosaurus as the sister taxon to other African gerrhosaurids. Rather, the molecular analyses consistently place Angolosaurus within Gerrhosaurus , rendering the latter genus paraphyletic.  © 2003 The Linnean Society of London, Biological Journal of the Linnean Society , 2003, 78 , 253–261.     

Phylogeny and biogeography of African Biomphalaria (Gastropoda: Planorbidae), with emphasis on endemic species of the great East African lakes

The pulmonate snails of the genus Biomphalaria are widely distributed in the tropics, and they are intermediate hosts of the digenean trematode Schistosoma mansoni that causes intestinal schistosomiasis in humans. Recent molecular evidence suggests that Biomphalaria originated in South America, and following a recent transatlantic migration colonized Africa, where it radiated into the currently recognized 12 species. In the present study we further investigate the internal phylogenetic relationships of African Biomphalaria with emphasis on the dispersal and speciation on the continent, especially in the Great Lakes in East Africa. Our results, based on 16S ribosomal DNA, cytochrome oxidase subunit I (COI), and ribosomal internal transcribed spacer I (ITS1), support the monophyly of an African clade with two separate lineages ( Biomphalaria pfeifferi / Biomphalaria camerunensis and the Nilotic species complex/ Biomphalaria angulosa ). Following the initial colonization of Africa, Biomphalaria spread towards the east where a later radiation occurred in the Lake Victoria basin and the Albertine Rift Valley Lakes. With further dispersal along the River Nile, additional speciation took place giving origin to the North-east African species Biomphalaria alexandrina . Our results present almost no support of the species groups of Mandahl-Barth (except for the pfeifferi group), which is in accordance with other molecular appraisals. Our results suggest that Biomphalaria stanleyi , which is endemic to Lake Albert, is not an ecophenotype of the continental B. pfeifferi as previously suggested by other molecular studies. B. angulosa is sequenced for the first time and it is inferred to have an important phylogenetic position as sister group to the Albertine Rift/Lake Victoria basin radiation.  © 2007 The Linnean Society of London, Zoological Journal of the Linnean Society , 2007, 151 , 337–349.     

Biogeography of the grasses (Poaceae): a phylogenetic approach to reveal evolutionary history in geographical space and geological time

The grasses (Poaceae) are the fifth most diverse family of angiosperms, including 800 genera and more than 10 000 species. Few phylogenetic studies have tried to investigate palaeo‐biogeographical and palaeo‐ecological scenarios that may have led to present‐day distribution and diversity of grasses at the family level. We produced a dated phylogenetic tree based on combined plastid DNA sequences and a comprehensive sample of Poaceae. Furthermore, we produced an additional tree using a supermatrix of morphological and molecular data that included all 800 grass genera so that ancestral biogeography and ecological habitats could be inferred. We used a likelihood‐based method, which allows the estimation of ancestral polymorphism in both biogeographical and ecological analyses for large data sets. The origin of Poaceae was retrieved as African and shade adapted. The crown node of the BEP + PACCMAD clade was dated at 57 Mya, in the early Eocene. Grasses dispersed to all continents by approximately 60 million years after their Gondwanan origin in the late Cretaceous. PACCMAD taxa adapted to open habitats as early as the late Eocene, a date consistent with recent phytolith fossil data for North America. C₄ photosynthesis first originated in Africa, at least for Chloridoideae in the Eocene at c. 30 Mya. The BEP clade members adapted to open habitats later than PACCMAD members; this was inferred to occur in Eurasia in the Oligocene. © 2010 The Linnean Society of London, Botanical Journal of the Linnean Society, 2010, 162 , 543–557.     

The importance of looking at small-scale patterns when inferring Gondwanan biogeography: a case study of the centipede Paralamyctes (Chilopoda, Lithobiomorpha, Henicopidae)

Gondwanan biogeography has fascinated zoologists and botanists for over a century, but most biogeographical work has used continent-scale areas as analytical units. More finely resolved patterns, as can be obtained from small invertebrates with limited dispersal abilities, will be obscured in those studies. A common case is treating Australia as a single biogeographical region. In the present study, the necessity of splitting Australia into multiple microareas is demonstrated using centipedes as an example. The lithobiomorph centipede Paralamyctes is distributed on fragments of Gondwana, with species in southern Africa, Madagascar, southern India, Patagonia, eastern Australia, and New Zealand. A cladogram for Paralamyctes is based on morphology and sequences for four molecular markers for 30 terminals that sample 20 of 26 known ingroup species and four outgroups. Analysis with direct optimization across a range of indel costs and transversion : transition cost ratios identifies two main clades: Paralamyctes ( Paralamyctes ) unites species from southern Africa, Madagascar, tropical and warm temperate Australia, and New Zealand. The other group includes the temperate Australian/New Zealand Paralamyctes ( Haasiella ) and Paralamyctes ( Thingathinga ) and a Chilean clade. Subtree analysis finds that different parts of Australia have closest affinities to other Gondwanan fragments, and some of these relationships (such as that between north Queensland and New Zealand) are based on taxonomically stable clades. Area delimitation for large continental fragments should use sufficiently fine resolution to test the 'monophyly' of those fragments and attempt to eliminate spurious geographical paralogy.  © 2006 The Linnean Society of London, Biological Journal of the Linnean Society , 2006, 89 , 65–78.     

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.     

Swimming out of Africa: mitochondrial DNA evidence for late Pliocene dispersal of a cichlid from Central Africa to the Levant

The rich Levantine fauna and flora were shaped by millions of years of migration across the region, from Africa to Eurasia and vice versa. Most large-scale processes that led to this diversity have been relatively well studied. However, small-scale processes, and details such as the area of origin of particular groups, and the route and time of dispersal are often not as clear. This is the case with the endemic Levantine representatives of the fish family Cichlidae. In this work we combine genetic, palaeontological and geological data in an attempt to understand the dispersal of the cichlid fish Astatotilapia flaviijosephi (Lortet, 1883) from sub-Saharan Africa to the Levant. A. flaviijosephi is unique among the Levantine cichlids in being the only non-tilapiine. It is also the only haplochromine cichlid to be found out of Africa. A partial sequence of the control region of the mitochondrial DNA was used to determine A. flaviijosephi 's phylogenetic relationships with other African haplochromines, and to estimate its time of divergence from this group. Combining our findings with palaeontological and geological data, we suggest that A. flaviijosephi separated from the other haplochromines during the middle to late Pliocene (2.5–3.3 Mya) and probably dispersed from Africa to the Levant via the Nile.  © 2004 The Linnean Society of London, Biological Journal of the Linnean Society , 2004, 82 , 103–109.     

Intercontinental distribution of Plagiochila corrugata (Plagiochilaceae, Hepaticae) inferred from nrDNA ITS sequences and morphology

Plagiochila sect. Vagae is a large pantropical clade that is characterized morphologically by frequent terminal branching, vegetative distribution by propagules on the ventral surface of the leaves and a capsule wall with thickenings in all layers. Plagiochila corrugata from Brazil is characterized by strongly undulate, toothed leaf margins and represents the only known neotropical species of sect. Vagae with unispiral elaters. Plagiochila cambuena from Madagascar is distinguished by the same features. Maximum likelihood and parsimony analyses of 38 nrDNA ITS sequences of Plagiochila reveal P. corrugata and P. cambuena in a weakly (ML) to well (MP) supported monophyletic lineage within P.  sect.  Vagae . As an outcome of the morphological and molecular investigation, P. cambuena is relegated to the synonymy of P. corrugata. Plagiochila corrugata is placed in a Vagae -subclade with 11 further American species. The range of P. corrugata can be ascribed to long-range dispersal from the Neotropics rather than a Gondwanan distribution. Species from tropical Asia and Africa are placed at the base of the Vagae clade. Branch length within P.  sect.  Vagae points to a sudden radiation.  © 2004 The Linnean Society of London, Botanical Journal of the Linnean Society , 2004, 146 , 469–481.     

Molecular evidence suggests an ancient radiation for the fairy shrimp genus Streptocephalus (Branchiopoda: Anostraca)

Phylogenetic relationships among assumed Gondwanan aquatic inland invertebrate fauna are generally largely neglected, and biogeographical hypotheses for these organisms are generally inferred from historic (palaeogeographical) and contemporary distribution patterns. The distribution of the monogeneric thermophilic freshwater fairy shrimp family Streptocephalidae ( Streptocephalus ) provides a particularly useful framework to test the three contrasting biogeographical scenarios proposed for the evolution of this group: (1) the genus evolved in Laurasia and subsequently dispersed into Africa and North America; (2) the genus evolved and dispersed out of Africa and (3) the current distribution of the genus is the result of vicariance following the fragmentation of Gondwana. In the present study, the phylogenetic relationships of species in this genus are examined with the use of two mitochondrial genes (12S rRNA and COI mtDNA), while the phylogenetic relationships among the North American species and selected African taxa was investigated using the nuclear fragment (5.8S-ITS-1-18S). Phylogenetic results indicate that Streptocephalus probably evolved in Gondwana and that the current distribution patterns are a consequence of a combination of vicariance and limited dispersal. The implications for the evolution of continental freshwater crustaceans are discussed.  © 2004 The Linnean Society of London, Biological Journal of the Linnean Society , 2004, 82 , 313–327.     

‘Back to Africa’: increased taxon sampling confirms a problematic Australia‐to‐Africa bee dispersal event in the Eocene

Colletidae is a predominantly southern hemisphere bee family with a Late Cretaceous origin and with an inferred ancestral region covering late Gondwanan South America, Antarctica and Australia. One highly diverse colletid subfamily, Euryglossinae, is entirely restricted to Australia and the strictly Afrotropical subfamily Scrapterinae has been inferred as its sister clade. This has led to suggestions that Scrapterinae represents a highly unusual post‐Gondwanan dispersal from Australia to Africa, but phylogenetic studies to date have included only minimal representatives from each subfamily. Here we greatly increase the level of species sampling of both subfamilies and develop a molecular phylogeny based on one mitochondrial and two nuclear genes. Our results indicate that the broad results of earlier studies are robust to substantially greater taxon sampling, and we infer a divergence date between the two subfamilies in the early Eocene. Dispersal pathways between Africa and Australia during that time are problematic, with several studies suggesting dispersals via the now largely submerged Kerguelen and Crozet Plateaus. Our results contribute another example of a puzzling sister‐clade relationship between African and Australian taxa and indicate the need to better understand southern hemisphere subaerial configurations, including Antarctica, and ocean and wind currents at those times.     

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.     

A new species of Orycteropodidae (Mammalia, Tubulidentata) in the Mio-Pliocene of northern Chad

A new species of fossil Tubulidentata has been found by the Mission Paléoanthropologique Franco-Tchadienne in Northern Chad. It is the first fossil Orycteropodidae (aardvark) from the Mio-Pliocene of Central Africa. The new taxon, Orycteropus abundulafus sp. nov. , is considered in the framework of the available Orycteropodidae fossil record. The Chadian specimen is characterized by the highest dental robustness index among all Tubulidentata, the presence of crests on the pterygoid, the triangular-shaped olecranon fossa and the reduction of the deltoid crest. All of these characters are linked to a less fossorial animal that had a tougher diet. This new African species is closer to the Eurasian O. gaudryi than to any other Tubulidentata. Together they form a clade distinct from that which includes O. afer . This is the first evidence of a relationship for aardvarks between Africa and Eurasia. An initial step is made towards revision of the phylogeny of the order.  © 2005 The Linnean Society of London, Zoological Journal of the Linnean Society , 2005, 143 , 109–131.     

Evaluating trans-tethys migration: an example using acrodont lizard phylogenetics

A phylogenetic tree for acrodont lizards (Chamaeleonidae and Agamidae) is established based on 1434 bases (1041 informative) of aligned DNA positions from a 1685-1778 base pair region of the mitochondrial genome. Sequences from three protein-coding genes (ND1, ND2, and COI) are combined with sequences from eight intervening tRNA genes for samples of 70 acrodont taxa and two outgroups. Parsimony analysis of nucleotide sequences identifies eight major clades in the Acrodonta. Most agamid lizards are placed into three distinct clades. One clade is composed of all taxa occurring in Australia and New Guinea; Physignathus cocincinus from Southeast Asia is the sister taxon to the Australia-New Guinea clade. A second clade is composed of taxa occurring from Tibet and the Indian Subcontinent east through South and East Asia. A third clade is composed of taxa occurring from Africa east through Arabia and West Asia to Tibet and the Indian Subcontinent. These three clades contain all agamid lizards except Uromastyx, Leiolepis, and Hydrosaurus, which represent three additional clades of the Agamidae. The Chamaeleonidae forms another clade weakly supported as the sister taxon to the Agamidae. All eight clades of the Acrodonta contain members occurring on land masses derived from Gondwanaland. A hypothesis of agamid lizards rafting with Gondwanan plates is examined statistically. This hypothesis suggests that the African/West Asian clade is of African or Indian origin, and the South Asian clade is either of Indian or Southeast Asian origin. The shortest tree suggests a possible African origin for the former and an Indian origin for the latter, but this result is not statistically robust. The Australia-New Guinea clade rafted with the Australia-New Guinea plate and forms the sister group to a Southeast Asian taxon that occurs on plates that broke from northern Australia-New Guinea. Other acrodont taxa are inferred to be associated with the plates of Afro-Arabia and Madagascar (Chameleonidae), India (Uromastyx), or southeast Asia (Hydrosaurus and Leiolepis). Introduction of different biotic elements to Asia by way of separate Gondwanan plates may be a major theme of Asian biogeography. Three historical events may be responsible for the sharp faunal barrier between Southeast Asia and Australia-New Guinea, known as Wallace's line: (1) primary vicariance caused by plate separations; (2) secondary contact of Southeast Asian plates with Eurasia, leading to dispersal from Eurasia into Southeast Asia, and (3) dispersal of the Indian fauna (after collision of that subcontinent) to Southeast Asia. Acrodont lizards show the first and third of these biogeographic patterns and anguid lizards exhibit the second pattern. Modern faunal diversity may be influenced primarily by historical events such as tectonic collisions and land bridge connections, which are expected to promote episodic turnover of continental faunas by introducing new faunal elements into an area. Repeated tectonic collisions may be one of the most important phenomena promoting continental biodiversity. Phylogenetics is a powerful method for investigating these processes.     

The genetic population structure of Buthus occitanus (Scorpiones: Buthidae) across the Strait of Gibraltar: calibrating a molecular clock using nuclear allozyme variation

I assess here the importance of the Strait of Gibraltar as a barrier to gene flow for populations of the scorpion Buthus occitanus . This polytypic buthid scorpion occurs in Europe and in North Africa where it is morphologically more diverse. The phylogenetic relationship between B. occitanus populations across the Strait of Gibraltar is investigated by nuclear allozymes analysis (15 loci scored). Phylogenetic analysis based on estimated gene frequency data results in a tree topology that divides the populations into three clades, i.e. a European, an Atlas (= Morocco samples) and a Tell-Atlas clade (= Tunisian samples). The Tell-Atlas clade grouped with the European clade with a rather high bootstrap support of 70%. Within these clades low levels of genetic variability are observed. Calibrating a molecular clock under the assumption that the European populations are autochthonous and have been isolated from the North African for at least 5.33 Myr reveals a divergence rate of 0.060 genetic distance (D) per Myr estimated between European and Moroccan samples and 0.036D Myr⁻¹ between European and Tunisian samples, respectively.  © 2004 The Linnean Society of London, Biological Journal of the Linnean Society , 2004, 81 , 519–534.     

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.     

Phylogeny and biogeographical history of Trogoniformes, a pantropical bird order

With highly conserved morphology throughout the family, a tropical distribution, and no close living relatives, the trogons (Aves: Trogonidae) pose a difficult problem for systematists. Disjunct tropical distributions are often attributed to Gondwanan vicariance, but the fossil record for trogons is mostly from the Tertiary of Europe. This study examined support for the basal relationships among trogons using a combination of nuclear (RAG-1) and mitochondrial (ND2) DNA sequence data. Although some nodes could not be resolved with significant support, there is strong support for the basal position of three New World genera ( Pharomachrus , Euptilotis , and Priotelus ). This phylogenetic hypothesis differs markedly from previous studies of trogon relationships and taxonomic treatments. Biogeographically, it implies an origin and early vicariance events for the crown clade in the New World. Molecular divergence estimates place all of the basal nodes of the trogon phylogeny in the Oligocene, precluding a Gondwanan origin for modern trogons.  © 2005 The Linnean Society of London, Biological Journal of the Linnean Society , 2005, 84 , 725–738.     

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.     

Age and historical biogeography of the pantropically distributed Spathelioideae (Rutaceae,Sapindales)

Aim The family Rutaceae (rue family) is the largest within the eudicot order Sapindales and is distributed mainly in the tropical and subtropical regions of both the New World and the Old World, with a few genera in temperate zones. The main objective of this study is to present molecular dating and biogeographical analyses of the subfamily Spathelioideae, the earliest branching clade (which includes eight extant genera), to interpret the temporal and spatial origins of this group, ascertaining possible vicariant patterns and dispersal routes and inferring diversification rates through time. Location Pantropics. Methods A dataset comprising a complete taxon sampling at generic level (83.3% at species level) of Spathelioideae was used for a Bayesian molecular dating analysis (beast ). Four fossil calibration points and an age constraint for Sapindales were applied. An ancestral area reconstruction analysis utilizing the dispersal–extinction–cladogenesis model and diversification rate analyses was conducted. Results Dating analyses indicate that Rutaceae and Spathelioideae are probably of Late Cretaceous origin, after which Spathelioideae split into a Neotropical and a Palaeotropical lineage. The Palaeotropical taxa have their origin inferred in Africa, with postulated dispersal events to the Mediterranean, the Canary Islands, Madagascar and Southeast Asia. The lineages within Spathelioideae evolved at a relatively constant diversification rate. However, abrupt changes in diversification rates are inferred from the beginning of the Miocene and during the Pliocene/Pleistocene. Main conclusions The geographical origin of Spathelioideae probably lies in Africa. The existence of a Neotropical lineage may be the result of a dispersal event at a time in the Late Cretaceous when South America and Africa were still quite close to each other (assuming that our age estimates are close to the actual ages), or by Gondwanan vicariance (assuming that our age estimates provide minimal ages only). Separation of land masses caused by sea level changes during the Pliocene and Pleistocene may have been triggers for speciation in the Caribbean genus Spathelia.     

Tri-locus sequence data reject a “Gondwanan origin hypothesis” for the African/South Pacific crab genus Hymenosoma

Crabs of the family Hymenosomatidae are common in coastal and shelf regions throughout much of the southern hemisphere. One of the genera in the family, Hymenosoma, is represented in Africa and the South Pacific (Australia and New Zealand). This distribution can be explained either by vicariance (presence of the genus on the Gondwanan supercontinent and divergence following its break-up) or more recent transoceanic dispersal from one region to the other. We tested these hypotheses by reconstructing phylogenetic relationships among the seven presently-accepted species in the genus, as well as examining their placement among other hymenosomatid crabs, using sequence data from two nuclear markers (Adenine Nucleotide Transporter [ANT] exon 2 and 18S rDNA) and three mitochondrial markers (COI, 12S and 16S rDNA). The five southern African representatives of the genus were recovered as a monophyletic lineage, and another southern African species, Neorhynchoplax bovis, was identified as their sister taxon. The two species of Hymenosoma from the South Pacific neither clustered with their African congeners, nor with each other, and should therefore both be placed into different genera. Molecular dating supports a post-Gondwanan origin of the Hymenosomatidae. While long-distance dispersal cannot be ruled out to explain the presence of the family Hymenosomatidae on the former Gondwanan land-masses and beyond, the evolutionary history of the African species of Hymenosoma indicates that a third means of speciation may be important in this group: gradual along-coast dispersal from tropical towards temperate regions, with range expansions into formerly inhospitable habitat during warm climatic phases, followed by adaptation and speciation during subsequent cooler phases.     

The phylogeny of the Pantropical genus Arrhipis Bonvouloir (Coleoptera, Eucnemidae)

The phylogeny of the genus Arrhipis Bonvouloir (Coleoptera, Eucnemidae) is clarified with a cladistic analysis based on five molecular markers and morphology. Sixteen species from Africa, America, Asia, and Australia are included in the analysis. Two separate Asian clades are recovered, one of them being the sister group to a clade with the American and African species. With the exception of the continental south-east Asian species, all Gondwanan regions have monophyletic faunas. According to the present data, the continental south-east Asian fauna comprises two monophyletic groups, one of which is the sister group to African and American species. Vicariance seems to be the logical explanation for the distribution of these lignicolous beetles.
© The Willi Hennig Society 2009.