Evolution in action: molecular evidence for recent emergence of secotioid genera Endxoptychum, Gyrophragmium and Longula from Agaricus ancestors

Phylogenetic analyses of 29 Agaricus species and one representative from genera Endoptychum, Gyrophragmium and Longula each were conducted based on sequence data of the entire internal transcribed spacers and partial large subunit of ribosomal DNA. The Agaricus species formed several distinct clades both confirming and challenging previous morphological sections in several cases. Endoptychum depressum, Gyrophragmium dunalii and Longula texensis were nested among species of the genus Agaricus. This study provides evidence for independent emergence of these secotioid fungi from Agaricus ancestors, most likely from species placed in the section Arvenses as inferred from molecular data.     

Differentiation of commercial strains of <Emphasis Type="Italic">Agaricus</Emphasis> species in China with inter-simple sequence repeat marker

In recent years, Agaricus mushroom production has undergone an important increase in the world. However, there is no consensus over the taxonomy of some commercial strains. So rapid and reliable methods to identify Agaricus strains are urgently needed. In this paper, six inter-simple sequence repeat (ISSR) primers were screened from 20 primers to analyse the genetic diversities of the 12 main commercial strains of Agaricus in China. The results showed that a total of 124 bands were produced, among which 80.6% was polymorphic, the similarity coefficients ranged from 0.69 to 0.98. Twelve tested strains were divided into 2 groups on the similarity coefficient of 0.75. Three Agaricus bitorquis strains were clustered in one group, and seven Agaricus bisporus strains were clustered with Agaricus strains S1 and S2 in the other group. As comparison with ISSR, intergenic spacers combined with restriction fragment length polymorphisms (IGS-RFLP) was employed to analyse genetic diversities of the tested strains by digesting IGS amplified products with restriction enzymes Hae III, Alu I, Rsa I, respectively, and IGS-RFLP cluster results were similar with those of ISSR. However, ISSR was superior to IGS-RFLP in differentiating closely related strains. It is suggested that the ISSR marker is an alternative method to differentiate Agaricus strains.     

Lacustrine radiations in African Synodontis catfish

Synodontis catfish are a species‐rich, tropical pan‐African genus that predominately occur in fluviatile environments, but which also form a small radiation within Lake Tanganyika (LT). Here we estimate Synodontis relationships, based on mitochondrial and nuclear DNA, greatly expanding previous sampling. Data were analysed using different methods of phylogenetic inference: Bayesian (also testing compositional heterogeneity), likelihood and parsimony, in order to investigate biogeographic history and the extent of intralacustrine speciation within this group. Bayesian‐relaxed clock analyses were used to estimate timings of radiations. Our analyses reveal a single origin of the LT flock with the inclusion of the nonendemic S. victoriae, and that these taxa evolved relatively recently (5.5 Ma), considerably later than the formation of LT (9–12 Ma). Two internal endemic clades diversified at a similar time (2–2.5 Ma), corresponding to a period of climate change, when lake levels dropped. We find evidence for a further species flock, composed of riverine southern African taxa, the diversification of which is very rapid, 0.8 Ma (95% HPD: 0.4–1.5) and infer a similar scenario for the diversification of this flock to southern African serrachromine cichlids in that they radiated in the now extinct lake Makgadikgadi. We also reveal that the biogeographic history of Synodontis catfish is more complex than previously thought, with nonmonophyletic geographic species groupings.     

A phylogenetic and morphological overview of sections Bohusia, Sanguinolenti, and allied sections within Agaricus subg. Pseudochitonia with three new species from France,Iran, and Portugal

The genus Agaricus was recently rearranged to accommodate numerous tropical taxa. Accordingly, the genus was split into six subgenera and 22 sections of which 12 are included in A. subg. Pseudochitonia. Preliminary data indicated that three putative new species belong to this subgenus. Our objectives were to describe these species, to determine to which sections they belong, and to experience the interest of some traditional traits in this new context. We morphologically described Agaricus coniferarum from France and Portugal, Agaricus iranicus from Iran, and Agaricus lusitanicus from Portugal. Multi-gene phylogenetic analyses based on ITS, LSU, and tef1 sequence data of representatives of the 12 sections clearly indicated that A. coniferarum and A. lusitanicus are placed in Agaricus sect. Bohusia, while A. iranicus is in A. sect. Sanguinolenti. Incidentally, we replaced the illegitimate name Agaricus magnivelaris by Agaricus fiardianus. In a phylogenetic tree, based on all available ITS sequence data and focussing on six related sections, we examined the phylogenetic distribution of various characters. The intensity of red discolouration when the sporocarp is rubbed or cut appeared as a phylogenetically weak informative trait. We propose a determination key leading to a group of three hardly distinguishable sections (Bohusia, Nigrobrunnescentes, and Sanguinolenti).     

Does secotioid inertia drive the evolution of false-truffles?

Secotioid inertia is a model implemented to explain the prevalence of highly derived false-truffles with no obvious connection to the Homobasidiomycetes. The model accommodates the apparent lack of epigeous sister taxa for some highly derived hypogeous lineages by assuming that gasteromycetation in some fungi leads to the extinction of their epigeous sister population. The derived state of some hypogeous lineages suggests that they arose early in the evolution of Homobasidiomycetes and that those groups were subject to conditions that favoured hypogeous lineages such that the hypogeous fruit body form became the predominant form for some lineages. The directional selection component of secotioid inertia, termed secotioid drive, led to the extinction of their epigeous sister taxon. Morphological and molecular data from Russulaceae are used to model the evolutionary stages of secotioid inertia. The resulting phylogenetic results are compared with data from the order Leucogastrales, and the genus Destuntzia. The implications of secotioid drive are discussed with reference to gasteromycete phylogenetics, evolution, and conservation. Specifically, secotioid inertia can be used to account for reversals in fruit body morphology and instability in mycorrhizal formation.     

Molecular phylogeny and historical biogeography of the genus Tuber, the 'true truffles'

Aim Various data sets and methods of analysis were combined to produce the first comprehensive molecular phylogeny of the genus Tuber and to analyse its biogeography. Location Europe, North Africa, China, Asia, North America. Methods Phylogenetic relationships among Tuber species were reconstructed based on a data set of internal‐transcribed spacer (ITS) sequences and various phylogenetic inference methods, specifically maximum parsimony, Bayesian analysis and neighbour joining. Tajima’s relative rate test showed that Tuber 18S rRNA, 5.8S rRNA, 5.8S‐ITS2 rRNA and β‐tubulin sequences evolved in a clock‐like manner. These genes, combined or not, were employed for molecular clock estimates after construction of linearized trees using mega 3.1. We reconstructed ancestral areas in the Northern Hemisphere by means of a dispersal–vicariance analysis (diva 1.1) based on current distribution patterns of the genus Tuber determined from the literature. Results The resulting molecular phylogeny divided the genus Tuber into five distinct clades, in agreement with our previously published studies. The Puberulum, Melanosporum and Rufum groups were diversified in terms of species and geographical distribution. In contrast, the Aestivum and Excavatum groups were less diversified and were located only in Europe or North Africa. Using a global molecular clock analysis, we estimated the divergence times for the origin of the genus and for the origin of several groups. diva inferred nine dispersal events and suggested that the ancestor of Tuber was originally present in Europe or was widespread in Eurasia. Equally optimal distributions were obtained for several nodes, suggesting different possible biogeographical patterns. Main conclusions Our analyses identified several discrepancies with the classical taxonomy of the genus, and we propose a new phylogenetic classification. According to molecular clocks, the radiation of the genus Tuber could have started between 271 and 140 Ma. Used in combination with the results obtained from time divergence estimates, this allows us to propose two equally probable scenarios of intra‐ and inter‐continental diversification of the genus according to the geographic distribution of the most recent common ancestor in Europe or Eurasia. The biogeographical patterns imply intra‐continental dispersal events between Europe and Asia and inter‐continental dispersal events between North America and Europe or Asia, which are compatible with land connections during the Tertiary.     

Island life – classification,speciation and cryptic species of Pycnandra (Sapotaceae) in New Caledonia

Pycnandra (Sapotaceae), the largest endemic genus in New Caledonia, comprises 66 species classified in six subgenera. We tested phylogenetic relationships and a proposed infrageneric classification by sampling 60 species for sequences of nuclear ribosomal DNA (ETS, ITS, RPB2) and plastid DNA (trnH–psbA) and nine morphological characters. Data were analysed with Bayesian inference, parsimony jackknifing and lineage through time. We recovered a phylogenetic tree supporting the recognition of six proposed subgenera (Achradotypus, Leptostylis, Pycnandra, Sebertia, Trouettia and Wagapensia). Because a subgeneric classification is used, the nomenclature will be stable when the members are transferred to Pycnandra. Morphological traits were optimized in the BEAST analysis, adding evidence to earlier work that morphology has limited value for successfully diagnosing groups in Sapotaceae. We confirm a previously suspected case of cryptic species that exhibit the same morphological features and require the same abiotic conditions, but are distantly related in the phylogenetic tree. We detected two possible new cases of cryptic sibling species that might warrant recognition. A slowdown in speciation rate in several genera has been suggested as evidence that New Caledonia was once submerged after rifting from Australia. Plotting lineages through time reveals two important intervals at 7.5–8.6 Ma and present to 1.5 Ma, when net molecular diversification within the genus was zero. This indicates that the genus presently has reached a dynamic equilibrium, providing additional evidence that New Caledonia is an old Darwinian island, being submerged during the Eocene and colonized after re‐emergence c. 37 Ma. © 2015 The Linnean Society of London, Botanical Journal of the Linnean Society, 2015, 179 , 57–77.     

Effect of different Agaricus species on the development of the mushroom sciarid fly Lycoriella ingenua

Isolates from eight species of Agaricus were investigated in laboratory experiments for their effect on the development of the mushroom sciarid fly, Lycoriella ingenua (Dufour) (Diptera: Sciaridae), which is an important pest of the white mushroom Agaricus bisporus (Lange) Imbach (Agaricaceae). The population levels of L. ingenua developing in compost inoculated with Agaricus mycelium varied with the Agaricus isolate used, with some isolates causing high levels of inhibition. The development of L. ingenua populations and the survival of larval instars were inversely proportional to the amount of Agaricus inoculum applied. There was also a negative relationship between L. ingenua survival and the extension rate of the Agaricus isolate in compost. The results suggest that inhibition of L. ingenua population development by Agaricus is linked to the rate at which compost is colonized by fungal mycelium. Therefore, on mushroom farms, sciarid fly control should focus on protection of the compost before it has become colonized by mycelium, as this is when it is most vulnerable to the pest.     

Out of Alaska: morphological diversity within the genus <Emphasis Type="Italic">Eurytemora</Emphasis> from its ancestral Alaskan range (Crustacea,Copepoda)

The copepod genus Eurytemora occupies a wide range of habitat types throughout the Northern Hemisphere, with among the broadest salinity ranges of any known copepod. The epicenter of diversity for this genus lies along coastal Alaska, where several species are endemic. Systematic analysis has been difficult, however, because of a tendency toward morphological stasis in this genus, despite large genetic divergences among populations and species. The goals of this study were to (1) analyze patterns of morphological variation and divergence within this genus, focusing on Eurytemora species that occur in North America, and (2) determine patterns of geographic and salinity distribution of Eurytemora species within the ancestral range in Alaska. We applied a comparative multivariate morphological analysis using 16–26 characters from 125 specimens from 20 newly collected sites in Alaska and 15 existing samples predominantly from North America. Results from principal component and hierarchical cluster analyses identified seven distinct morphological species of Eurytemora in North America (E. affinis, E. americana, E. canadensis, E. composita, E. herdmani, E. pacifica, and E. raboti), and identified diagnostic characters that distinguish the species (forming the basis for a new identification key). Several previously named species were regarded as synonyms. The sites we sampled in Alaska were remarkable in the high levels of sympatry of Eurytemora species, to a degree not seen outside of Alaska. Future studies of Eurytemora should shed light on patterns of habitat invasions and physiological evolution within the genus, and yield insights into mechanisms leading to its remarkably broad geographic and habitat range.     

Phylogeny and evolution of the cryptic fungus‐farming ant genus Myrmicocrypta F. Smith (Hymenoptera: Formicidae) inferred from multilocus data

Fungus‐farming ants (Hymenoptera: Formicidae) have become model systems for exploring questions regarding the evolution of symbiosis. However, robust phylogenetic studies of both the ant agriculturalists and their fungal cultivars are necessary for addressing whether or not observed ant–fungus associations are the result of coevolution and, if so, whether that coevolution has been strict or diffuse. Here we focus on the evolutionary relationships of the species within the ant genus Myrmicocrypta and of their fungal cultivars. The fungus‐farming ant genus Myrmicocrypta was created by Fr. Smith in 1860 based on a single alate queen. Since then, 31 species and subspecies have been described. Until now, the genus has not received any taxonomic treatment and the relationships of the species within the genus have not been tested. Our molecular analyses, using ～40 putative species and six protein‐coding (nuclear and mitochondrial) gene fragments, recover Myrmicocrypta as monophyletic and as the sister group of the genus Mycocepurus Forel. The species M. tuberculata Weber is recovered as the sister to the rest of Myrmicocrypta. The time‐calibrated phylogeny recovers the age of stem group Myrmicocrypta plus its sister group as 45 Ma, whereas the inferred age for the crown group Myrmicocrypta is recovered as 27 Ma. Ancestral character‐state analyses suggest that the ancestor of Myrmicocrypta had scale‐like or squamate hairs and that, although such hairs were once considered diagnostic for the genus, the alternative state of erect simple hairs has evolved at least seven independent times. Ancestral‐state analyses of observed fungal cultivar associations suggest that the most recent common ancestor of Myrmicocrypta cultivated clade 2 fungal species and that switches to clade 1 fungi have occurred at least five times. It is our hope that these results will encourage additional species‐level phylogenies of fungus‐farming ants and their fungal cultivars, which are necessary for understanding the evolutionary processes that gave rise to agriculture in ants and that produced the current diversity of mutualistic ant–fungus interactions.     

Phylogenetic relationships of the pygmy rice rats of the genus Oligoryzomys Bangs, 1900 (Rodentia: Sigmodontinae)

Sequences from two mitochondrial genes (cytochrome b and NADH1) were used to produce a molecular phylogeny for 12 named and two undescribed species of the genus Oligoryzomys. All analyses placed Oligoryzomys microtis as the most basal taxon, a finding consistent with previous studies that suggested the west‐central Amazon as a centre of origin for the tribe Oryzomyini to which Oligoryzomys belongs. Biogeographically, this suggests that Oligoryzomys had a South American origin, and later advanced northwards, entering Central America and Mexico more recently. Different analyses have provided consistent support for several additional clades that did not necessarily agree with the species groups hypothesized by previous studies. A molecular clock derived for these data suggests an origin for the genus of 6.67 Mya, with most speciation within the genus occurring between 3.7 and 1.5 Mya. © 2010 The Linnean Society of London, Zoological Journal of the Linnean Society, 2010, 160 , 551–566.     

18S rDNA AND EVOLUTION IN THE DASYCLADALES (CHLOROPHYTA): MODERN LIVING FOSSILS1

Phylogenetic relationships were inferred from parsimony and distance analyses of nuclear small-subunit ribosomal DNA sequences taken from 14 species representing 8 of the 11 extant genera in the Dasycladales. Of 1733 aligned positions, 412 (23.8%) were variable and 251 (61%) of those were phylogenetically informative within the Dasycladales. Secondary structure was analyzed and taken into account during all phases of data analysis. Robustness of the trees was assessed using bootstrap analysis and g₁ statistics of tree-length decay. Strongly supported branches were robust to all methods of analysis regardless of weighting schemes used. The secondary structure of the 18S within the Dasycladales agrees with that of other green algae with the exception of a shared deletion in stemloop E10-1 (ca. 13 nucleotides long), which provides additional support for the uniqueness of this monophyletic group. A molecular clock was calibrated from the dasyclad fossil record and suggests a radiation of the Acetabulariaceae at 120 ± 30 million years (Ma) ago and the Dasycladaceae 215 ± 40 Ma ago. The split of the two lineages from a shared ancestor is estimated at 265 ± 50 Ma ago. Within the Dasycladaceae, Neomeris and Cymopolia are sister taxa, as are Batophora and Chlorocladus. Bornetella groups with the Neomeris and Cymopolia clade in 78% of the bootstrap replicates. Relationships among the Acetabulariaceae show that Acetabularia and Polyphysa do not form monophyletic groups as presently circumscribed. No evidence indicates that Acicularia is the oldest genus. Halicoryne, Chalmasia, and Dasycladus were not included in the analysis. Molecular data provide afresh background perspective from which to discuss the evolution of one of the most ancient lineages of green plants.     

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.     

Mitochondrial gene rearrangements suggest a new genus in the subfamily Cantharinae (Coleoptera)

The Cephalomalthinus semifumatus species group, referred to as the “semifumatus” group henceforth, is interesting because of its heterogeneous morphology resembling either Cephalomalthinus Pic, 1921 or Rhagonycha Eschscholtz, 1830. To elucidate its phylogenetic status, mitochondrial genomes of four species of the “semifumatus” group, 11 Cephalomalthinus species, and 11 Rhagonycha species were sequenced and examined. All analysed mitogenomes were similar with respect to genome size, nucleotide composition, and AT content. Surprisingly, a rearrangement of the trnW-trnC and trnY genes was detected in the “semifumatus” group, presumably caused by tandem duplication and random loss events. Furthermore, genetic distance analyses showed that the proximity of the “semifumatus” group to Cephalomalthinus and to Rhagonycha was comparable to that between the latter two. Moreover, the produced phylogeny strongly supported the monophyly of the “semifumatus” group, and molecular clock analyses dated its divergence from Cephalomalthinus to 32.52 Ma. Thus, the new genus Amphimorphus gen. nov. is suggested to comprise the “semifumatus” group, in which the observed gene rearrangement was a synapomorphy. Moreover, morphological evidence regarding the unique structure of the aedeagus supported this separation. These results indicate that mitochondrial gene rearrangement provide important phylogenetic implications for revising Cephalomalthinus, a speciose genus that is puzzling in the morphology-based taxonomy.     

Multi-locus phylogenetic analyses support the monophyly and the Miocene diversification of Iksookimia (Teleostei: Cypriniformes: Cobitidae)

The genus Iksookimia contains six species of primary freshwater fishes that are endemic to South Korea. Previous phylogenetic studies, based on DNA sequence data from three or fewer loci, have suggested non-monophyly of the genus, providing inconsistent resolutions of the relationships of Iksookimia. Our coalescent and concatenation-based phylogenetic analyses, utilizing seven unlinked nuclear-encoded genes, strongly supported Iksookimia as a monophyletic group, emphasizing the importance of multi-locus data in investigating complicated phylogenetic relationships. A relaxed molecular clock analysis using fossil calibrations, indicated that the origin of the major lineages of Iksookimia occurred between ～12 to 5 Ma, which is consistent with the Miocene uplift of the Taebaek and Sobaek Mountains and the Miocene activation of the major south-eastern faults. These palaeogeographic events may have served as vicariant events in the diversification of Iksookimia.     

Multiple nuclear genes stabilize the phylogenetic backbone of the genus Quercus

Phylogenetic relationships among 108 oak species (genus Quercus L.) were inferred using DNA sequences of six nuclear genes selected from the existing genomic resources of the genus. Previous phylogenetic reconstructions based on traditional molecular markers are inconclusive at the deeper nodes. Overall, weak phylogenetic signals were obtained for each individual gene analysis, but stronger signals were obtained when gene sequences were concatenated. Our data support the recognition of six major intrageneric groups Cyclobalanopsis, Cerris, Ilex, Quercus, Lobatae and Protobalanus. Our analyses provide resolution at deeper nodes but with moderate support and a more robust infrageneric classification within the two major clades, the ‘Old World Oaks’ (Cyclobalanopsis, Cerris, Ilex) and ‘New World Oaks’ (Quercus, Lobatae, Protobalanus). However, depending on outgroup choice, our analysis yielded two alternative placements of the Cyclobalanopsis clade within the genus Quercus. When Castanea Mill. was chosen as outgroup, our data suggested that the genus Quercus comprised two clades corresponding to two subgenera as traditionally recognized by Camus: subgenus Euquercus Hickel and Camus and subgenus Cyclobalanopsis Øersted (Schneider). However, when Notholithocarpus Manos, Cannon and S. Oh was chosen as an outgroup subgenus Cyclobalanopsis clustered with Cerris and Ilex groups to form the Old World clade. To assess the placement of the root, we complemented our dataset with published data of ITS and CRC sequences. Based on the concatenated eight gene sequences, the most likely root position is at the split between the ‘Old World Oaks’ and the ‘New World Oaks’, which is one of the alternative positions suggested by our six gene analysis. Using a dating approach, we inferred an Eocene age for the primary divergences in Quercus and a root age of about 50–55 Ma, which agrees with palaeobotanical evidence. Finally, irrespective of the outgroup choice, our data boost the topology within the New World clade, where (Protobalanus + Quercus) is a sister clade of Lobatae. Inferred divergence ages within this clade and the Cerris–Ilex clade are generally younger than could be expected from the fossil record, indicating that morphological differentiation pre-dates genetic isolation in this clade.     

Trapped in desert springs: phylogeography of Australian desert spring snails

Aim We investigate the phylogeographical history and determine the time‐scale of population divergence of hydrobiid freshwater snails (genus Trochidrobia) inhabiting groundwater springs in the Australian desert. We test the hypothesis that divergence between geographically distinct snail populations occurred simultaneously due to their isolation in hydrologically discrete spring systems, i.e. ‘trapped in desert springs’. Location Groundwater springs of the Great Artesian Basin (GAB) in central Australia. Methods DNA sequence data from the mitochondrial cytochrome c oxidase subunit I gene and the nuclear 28S and internal transcribed spacer rRNA genes were used to reconstruct phylogenetic relationships within and among three species of Trochidrobia (Hydrobiidae): T. punicea (13 spring groups, n = 90), T. smithi (12 spring groups, n = 62) and T. minuta (2 spring groups, n = 4). Bayesian relaxed molecular clock analyses and approximate Bayesian computation were used to date lineage divergence and distinguish between alternative biogeographical scenarios. Results The diversification of the three Trochidrobia species probably occurred between 2.54 and 9.3 Ma, prior to the formation of the springs c. 1 Ma. Intraspecific divergences within the two widespread species occurred after the formation and colonization of the springs. Coalescent modelling and molecular clock analyses supported a simultaneous radiation of five allopatric intraspecific snail lineages within T. punicea (two lineages) and T. smithi (three lineages) across the GAB springs examined. Main conclusions The analyses support the ‘trapped in desert springs’ hypothesis for the diversification of intraspecific lineages within the species T. punicea and T. smithi. This hypothesis suggests that the formation of deserts around Lake Eyre in the early Pleistocene led to the hydrological isolation of spring complexes in the GAB, resulting in significant molecular divergence, but no morphological divergence, of Trochidrobia snail populations.     

Human evolution at the Matuyama‐Brunhes boundary

The cranial morphology of fossil hominids between the end of the Early Pleistocene and the beginning of the Middle Pleistocene provides crucial evidence to understand the distribution in time and space of the genus Homo. This evidence is critical for evaluating the competing models regarding diversity within our genus. The debate focuses on two alternative hypotheses, one basically anagenetic and the other cladogenetic. The first suggests that morphological change is so diffused, slow, and steady that it is meaningless to apply species names to segments of a single lineage. The second is that the morphological variation observed in the fossil record can best be described as a number of distinct species that are not connected in a linear ancestor‐descendant sequence. Today much more fossil evidence is available than was in the past to test these alternative hypotheses, as well as intermediate variants. Special attention must be paid to Africa because this is the most probable continental homeland for both the origin of the genus Homo (around 2.5–2 Ma), 1 as well as the site, two million or so years later, of the emergence of the species H. sapiens. 2 However, the African fossil record is very poorly represented between 1 Ma and 600 ka. Europe furnishes recent discoveries in this time range around the Matuyama‐Brunhes chron boundary (780,000 years ago), a period for which, at present, we have no noteworthy fossil evidence in Africa or the Levant. Two penecontemporaneous sources of European fossil evidence, the Ceprano calvaria (Italy) 3 and the TD6 fossil assemblage of Atapuerca (Spain) 4 are thus of great interest for testing hypotheses about human evolution in the fundamental time span bracketed between the late Early and the Middle Pleistocene. This paper is based on a phenetic approach to cranial variation aimed at reviewing the Early‐to‐Middle Pleistocene trajectories of human evolution. The focus of the paper is on neither the origin nor the end of the story of the genus Homo, but rather its chronological and phylogenetic core. Elucidation of the evolutionary events that happened around 780 ka during the transition from the Early to Middle Pleistocene is one of the new frontiers for human paleontology, and is critical for understanding the processes that ultimately led to the origin of H. sapiens.     

Phylogeography of the Namib Desert dung beetles Scarabaeus (Pachysoma) MacLeay (Coleoptera: Scarabaeidae)

Aim Namib biogeography in many instances remains reliant on advanced and detailed systematic studies. This study attempts to combine molecular phylogenetic data, geology and palaeo‐climatic data to (i) resolve the relationships of the 13 morphological species of Scarabaeus (Pachysoma) and (ii) relate their evolution to past climatic and geological events. Location South Africa and Namibia. Methods Sequencing of a 1197 bp segment of the mitochondrial cytochrome oxidase I (COI) gene of the 13 species within Scarabaeus (Pachysoma) was undertaken. Analyses performed included Parsimony and Maximum Likelihood as well as imposing a molecular clock. Results The molecular phylogeny showed strong support for 11 of the 13 morphological species. The remaining two species, S. (P.) glentoni and S. (P.) hippocrates, formed a complex and could not be assigned specific status on the basis of the COI gene phylogeny. Strong support for the three species formerly classified within the genus Neopachysoma was consistently obtained. The subgenus appears to have arisen c. 2.9 Ma. Species within the subgenus arose at different times, with the common ancestor to Neopachysoma and the hippocrates complex having evolved 2.65 and 2.4 Ma, respectively. Scarabaeus (P.) denticollis, S. (P.) rotundigenus, S. (P.) rodriguesi and S. (P.) schinzi are some of the youngest species, having diverged between 2 million and 600,000 years ago. Main conclusions Scarabaeus (Pachysoma) is a derived monophyletic clade within the Scarabaeini. The subgenus appears to be young in comparison with the age of the Namib Desert, which dates back to the Miocene (c. 15 Ma). The psammophilous taxa are shown to disperse with their substratum and habitat, barchan dunes. Clear south/north evolutionary gradients can be seen within the species of this subgenus, which are consistent with the unidirectional wind regime. Species with a suite of mostly plesiomorphic characters have a southerly distribution while their derived psammophilous relatives have central to northern Namib distributions. Major rivers such as the Orange, Buffels and Holgat appear to be gene barriers to certain species as well as areas of origin of speciation events.