The role of pollinator diversity in the evolution of corolla-shape integration in a pollination-generalist plant clade

Flowers of animal-pollinated plants are integrated structures shaped by the action of pollinator-mediated selection. It is widely assumed that pollination specialization increases the magnitude of floral integration. However, empirical evidence is still inconclusive. In this study, we explored the role of pollinator diversity in shaping the evolution of corolla-shape integration in Erysimum, a plant genus with generalized pollination systems. We quantified floral integration in Erysimum using geometric morphometrics and explored its evolution using phylogenetic comparative methods. Corolla-shape integration was low but significantly different from zero in all study species. Spatial autocorrelation and phylogenetic signal in corolla-shape integration were not detected. In addition, integration in Erysimum seems to have evolved in a way that is consistent with Brownian motion, but with frequent convergent evolution. Corolla-shape integration was negatively associated with the number of pollinators visiting the flowers of each Erysimum species. That is, it was lower in those species having a more generalized pollination system. This negative association may occur because the co-occurrence of many pollinators imposes conflicting selection and cancels out any consistent selection on specific floral traits, preventing the evolution of highly integrated flowers.     

Why Africa matters: evolution of Old World Salvia (Lamiaceae) in Africa

Background and Aims

Salvia is the largest genus in Lamiaceae and it has recently been found to be non-monophyletic. Molecular data on Old World Salvia are largely lacking. In this study, we present data concerning Salvia in Africa. The focus is on the colonization of the continent, character evolution and the switch of pollination systems in the genus.

Methods

Maximum likelihood and Bayesian inference were used for phylogenetic reconstruction. Analyses were based on two nuclear markers [internal transcribed spacer (ITS) and external transcribed spacer (ETS)] and one plastid marker (rpl32-trnL). Sequence data were generated for 41 of the 62 African taxa (66 %). Mesquite was used to reconstruct ancestral character states for distribution, life form, calyx shape, stamen type and pollination syndrome.

Key Results

Salvia in Africa is non-monophyletic. Each of the five major regions in Africa, except Madagascar, was colonized at least twice, and floristic links between North African, south-west Asian and European species are strongly supported. The large radiation in Sub-Saharan Africa (23 species) can be traced back to dispersal from North Africa via East Africa to the Cape Region. Adaptation to bird pollination in southern Africa and Madagascar reflects parallel evolution.

Conclusions

The phenotypic diversity in African Salvia is associated with repeated introductions to the continent. Many important evolutionary processes, such as colonization, adaptation, parallelism and character transformation, are reflected in this comparatively small group. The data presented in this study can help to understand the evolution of Salvia sensu lato and other large genera.     

Cercopithecidae from the Pliocene Chiwondo beds, Malawi-rift

There are a total of 16 fossil cercopithecid specimens, representing at least 10 individuals, from the Chiwondo Beds of northern Malawi. The majority of this material is derived from the Middle Pliocene Unit 3A, but one specimen is from the Early Pliocene Unit 2. This latter specimen is from a papionin of indeterminate genus similar in size to Parapapio ado and Pliopapio alemui. Among the specimens from Unit 3A, two species can be diagnosed: an indeterminate species of Theropithecus, and a species of Parapapio similar in dental size to P. broomi. Neither of the genera from Unit 3A are exclusive to either East Africa or South Africa. Their relative abundances, however, are more similar to Middle Pliocene South African sites where Parapapio is the most common primate genus, and Theropithecus is comparatively rare. This is in contrast to similarly-aged East African sites where Theropithecus is by far the most abundant genus.     

The pace of morphological change: historical transformation of skull shape in St Bernard dogs

Owing to the great morphological diversity of domestic dogs (Canis familiaris), the study of historical shape change in dog skulls provides an excellent opportunity for investigating the dynamics of morphological evolution. Breed standards make known which features were selected by breeders. Here we use the methods of geometric morphometrics to study change of skull shape in a series of purebred St Bernard dogs spanning nearly 120 years. A regression of shape on time was highly significant and revealed a consistent trend of shape change that corresponded to the features deemed desirable by the breed standard. Historical shape change in St Bernards involves a broadening of the skull and a tilting of the palate and upper jaw relative to the rest of the skull. This trend appears to be linear throughout the entire period and appears to be continuing. Allometry was ruled out as a contributing factor to this change because there was no consistent trend of historical change in skull size and because neither the patterns of static nor ontogenetic allometry corresponded to the historical shape change. The dramatic modification of the St Bernard skull demonstrates that selection can achieve sustained and substantial change and can completely overcome constraints such as allometry.     

Cytotype diversity and genome size variation in eastern Asian polyploid Cardamine (Brassicaceae) species

Background and Aims

Intraspecific ploidy-level variation is an important aspect of a species'' genetic make-up, which may lend insight into its evolutionary history and future potential. The present study explores this phenomenon in a group of eastern Asian Cardamine species.

Methods

Plant material was sampled from 59 localities in Japan and Korea, which were used in karyological (chromosome counting) and flow cytometric analyses. The absolute nuclear DNA content (in pg) was measured using propidium iodide and the relative nuclear DNA content (in arbitrary units) was measured using 4,6-diamidino-2-phenylindole fluorochrome.

Key Results

Substantial cytotype diversity was found, with strikingly different distribution patterns between the species. Two cytotypes were found in C. torrentis sensu lato (4x and 8x, in C. valida and C. torrentis sensu stricto, respectively), which displays a north–south geographical pattern in Japan. Hypotheses regarding their origin and colonization history in the Japanese archipelago are discussed. In Korean C. amaraeiformis, only tetraploids were found, and these populations may in fact belong to C. valida. C. yezoensis was found to harbour as many as six cytotypes in Japan, ranging from hexa- to dodecaploids. Ploidy levels do not show any obvious geographical pattern; populations with mixed ploidy levels, containing two to four cytotypes, are frequently observed throughout the range. C. schinziana, an endemic of Hokkaido, has hexa- and octoploid populations. Previous chromosome records are also revised, showing that they are largely based on misidentified material or misinterpreted names.

Conclusions

Sampling of multiple populations and utilization of the efficient flow cytometric approach allowed the detection of large-scale variation in ploidy levels and genome size variation attributable to aneuploidy. These data will be essential in further phylogenetic and evolutionary studies.     

The making of a pest: the evolution of a fruit-penetrating ovipositor in Drosophila suzukii and related species

Evolutionary innovation can allow a species access to a new ecological niche, potentially reducing competition with closely related species. While the vast majority of Drosophila flies feed on rotting fruit and other decaying matter, and are harmless to human activity, Drosophila suzukii, which has a morphologically modified ovipositor, is capable of colonizing live fruit that is still in the process of ripening, causing massive agricultural damage. Here, we conducted the first comparative analysis of this species and its close relatives, analysing both ovipositor structure and fruit susceptibility. We found that the ovipositor of the species most closely related to D. suzukii, Drosophila subpulchrella, has a similar number of enlarged, evolutionarily derived bristles, but a notably different overall shape. Like D. suzukii, D. subpulchrella flies are capable of puncturing the skin of raspberries and cherries, but we found no evidence that they could penetrate the thicker skin of two varieties of grapes. More distantly related species, one of which has previously been mistaken for D. suzukii, have blunt ovipositors with small bristles. While they did not penetrate fruit skin in any of the assays, they readily colonized fruit interiors where the skin was broken. Our results suggest that considering evolutionary context may be beneficial to the management of invasive species.     

Postnatal temporal bone ontogeny in Pan,Gorilla, and Homo,and the implications for temporal bone ontogeny in Australopithecus afarensis

Assessments of temporal bone morphology have played an important role in taxonomic and phylogenetic evaluations of fossil taxa, and recent three‐dimensional analyses of this region have supported the utility of the temporal bone for testing taxonomic and phylogenetic hypotheses. But while clinical analyses have examined aspects of temporal bone ontogeny in humans, the ontogeny of the temporal bone in non‐human taxa is less well documented. This study examines ontogenetic allometry of the temporal bone in order to address several research questions related to the pattern and trajectory of temporal bone shape change during ontogeny in the African apes and humans. We further apply these data to a preliminary analysis of temporal bone ontogeny in Australopithecus afarensis. Three‐dimensional landmarks were digitized on an ontogenetic series of specimens of Homo sapiens, Pan troglodytes, Pan paniscus, and Gorilla gorilla. Data were analyzed using geometric morphometric methods, and shape changes throughout ontogeny in relation to size were compared. Results of these analyses indicate that, despite broadly similar patterns, African apes and humans show marked differences in development of the mandibular fossa and tympanic portions of the temporal bone. These findings indicate divergent, rather than parallel, postnatal ontogenetic allometric trajectories for temporal bone shape in these taxa. The pattern of temporal bone shape change with size exhibited by A. afarensis showed some affinities to that of humans, but was most similar to extant African apes, particularly Gorilla. Am J Phys Anthropol 151:630–642, 2013. © 2013 Wiley Periodicals, Inc.     

A new Middle Miocene tarsier from Thailand and the reconstruction of its orbital morphology using a geometric-morphometric method

Tarsius is an extant genus of primates endemic to the islands of Southeast Asia that is characterized by enormously enlarged orbits reflecting its nocturnal activity pattern. Tarsiers play a pivotal role in reconstructing primate phylogeny, because they appear to comprise, along with Anthropoidea, one of only two extant haplorhine clades. Their fossils are extremely rare. Here, we describe a new species of Tarsius from the Middle Miocene of Thailand. We reconstructed aspects of its orbital morphology using a geometric-morphometric method. The result shows that the new species of Tarsius had a very large orbit (falling within the range of variation of modern Tarsius) with a high degree of frontation and a low degree of convergence. Its relatively divergent lower premolar roots suggest a longer mesial tooth row and therefore a longer muzzle than in extant species. The new species documents a previous unknown Miocene group of Tarsius, indicating greater taxonomic diversity and morphological complexity during tarsier evolution. The current restriction of tarsiers to offshore islands in Southeast Asia appears to be a relatively recent phenomenon.     

Evolution of body shape in sympatric versus non-sympatric Tropheus populations of Lake Tanganyika

Allopatric speciation often yields ecologically equivalent sister species, so that their secondary admixis enforces competition. The shores of Lake Tanganyika harbor about 120 distinct populations of the cichlid genus Tropheus, but only some are sympatric. When alone, Tropheus occupies a relatively broad depth zone, but in sympatry, fish segregate by depth. To assess the effects of competition, we studied the partial co-occurrence of Tropheus moorii ‘Kaiser'' and ‘Kirschfleck'' with Tropheus polli. A previous study demonstrated via standardized breeding experiments that some observed differences between Tropheus ‘Kaiser'' living alone and in sympatry with T. polli have a genetic basis despite large-scale phenotypic plasticity. Using geometric morphometrics and neutral genetic markers, we now investigated whether sympatric populations differ consistently in body shape from populations living alone and if the differences are adaptive. We found significant differences in mean shape between non-sympatric and sympatric populations, whereas all sympatric populations of both color morphs clustered together in shape space. Sympatric populations had a relatively smaller head, smaller eyes and a more anterior insertion of the pectoral fin than non-sympatric populations. Genetically, however, non-sympatric and sympatric ‘Kaiser'' populations clustered together to the exclusion of ‘Kirschfleck''. Genetic distances, but not morphological distances, were correlated with geographic distances. Within- and between-population covariance matrices for T. moorii populations deviated from proportionality. It is thus likely that natural selection acts on both phenotypic plasticity and heritable traits and that both factors contribute to the observed shape differences. The consistency of the pattern in five populations suggests ecological character displacement.     

A PCR system for detection of species and genotypes of the Echinococcus granulosus-complex, with reference to the epidemiological situation in eastern Africa

We describe the development of a specific and sensitive PCR/semi-nested PCR system for the rapid diagnosis of Echinococcus granulosus genotype G1, E. granulosus genotype G6/7, and Echinococcus ortleppi (G5). Diagnosis of G1 and the group G5/6/7 is performed by a simple PCR, while discrimination between E. ortleppi (G5) and G6/7 involves a subsequent semi-nested PCR step. The target sequence for amplification is part of the mitochondrial 12S rRNA gene. Specificity of the PCRs was 100% when evaluated with isolates of 16 species of cestodes, including Echinococcus multilocularis, Echinococcus equinus, E. ortleppi and three strains of E. granulosus (G1, G6 and G7). Sensitivity threshold was 0.25pg of DNA. This new approach was compared with published protocols of restriction fragment length polymorphism-PCR and sequencing of mitochondrial cytochrome c oxidase subunit 1 and NADH dehydrogenase 1 genes using Echinococcus isolates of human, sheep, goat, camel, cattle and pig origin from Kenya and Sudan. Additionally, two internal DNA probes were developed, one hybridising only with G1, the other with G5, G6 and G7 amplification products. Preliminary epidemiological results obtained with this PCR approach include the detection of a camel strain (G6) infection for the first time in a human patient from eastern Africa, and the first reports of E. ortleppi (G5) in livestock from Kenya and the Sudan.     

A review of the alderfly genus Leptosialis Esben-Petersen (Megaloptera, Sialidae) with description of a new species from South Africa

The monotypic South African alderfly genus Leptosialis Esben-Petersen, 1920 is reviewed and Leptosialis africana Esben-Petersen, 1920 is redescribed. In the process a new species of alderfly Leptosialis necopinatasp. n. from the Eastern Cape and KwaZulu-Natal provinces of South Africa is recognised and described. Within Sialidae the new species most closely resembles Leptosialis africana. A key to the two species of Leptosialis using both adult and larval characters is provided.     

Life history,sexual dimorphism and ‘ornamental’ feathers in the mesozoic bird Confuciusornis sanctus

The life history of Confuciusornis sanctus is controversial. Recently, the species’ body size spectrum was claimed to contradict osteohistological evidence for a rapid, bird-like development. Moreover, sexual size dimorphism was rejected as an explanation for the observed bimodal size distribution since the presence of elongated rectrices, an assumed ‘male’ trait, was uncorrelated with size. However, this interpretation (i) fails to explain the size spectrum of C. sanctus which is trimodal rather than bimodal, (ii) requires implausible neonate masses and (iii) is not supported by analogy with sexual dimorphisms in modern birds, in which elongated central rectrices are mostly sex-independent. Available information on C. sanctus is readily reconciled if we assume a bird-like life history, as well as a pronounced sexual size dimorphism and sexually isomorphic extravagant feathers as frequently observed in extant species.     

Massive expansions of Dscam splicing diversity via staggered homologous recombination during arthropod evolution

The arthropod Down syndrome cell adhesion molecule (Dscam) gene can generate tens of thousands of protein isoforms via combinatorial splicing of numerous alternative exons encoding immunoglobulin variable domains organized into three clusters referred to as the exon 4, 6, and 9 clusters. Dscam protein diversity is important for nervous system development and immune functions. We have performed extensive phylogenetic analyses of Dscam from 20 arthropods (each containing between 46 and 96 alternative exons) to reconstruct the detailed history of exon duplication and loss events that built this remarkable system over 450 million years of evolution. Whereas the structure of the exon 4 cluster is ancient, the exon 6 and 9 clusters have undergone massive, independent expansions in each insect lineage. An analysis of nearly 2000 duplicated exons enabled detailed reconstruction of the timing, location, and boundaries of these duplication events. These data clearly show that new Dscam exons have arisen continuously throughout arthropod evolution and that this process is still occurring in the exon 6 and 9 clusters. Recently duplicated regions display boundaries corresponding to a single exon and the adjacent intron. The boundaries, homology, location, clustering, and relative frequencies of these duplication events strongly suggest that staggered homologous recombination is the major mechanism by which new Dscam exons evolve. These data provide a remarkably detailed picture of how complex gene structure evolves and reveal the molecular mechanism behind this process.     

Discovery of an unknown diversity of Leucinodes species damaging Solanaceae fruits in sub-Saharan Africa and moving in trade (Insecta,Lepidoptera, Pyraloidea)

The larvae of the Old World genera Leucinodes Guenée, 1854 and Sceliodes Guenée, 1854 are internal feeders in the fruits of Solanaceae, causing economic damage to cultivated plants like Solanum melongena and Solanum aethiopicum. In sub-Saharan Africa five nominal species of Leucinodes and one of Sceliodes occur. One of these species, the eggplant fruit and shoot borer Leucinodes orbonalis Guenée, 1854, is regarded as regularly intercepted from Africa and Asia in Europe, North and South America and is therefore a quarantine pest on these continents. We investigate the taxonomy of African Leucinodes and Sceliodes based on morphological characters in wing pattern, genitalia and larvae, as well as mitochondrial DNA, providing these data for identification of all life stages. The results suggest that both genera are congeneric, with Sceliodes syn. n. established as junior subjective synonym of Leucinodes. Leucinodes orbonalis is described from Asia and none of the samples investigated from Africa belong to this species. Instead, sub-Saharan Africa harbours a complex of eight endemic Leucinodes species. Among the former nominal species of Leucinodes (and Sceliodes) from Africa, only Leucinodes laisalis (Walker, 1859), comb. n. (Sceliodes) is confirmed, with Leucinodes translucidalis Gaede, 1917, syn. n. as a junior subjective synonym. The other African Leucinodes species were unknown to science and are described as new: Leucinodes africensis sp. n., Leucinodes ethiopica sp. n., Leucinodes kenyensis sp. n., Leucinodes malawiensis sp. n., Leucinodes pseudorbonalis sp. n., Leucinodes rimavallis sp. n. and Leucinodes ugandensis sp. n. An identification key based on male genitalia is provided for the African Leucinodes species. Most imports of Leucinodes specimens from Africa into Europe refer to Leucinodes africensis, which has been frequently imported with fruits during the last 50 years. In contrast, Leucinodes laisalis has been much less frequently recorded, and Leucinodes pseudorbonalis as well as Leucinodes rimavallis only very recently in fruit imports from Uganda. Accordingly, interceptions of Leucinodes from Africa into other continents will need to be re-investigated for their species identity and will likely require, at least in parts, revisions of the quarantine regulations. The following African taxa are excluded from Leucinodes: Hyperanalyta Strand, 1918, syn. rev. as revised synonym of Analyta Lederer, 1863; Analyta apicalis (Hampson, 1896), comb. n. (Leucinodes); Lygropia aureomarginalis (Gaede, 1916), comb. n. (Leucinodes); Syllepte hemichionalis Mabille, 1900, comb. rev., Syllepte hemichionalis idalis Viette, 1958, comb. rev. and Syllepte vagans (Tutt, 1890), comb. n. (Aphytoceros). Deanolis iriocapna (Meyrick, 1938), comb. n. from Indonesia is originally described and misplaced in Sceliodes, and Leucinodes cordalis (Doubleday, 1843), comb. n. (Margaritia) from New Zealand, Leucinodes raondry (Viette, 1981), comb. n. (Daraba) from Madagascar as well as Leucinodes grisealis (Kenrick, 1912), comb. n. (Sceliodes) from New Guinea are transferred from Sceliodes to Leucinodes. While Leucinodes is now revised from Africa, it still needs further revision in Asia.     

Trap diversity and evolution in the family Droseraceae

We review trapping mechanisms in the carnivorous flowering plant family Droseraceae (order Caryophyllales). Its members are generally known to attract, capture, retain and digest prey animals (mainly arthropods) with active snap-traps (Aldrovanda, Dionaea) or with active sticky flypaper traps (Drosera) and to absorb the resulting nutrients. Recent investigations revealed how the snap-traps of Aldrovanda vesiculosa (waterwheel plant) and Dionaea muscipula (Venus’ flytrap) work mechanically and how these apparently similar devices differ as to their functional morphology and shutting mechanics. The Sundews (Drosera spp.) are generally known to possess leaves covered with glue-tentacles that both can bend toward and around stuck prey. Recently, it was shown that there exists in this genus a higher diversity of different tentacle types and trap configurations than previously known which presumably reflect adaptations to different prey spectra. Based on these recent findings, we finally comment on possible ways for intrafamiliar trap evolution.     

Nucleotide diversity and molecular evolution of the WAG-2 gene in common wheat (Triticum aestivum L) and its relatives

In this work, we examined the genetic diversity and evolution of the WAG-2 gene based on new WAG-2 alleles isolated from wheat and its relatives. Only single nucleotide polymorphisms (SNP) and no insertions and deletions (indels) were found in exon sequences of WAG-2 from different species. More SNPs and indels occurred in introns than in exons. For exons, exons+introns and introns, the nucleotide polymorphism π decreased from diploid and tetraploid genotypes to hexaploid genotypes. This finding indicated that the diversity of WAG-2 in diploids was greater than in hexaploids because of the strong selection pressure on the latter. All dn/ds ratios were < 1.0, indicating that WAG-2 belongs to a conserved gene affected by negative selection. Thirty-nine of the 57 particular SNPs and eight of the 10 indels were detected in diploid species. The degree of divergence in intron length among WAG-2 clones and phylogenetic tree topology suggested the existence of three homoeologs in the A, B or D genome of common wheat. Wheat AG-like genes were divided into WAG-1 and WAG-2 clades. The latter clade contained WAG-2, OsMADS3 and ZMM2 genes, indicating functional homoeology among them.     

Past climate changes explain the phylogeography of Vitellaria paradoxa over Africa

The evolution of the savanna biome has been deeply marked by repeated contraction/expansion phases due to climate perturbations during the Quaternary period. In this study, we investigated the impact of the last glacial maximum (LGM) on the present genetic pattern of Vitellaria paradoxa (shea tree), a major African savanna tree. A range-wide sampling of the species enabled us to sample 374 individuals from 71 populations distributed throughout sub-Sahelian Africa. Trees were genotyped using 3 chloroplasts and 12 nuclear microsatellites, and were sequenced for 2 polymorphic chloroplast intergenic spacers. Analyses of genetic diversity and structure were based on frequency-based and Bayesian methods. Potential distributions of V. paradoxa at present, during the LGM and the last interglacial period, were examined using DIVA-GIS ecological niche modelling (ENM). Haplotypic and allelic richness varied significantly across the range according to chloroplast and nuclear microsatellites, which pointed to higher diversity in West Africa. A high but contrasted level of differentiation was revealed among populations with a clear phylogeographic signal, with both nuclear (F(ST) = 0.21; R(ST) = 0.28; R(ST) > R(ST) (permuted)) and chloroplast simple sequence repeats (SSRs) (G(ST) = 0.81; N(ST) = 0.90; N(ST) > N(ST) (permuted)). We identified a strong geographically related structure separating western and eastern populations, and a substructure in the eastern part of the area consistent with subspecies distinction. Using ENM, we deduced that perturbations during the LGM fragmented the potential eastern distribution of shea tree, but not its distribution in West Africa. Our main results suggest that climate variations are the major factor explaining the genetic pattern of V. paradoxa.     

Morphology and gonad development of normal soldiers and reproductive soldiers of the termite Zootermopsis nevadensis nevadensis (Isoptera, Archotermopsidae)

Reproductive or neotenic soldiers of the Archotermopsid Zootermopsis nevadensisnevadensis (Hagen) are compared to sterile soldiers and primary male reproductives. Several head capsule morphometrics correlate significantly with gonad size across all forms and both sexes of soldiers. The easily observed field character of ratio of mandible length to labrum length is a consistent and reliable feature of head capsule external morphology for predicting gonad development and reproductive potential of soldier forms regardless of age, sex, or live weight.