Steph B. J. Menken

Sex pheromones and their potential role in the evolution of reproductive isolation in small ermine moths (Yponomeutidae)

SummarySex pheromone communication in the nine European species of small ermine moths (Yponomeuta) is reviewed in regard to the potential role of pheromones in the speciation process. Six of the nine species studied (viz.,Y. evonymellus, Y. cagnagellus, Y. padellus, Y. irrorellus, Y. plumbellus, andY. vigintipunctatus) use a mixture of (E)-11-and (Z)-11-tetradecenyl acetate in different ratios as primary pheromone components, with combinations of tetradecyl acetate, (Z)-9-tetradecenyl acetate, (Z)-11-hexadecenyl acetate and the corresponding alcohols of the acetates as additional pheromone components. Analysis of (Z)- to (E)-11-tetradecenyl acetate ratios produced by individual females of these species demonstrated significant variation among females of all species. However, the ranges of ratios produced byY. cagnagellus, Y. irrorellus, andY. plumbellus, sharing the same host-plant species, spindle tree, did not overlap. Niche separation of all six species mentioned required consideration of at least one additional pheromone component or of temporal aspects. The remaining three species,i.e. Y. malinellus, Y. mahalebellus andY. rorellus, have pheromones that differ qualitatively.Biosynthetic routes to the pheromone components identified are proposed on the basis of fatty acid pheromone precursors found in the pheromone glands. A phylogenetic tree for the genus is constructed based on allozyme frequency data and changes in pheromone composition are superimposed on this tree. We suggest that the ancestral ermine moth pheromone is a mixture of (Z)-11- and (E)-11-tetradecenyl acetate and the corresponding alcohols, and a scenario of how present-day patterns evolved is outlined. The pheromone differences among the three species using spindle tree as their host-plant might have evolved throughreproductive character displacement upon secondary contact between populations that had already diverged genetically in allopatry. Pheromone differences within the so-calledpadellus-complex (includingY. cagnagellus, Y. mahalebellus, Y. malinellus, Y. padellus, andY. rorellus) in which species might have originated sympatrically, may have evolved byreinforcing selection as these species still hybridise and produce viable offspring when confined in cages. The role of pheromones in reproductive isolation amongYponomeuta species is emphasised by (1) the function of pheromone components of some of the species as behavioural antagonists to other species, (2) the cross-attraction under experimental conditions between allochronic species with similar pheromones, and (3) the formation of hybrids in the laboratory between species that are isolated in nature by pheromone differences.

Genetical population structure in plants: Gene flow between diploid sexual and triploid asexual dandelions (Taraxacum section Ruderalia)

Levels and distribution of genetic variation were studied in central and western European populations of Taraxacum section Ruderalia containing differing mixtures of sexual diploid and asexual triploid plants. All sexual populations were panmictic with their variation partitioned mainly among populations. Genotypic diversity in triploid samples was very high with few clones widespread and many clones restricted to one or a few populations. Extensive amounts of gene (pollen) flow between the diploid and triploid components of a population were inferred from the following data: (1) the two ploidy levels share all major allozyme polymorphisms; (2) the intrapopulational homogeneity in genic variation between diploids and triploids contrasts strongly with the geographic differentiation at each ploidy level separately; (3) population‐unique alleles simultaneously occur at the two ploidy levels; (4) not only sexuals but also asexuals generally simulate Hardy‐Weinberg expectations. Most likely, intrapopulational gene exchange occurs bidirectionally by mechanisms such as reductional pollen meiosis in apomictic plants, facultative apomixis, and formation of unreduced gametes in sexuals. Thus, diploid and triploid Taraxacum section Ruderalia are less genetically isolated than has previously been supposed and probably form a cohesive evolutionary unit with the level at which gene pools are shared differing by population.

Host race formation in Tetranychus urticae: genetic differentiation, host plant preference, and mate choice in a tomato and a cucumber strain

The two‐spotted spider mite, Tetranychus urticae Koch, occurs in two colour forms in greenhouses in the Netherlands: a red form on tomato and a green form on cucumber. The evolutionary status of these strains was analysed by studying genetic differentiation, host plant preference, and mate choice. Males of the tomato strain preferred the female (30 h‐old) teleiochrysales from the same strain to those of the cucumber strain, independent of the host plants (tomato, cucumber, bean) on which the teleiochrysales were placed. In contrast, males of the cucumber strain were not selective. In a Y‐tube olfactometer, females of the cucumber strain were not responsive to host plant volatiles alone. However, in two‐choice disc experiments, where females were exposed to both volatile and contact cues, they settled on cucumber leaves in preference to tomato leaves. Females of the tomato strain preferred the odour of tomato leaves and settled on tomato leaves in preference to cucumber leaves. These experimental results provide the first evidence for (1) host‐plant independent mate selection in male spider mites and (2) olfactory discrimination between host plants in female spider mites.

BIRD SPECIES AND TRAITS ASSOCIATED WITH LOGGED AND UNLOGGED FOREST IN BORNEO

The ecological consequences of logging have been and remain a focus of considerable debate. In this study, we assessed bird species composition within a logging concession in Central Kalimantan, Indonesian Borneo. Within the study area (approximately 196 km2) a total of 9747 individuals of 177 bird species were recorded. Our goal was to identify associations between species traits and environmental variables. This can help us to understand the causes of disturbance and predict whether species with given traits will persist under changing environmental conditions. Logging, slope position, and a number of habitat structure variables including canopy cover and liana abundance were significantly related to variation in bird composition. In addition to environmental variables, spatial variables also explained a significant amount of variation. However, environmental variables, particularly in relation to logging, were of greater importance in structuring variation in composition. Environmental change following logging appeared to have a pronounced effect on the feeding guild and size class structure but there was little evidence of an effect on restricted range or threatened species although certain threatened species were adversely affected. For example, species such as the terrestrial insectivore Argusianus argus and the hornbill Buceros rhinoceros, both of which are threatened, were rare or absent in recently logged forest. In contrast, undergrowth insectivores such as Orthotomus atrogularis and Trichastoma rostratum were abundant in recently logged forest and rare in unlogged forest. Logging appeared to have the strongest negative effect on hornbills, terrestrial insectivores, and canopy bark-gleaning insectivores while moderately affecting canopy foliage-gleaning insectivores and frugivores, raptors, and large species in general. In contrast, undergrowth insectivores responded positively to logging while most understory guilds showed little pronounced effect. Despite the high species richness of logged forest, logging may still have a negative impact on extant diversity by adversely affecting key ecological guilds. The sensitivity of hornbills in particular to logging disturbance may be expected to alter rainforest dynamics by seriously reducing the effective seed dispersal of associated tree species. However, logged forest represents an increasingly important habitat for most bird species and needs to be protected from further degradation. Biodiversity management within logging concessions should focus on maintaining large areas of unlogged forest and mitigating the adverse effects of logging on sensitive groups of species.

Multiple paternity and female-biased mutation at a microsatellite locus in the olive ridley sea turtle (Lepidochelys olivacea)

Multiple paternity in the olive ridley sea turtle (Lepidochelys olivacea) population nesting in Suriname was demonstrated using two microsatellite loci, viz., Ei8 and Cm84. The large number of offspring sampled per clutch (70 on average, ranging from 15 to 103) and the number of alleles found at the two loci (18 and eight alleles, respectively) enabled unambiguous assessment of the occurrence of multiple paternity. In two out of 10 clutches analysed, the offspring had been sired by at least two males, which was confirmed at both loci. In both clutches, unequal paternity occurred: 73% and 92% of the offspring had been sired by the primary male. The probability of detecting multiple paternity was 0.903, and therefore there is a small chance that multiple paternity occurred but remained undetected in some of the eight clutches that appeared to be singly sired. Analysis of 703 offspring revealed a high mutation rate for locus Ei8 (μ = 2.3 × 10−2) with all 33 mutations occurring in maternal alleles. In particular, one allele of 274 bp mutated at a high frequency in a clutch to which the mother contributed the allele, but in another clutch where the father contributed the same allele, no such mutations were observed. Inferred allele-specific mutation rates for Ei8 and expected numbers of mutations per clutch confirmed that maternal alleles for Ei8 are more likely to mutate in the olive ridley sea turtle than paternal alleles. Possible explanations are discussed.

Phylogeography of the planktonic chaetognath Sagitta setosa reveals isolation in European seas.

Abstract Numerous planktonic species have disjunct distribution patterns in the worlds oceans. However, it is unclear whether these are truly unconnected by gene flow, or whether they are composed of morphologically cryptic species. The marine planktonic chaetognath Sagitta setosa Müller has a discontinuous geographic distribution over the continental shelf in the northeastern Atlantic, Mediterranean Sea, and Black Sea. Morphological variation between these populations has been described, but overlaps and is therefore unsuitable to determine the degree of isolation between populations. To test whether disjunct populations are also genetically disjunct, we sequenced a 504‐bp fragment of mitochondrial DNA comprising the cytochrome oxidase II region of 86 individuals. Sequences were highly variable; each represented a different haplotype. Within S. setosa, sequence divergence ranged from 0.2 to 8.1% and strong phylogeographic structure was found, with four main groups corresponding to the northeastern Atlantic, Mediterranean Sea (including Ligurian Sea, Tyrrhenian Sea and Gulf of Gabès), Adriatic Sea, and Black Sea. Two of these (Atlantic and Black Sea) were resolved as monophyletic clades, thus gene flow between disjunct populations of S. setosa has been extremely limited and lineage sorting has taken place. The deepest divergence was between Atlantic and Mediterranean/Black Sea populations followed by a split between Mediterranean and Black Sea populations. The Mediterranean/Black Sea clade comprised three groups, with the Adriatic Sea as the most likely sister clade of the Black Sea. These data are consistent with a colonization of the Black Sea from the Mediterranean. Furthermore, a possible cryptic species was found in the Black Sea with 23.1% sequence divergence from S. setosa. Two possibilities for the evolutionary origin of this species are proposed, namely, that it represents a relict species from the ancient Paratethys, or that it represents another chaetognath species that colonized the Black Sea more recently. Even though the exact timing of disjunction of S. setosa populations remains unclear, on the basis of the geological and paleoclimatic history of the European basins and our estimates of net nucleotide divergence, we suggest that disjunct populations arose through vicariance resulting from the cyclical changes in temperature and sea levels during the Pleistocene. We conclude that these populations have remained disjunct, not because of limited dispersal ability, but because of the inability to maintain viable populations in suboptimal, geographically intermediate areas.

Pattern and process in the evolution of insect-plant associations: Yponomeuta as an example

Phylogenetic studies are increasing our understanding of the evolution of associations between phytophagous insects and their host plants. Sequential evolution, i.e. the shift of insect herbivores onto pre‐existing plant species, appears to be much more common than coevolution, where reciprocal selection between interacting insects and plants is thought to induce chemical diversification and resistance in plants and food specialization in insects.

Molecular Epidemiology ofFonsecaeaSpecies

To assess population diversities among 81 strains of fungi in the genus Fonsecaea that had been identified down to species level, we applied amplified fragment-length polymorphism (AFLP) technology and sequenced the internal transcribed spacer regions and the partial cell division cycle, beta-tubulin, and actin genes. Many species of the genus Fonsecaea cause human chromoblastomycosis. Strains originated from a global sampling of clinical and environmental sources in the Western Hemisphere, Asia, Africa, and Europe. According to AFLP fingerprinting, Fonsecaea isolates clustered in 5 groups corresponding with F. pedrosoi, F. monophora, and F. nubica: the latter 2 species each comprised 2 groups, and F. pedrosoi appeared to be of monophyletic origin. F. pedrosoi was found nearly exclusively in Central and South America. F. monophora and F. nubica were distributed worldwide, but both showed substantial geographic structuring. Clinical cases outside areas where Fonsecaea is endemic were probably distributed by human migration.

Global molecular epidemiology and genetic diversity of Fusarium, a significant emerging group of human opportunists from 1958 to 2015

Fusarium is a rapidly emerging, multidrug-resistant genus of fungal opportunists that was first identified in 1958 and is presently recognized in numerous cases of fusariosis each year. The authors examined trends in global Fusarium distribution, clinical presentation and prevalence since 1958 with the assumption that their distributions in each region had remained unaltered. The phylogeny and epidemiology of 127 geographically diverse isolates, representing 26 Fusarium species, were evaluated using partial sequences of the RPB2 and TEF1 genes, and compared with AFLP fingerprinting data. The molecular data of the Fusarium species were compared with archived data, which enabled the interpretation of hundreds of cases published in the literature. Our findings indicate that fusariosis is globally distributed with a focus in (sub)tropical areas. Considerable species diversity has been observed; genotypic features did not reveal any clustering with either the clinical data or environmental origins. This study suggests that infections with Fusarium species might be truly opportunistic. The three most common species are F. falciforme and F. keratoplasticum (members of F. solani species complex), followed by F. oxysporum (F. oxysporum species complex).

Experimental interspecific hybridization in Daphnia

Hybridization is a common phenomenon in Daphnia (Cladocera; Anomopoda); interspecific hybrids have been found between several species and hybrids are found in many European lakes. Although much information on the morphology, ecology and genetics of hybrids is available, little is known about the level of reproductive isolation among species or about the relative fitness of hybrids and parental species. In order to facilitate studies on differentiation and speciation processes and comparative experimental studies on hybrids and recombinant genotypes, we present the first successful laboratory crossing experiments of two different Daphnia species, D. galeata and D. cucullata. Males and sexual females from two D. galeata and two D. cucullata clones were reciprocally crossed, juveniles hatched from resting eggs and reared until maturity. Hatching and juvenile survival rates of hybrids were relatively low (12.1% and 24%, respectively). D. galeata and D. cucullata clones vary in their level of successful interspecific matings and in the number of subsequent offspring. In general, hybrid crosses between D. cucullata females and D. galeata males were more successful than reciprocal crosses.