TEMPERATURE-BASED EXTRINSIC REPRODUCTIVE ISOLATION IN TWO SPECIES OF DROSOPHILA

Drosophila santomea and D. yakuba are sister species that live on the African volcanic island of São Tomé, where they are ecologically isolated: D. yakuba inhabits low-altitude open and semiopen habitats while D. santomea lives in higher-elevation rain and mist forest. To determine whether this spatial isolation reflected differential preference for and tolerance of temperature, we estimated fitness components of both species at different temperatures as well as their behavioral preference for certain temperatures. At higher temperatures, especially 28°C, D. santomea was markedly inferior to D. yakuba in larval survival, egg hatchability, and longevity. Moreover, D. santomea females, unlike those of D. yakuba , become almost completely sterile after exposure to a temperature of 28°C, and conspecific males become semisterile. Drosophila santomea adults prefer temperatures 2–3°C lower than do adults of D. yakuba . Drosophila santomea , then, is poorly adapted to high temperature, partially explaining its restriction to cool, high habitats, which leads to extrinsic premating isolation and immigrant inviability. Rudimentary genetic analysis of the interspecific difference in egg hatchability and larval survival showed that these differences are due largely to cytoplasmic effects and to autosomal genes, with sex chromosomes playing little or no role.     

Do parasitoid preferences for different host species match virulence?

Abstract.  Leptopilina boulardi is a parasitoid wasp specialist of Drosophila larvae of the melanogaster subgroup. In Mediterranean areas, natural populations are highly virulent against their main host Drosophila melanogaster . In Congo, populations are less virulent against D. melanogaster but are able to develop successfully inside the tropical African species Drosophila yakuba . Host preferences are compared between two laboratory isofemale lines of L. boulardi , obtained from populations of Congo and Tunisia, respectively, and differing in virulence levels against D. melanogaster and D. yakuba . Host selection is studied by offering female parasitoids a choice between larvae of the two host species. In agreement with optimal foraging models, the line highly virulent against D. melanogaster shows a clear preference for this host species. The other line, less virulent against D. melanogaster but more virulent against D. yakuba , prefers to oviposit on D. yakuba . Such preferences can be observed after a period of host-patch exploitation only, suggesting that experience plays an important role in the host-selection process. These results evidence the existence of intraspecific variability in preference between two host species in L. boulardi , a major requisite in theoretical models of parasite specialization by the host. They also sustain the hypothesis that intraspecific variation in parasitoid preferences between host species might mirror intraspecific variation in virulence.     

Conspecific sperm precedence in sister species of Drosophila with overlapping ranges

Abstract Barriers to gene flow that act after mating but before fertilization are often overlooked in studies of reproductive isolation. Where species are sympatric, such "cryptic' isolating barriers may be important in maintaining species as distinct entities. Drosophilayakuba and its sister species D. santomea have overlapping ranges on the island of Sao Tome, off the coast of West Africa. Previous studies have shown that the two species are strongly sexually isolated. However, the degree of sexual isolation observed in the laboratory cannot explain the low frequency (–1%) of hybrids observed in nature. This study identifies two "cryptic" isolating barriers that may further reduce gene flow between D. yakuba andD. santomea where they are sympatric. First, noncompetitive gametic isolation has evolved between D. yakuba and D. santomea: heterospecific matings between the two species produce significantly fewer offspring than do conspecific matings. Second, conspecific sperm precedence (CSP) occurs when D. yakuba females mate with conspecific and heterospecific males. However, CSP is asymmetrical: D. santomea females do not show patterns of sperm usage consistent with CSP. Drosophila yakuba and D. santomea females also differ with respect to remating propensity after first mating with conspecific males. These results suggest that noncompetitive and competitive gametic isolating barriers may contribute to reproductive isolation between D. yakuba and D. santomea.     

Genomewide comparative analysis of the highly abundant transposable element DINE-1 suggests a recent transpositional burst in Drosophila yakuba

DINE-1 (Drosophila interspersed element) is the most abundant repetitive sequence in the Drosophila genome derived from transposable elements. It comprises >1% of the Drosophila melanogaster genome (DMG) and is believed to be a relic from an ancient transpositional burst that occurred approximately 5-10 MYA. We performed a genomewide comparison of the abundance, sequence variation, and chromosomal distribution of DINE-1 in D. melanogaster and D. yakuba. Unlike the highly diverged copies in the DMG (pairwise distance approximately 15%), DINE-1's in the Drosophila yakuba genome (DYG) have diverged by only 3.4%. Moreover, the chromosomal distribution of DINE-1 in the two species is very different, with a significant number of euchromatic insertions found only in D. yakuba. We propose that these different patterns are caused by a second transpositional burst of DINE-1's in the D. yakuba genome approximately 1.5 MYA. On the basis of the sequence of these recently transposed copies, we conclude that DINE-1 is likely to be a family of nonautomomous DNA transposons. Analysis of the chromosomal distribution of two age groups of DINE-1's in D. yakuba indicates that (1) there is a negative correlation between recombination rates and the density of DINE-1's and (2) younger copies are more evenly distributed in the chromosome arms, while older copies are mostly located near the centromere regions. Our results fit the predictions of a selection-transposition balance model. Our data on whole-genome comparison of a highly abundant TE among Drosophila sibling species demonstrate the unexpectedly dynamic nature of TE activity in different host genomes.     

Local changes in GC/AT substitution biases and in crossover frequencies on Drosophila chromosomes

I present here evidence of remarkable local changes in GC/AT substitution biases and in crossover frequencies on Drosophila chromosomes. The substitution pattern at 10 loci in the telomeric region of the X chromosome was studied for four species of the Drosophila melanogaster species subgroup. Drosophila orena and Drosophila erecta are clearly the most closely related species pair (the erecta complex) among the four species studied; however, the overall data at the 10 loci revealed a clear dichotomy in the silent substitution patterns between the AT-biased- substitution melanogaster and erecta lineages and the GC-biased-substitution yakuba and orena lineages, suggesting two or more independent changes in GC/AT substitution biases. More importantly, the results indicated a between- loci heterogeneity in GC/AT substitution bias in this small region independently in the yakuba and orena lineages. Indeed, silent substitutions in the orena lineage were significantly biased toward G and C at the consecutive yellow, lethal of scute, and asense loci, but they were significantly biased toward A and T at sta. The substitution bias toward G and C was centered in different areas in yakuba (significantly biased at EG:165H7.3, EG:171D11.2, and suppressor of sable). The similar silent substitution patterns in coding and noncoding regions, furthermore, suggested mutational biases as a cause of the substitution biases. On the other hand, previous study reveals that Drosophila yakuba has about 20-fold higher crossover frequencies in the telomeric region of the X chromosome than does D. melanogaster; this study revealed that the total genetic map length of the yakuba X chromosome was only about 1.5 times as large as that of melanogaster and that the map length of the X-telomeric y-sta region did not differ between Drosophila yakuba and D. erecta. Taken together, the data strongly suggested that an approximately 20- fold reduction in the X-telomeric crossover frequencies occurred in the ancestral population of D. melanogaster after the melanogaster-yakuba divergence but before the melanogaster-simulans divergence.     

The Genus Spiniferites Mantell, 1850 (Dinophyceae)

Following the recent official rejection of proposals for conservation of the generic name Hystrichosphaera, the diagnosis of its senior synonym Spiniferites is expanded in order to incorporate additional information gained since 1850 and all species according with the expanded diagnosis are listed (24 species are transferred to Spiniferites). The cyst-family name “Hystrichosphaeraceae” is replaced by the new name “Spiniferitaceae”.     

Evolutionary relationships to parasitism by seven species of the Drosophila melanogaster subgroup

Parasitic wasps are an important component of the niche of Drosophila species. The susceptibility to the Cynipid Leptopilina boulardi was estimated in the seven sibling species of Drosophila belonging to the melanogaster subgroup. Three categories of Hies can be distinguished, according to the level of cellular immune reaction and success of parasitism. Drosophila melanogaster and D. mauritiana belong to the category 1, specified by no encapsulative reaction and a high rate of successful parasitism. Category 2, characterized by a moderate encapsulation rate and a high mortality include D. simulans.5, D. erecta and D. orena . Category 3, with D. yakuba and D. tcissien , is specified by a very low rate or absence of successful parasitism due to a highly efficient immune cellular reaction. This classification parallels the phylogenic relationship based upon polytene chromosome banding sequences. Such specific ditferences in susceptibility to parasites may plan an important role in the competition between these species in Africa.     

Multilocus analysis of introgression between two sympatric sister species of Drosophila: Drosophila yakuba and D. santomea

Drosophila yakuba is widely distributed in sub-Saharan Africa, while D. santomea is endemic to the volcanic island of S?o Tomé in the Atlantic Ocean, 280 km west of Gabon. On S?o Tomé, D. yakuba is found mainly in open lowland forests, and D. santomea is restricted to the wet misty forests at higher elevations. At intermediate elevations, the species form a hybrid zone where hybrids occur at a frequency of approximately 1%. To determine the extent of gene flow between these species we studied polymorphism and divergence patterns in 29 regions distributed throughout the genome, including mtDNA and three genes on the Y chromosome. This multilocus approach, together with the comparison to the two allopatric species D. mauritiana and D. sechellia, allowed us to distinguish between forces that should affect all genes and forces that should act on some genes (e.g., introgression). Our results show that D. yakuba mtDNA has replaced that of D. santomea and that there is also significant introgression for two nuclear genes, yellow and salr. The majority of genes, however, has remained distinct. These two species therefore do not form a "hybrid swarm" in which much of the genome shows substantial introgression while disruptive selection maintains distinctness for only a few traits (e.g., pigmentation and male genitalia).     

Mitochondrial DNA evolution in themelanogaster species subgroup ofDrosophila

Detailed restriction maps (40 cleavage sites on average) of mitochondrial DNAs (mtDNAs) from the eight species of the melanogaster species subgroup of Drosophila were established. Comparison of the cleavage sites allowed us to build a phylogenetic tree based on the matrix of nucleotide distances and to select the most parsimonious network. The two methods led to similar results, which were compared with those in the literature obtained from nuclear characters. The three chromosomally homosequential species D. simulans, D. mauritiana, and D. sechellia are mitochondrially very related, but exhibit complex phylogenetic relationships. D. melanogaster is their closest relative, and the four species form a monophyletic group (the D. melanogaster complex), which is confirmed by the shared unusual length of their mt genomes (18-19 kb). The other four species of the subgroup (D. yakuba, D. teissieri, D. erecta, and D. orena) are characterized by a much shorter mt genome (16-16.5 kb). The monophyletic character of the D. yakuba complex, however, is questionable. Two species of this complex, D. yakuba and D. teissieri, are mitochondrially indistinguishable (at the level of our investigation) in spite of their noticeable allozymic and chromosomal divergence. Finally, mtDNA distances were compared with the nuclear-DNA distances thus far established. These sequences seem to evolve at rather similar rates, the mtDNA rate being barely double that of nuclear DNA.     

Comparative analysis of transposable elements in the melanogaster subgroup sequenced genomes

Transposable elements (TEs) are indwelling components of genomes, and their dynamics have been a driving force in genome evolution. Although we now have more information concerning their amounts and characteristics in various organisms, we still have little data from overall comparisons of their sequences in very closely-related species. While the Drosophila melanogaster genome has been extensively studied, we have only limited knowledge regarding the precise TE sequences in the genomes of the related species Drosophila simulans, Drosophila sechellia and Drosophila yakuba. In this study we analyzed the number and structure of TE copies in the sequenced genomes of these four species. Our findings show that, unexpectedly, the number of TE insertions in D. simulans is greater than that in D. melanogaster, but that most of the copies in D. simulans are degraded and in small fragments, as in D. sechellia and D. yakuba. This suggests that all three species were invaded by numerous TEs a long time ago, but have since regulated their activity, as the present TE copies are degraded, with very few full-length elements. In contrast, in D. melanogaster, a recent activation of TEs has resulted in a large number of almost-identical TE copies. We have detected variants of some TEs in D. simulans and D. sechellia, that are almost identical to the reference TE sequences in D. melanogaster, suggesting that D. melanogaster has recently been invaded by active TE variants from the other species. Our results indicate that the three species D. simulans, D. sechellia, and D. yakuba seem to be at a different stage of their TE life cycle when compared to D. melanogaster. Moreover, we show that D. melanogaster has been invaded by active TE variants for several TE families likely to come from D. simulans or the ancestor of D. simulans and D. sechellia. The numerous horizontal transfer events implied to explain these results could indicate introgression events between these species.     

Divergence in Multiple Courtship Song Traits between Drosophila santomea and D. yakuba

The courtship song of Drosophila is useful for species recognition and sexual selection. A new species of the melanogaster group of Drosophila , D. santomea , has recently been described from the island of São Tomé in the Gulf of Guinea. We describe the courtship song of D. santomea and compare it with that of its sibling species D. yakuba . Both species have a relatively unusual song pattern for melanogaster-group species, in that they have two types of pulse song but no sine song. There are large differences in the inter-pulse interval of both types of song, but no major differences in pulse shape or intrapulse frequency between the species. The song of D. yakuba is similar in lines from the African mainland (allopatric to D. santomea ) and from São Tomé (sympatric). We test if song pattern might influence sexual isolation by examining the mating success of wingless males with homo- and hetero-specific females. We show that song pattern contributes to sexual stimulation, but the differences in song patterns alone are unlikely to explain patterns of sexual isolation such as the asymmetrical isolation seen between species.     

The Recent species of Adeonella (Bryozoa: Cheilostomata) including descriptions of fifteen new species

The genus Adeonella Busk (1884) comprises 41 species, 15 of which are considered to be new to science. Adeonella lichenoides (Lamarck) is regarded as the senior synonym of A. platalea (Busk) (= A. polymorpha Busk), the type species of the genus. Adeonella is distributed throughout the Indo-West Pacific realm, the Mediterranean, and the South Atlantic Ocean. Its highest diversity is found off the east coasts of southern Africa. Thirty-one species (15 new to science) are described and illustrated, morphology and geographical distribution are discussed and a key is provided to all recognized species.     

The magnitude of behavioral isolation is affected by characteristics of the mating community

Gene exchange between species occurs in areas of secondary contact, where two species have the opportunity to hybridize. If heterospecific males are more common than conspecific males, females will experience more encounters with males of other species. These encounters might increase the likelihood of heterospecific matings, and lead to the production of hybrid progeny. I studied the mating behavior of two pairs of sibling species endemic to Africa: Drosophila yakuba/Drosophila santomea and Drosophila simulans/Drosophila sechellia. Drosophila yakuba and D. simulans are cosmopolitan species widely distributed in the African continent, while D. santomea and D. sechellia are island endemics. These pairs of species hybridize in nature and have the potential to exchange genes in natural conditions. I used these two pairs of Drosophila species, and constructed mating communities of different size and different heterospecific:conspecific composition. I found that both the total number of potential mates and the relative frequency of conspecific versus heterospecific males affect female mating decisions in the cosmopolitan species but not in the island endemics. These results suggest that the population characteristics, in which mating occurs, may affect the magnitude of premating isolation. Community composition might thus facilitate, or impair, gene flow between species.     

A key to the species of Mugilidae (Pisces) in the Northeastern Atlantic and Mediterranean, with explanatory notes

A key is given for identifying the eight species of Mugilidae of the Northeast Atlantic and Mediterranean. The use of Risso's names for four species is justified and explained and recognition is given to Tortonese's discovery that Mugil capurrii (Perugia) is a senior synonym of M. monodi Chabanaud. The relationship of Liza saliens to certain species of West and southern Africa is held to be additional support for the recognition of Protomugil Popov as a subgenus of Liza including these species. Chelon Röse, with type-species Mugil chelo Cuvier (= Chelon labrosus (Risso)), is recognized, following Jordan & Evermann (1917) and Schultz (1946), but its limits are differently conceived so as to include Crenimugil Schultz rather than Liza Jordan & Swain. The latter is held to comprise about 30 Old World species. The status of Chelon bispinosus (Bowdich) is discussed, with its bearing on possible population-differences in C. labrosus and the relationship of this species with C. crenilabis. The identity of Liza seheli (Forskål) is discussed and reasons are given for disagreeing with Wimpenny and others in applying this name to the species found at both ends of the Suez Canal zone. The latter is L. carinata (Ehrenberg in Cuvier & Valenciennes). Myxus maroccensis Mohr is believed to be a young Liza ramada.     

Identification of the Diadegma species (Hymenoptera: Ichneumonidae, Campopleginae) attacking the diamondback moth, Plutella xylostella (Lepidoptera: Plutellidae)

The species of Diadegma that attack Plutella xylostella (Linnaeus) are revised. Following a morphometric study involving principal components and discriminant analyses, seven distinct morphospecies are recognized. One species is described as new: D. novaezealandiae from New Zealand. Diadegma mollipla (Holmgren) is the name for the species from sub-Saharan Africa and some Indian Ocean and South Atlantic islands. Diadegma varuna Gupta syn. nov. and D. niponica Kusigemati syn. nov. are both synonymized with D. fenestrale (Holmgren). Diadegma xylostellae Kusigemati is strongly presumed to be a synonym of D. semiclausum (Hellén). An illustrated identification key is provided and each species is described in a standard way.     

Synthesis patterns of a set of follicle cell proteins in Drosophila melanogaster sibling species

The protein synthesis pattern of a set of stage and tissue specific proteins has previously been described in Drosophila melanogaster. The analysis of this set of follicle cell proteins (Fc proteins) is here extended to cover several sibling species of Drosophila melanogaster, namely D. simulans, D. mauritiana, D. erecta and D. yakuba. Even though a similar set of proteins were synthesized in these species, minor differences in size of the proteins were found between the species. Some of the species exhibited variation within species.