Monogamy in the maternally mouthbrooding Lake Tanganyika cichlid fish Tropheus moorii

Supported by evidence for assortative mating and polygynandry, sexual selection through mate choice was suggested as the main force driving the evolution of colour diversity of haplochromine cichlids in Lakes Malawi and Victoria. The phylogenetically closely related tribe Tropheini of Lake Tanganyika includes the genus Tropheus, which comprises over 100 colour variants currently classified into six morphologically similar, polyphyletic species. To assess the potential for sexual selection in this sexually monochromatic maternal mouthbrooder, we used microsatellite-based paternity inference to investigate the mating system of Tropheus moorii. In contrast to haplochromines in Lake Malawi, multiple paternity is rare or even absent in broods of T. moorii. Eighteen of the 19 analysed families were consistent with genetic monogamy, while either a mutation or more than one sire explained the genotype of one offspring in another brood. We discuss the differences in breeding behaviour between T. moorii and the Lake Malawi haplochromines, and evaluate additional factors or alternatives to sexual selection as promoters of colour diversification. A preliminary survey of other Tropheini species suggested that multiple paternity is infrequent in the entire tribe.     

Parasite escape through trophic specialization in a species flock

Adaptive radiation occurs when species diversify rapidly to occupy an array of ecological niches. As opportunities for parasite infection and transmission may greatly vary among these niches, adaptive radiation is expected to be associated with a turnover of the parasite community. As major agents of natural and sexual selection, parasites may play a central role in host diversification. The study of parasite turnover may thus be of general relevance and could significantly improve our understanding of adaptive radiation. In this study, we examined the parasite faunas of eleven species belonging to the tribe Tropheini, one of several adaptive radiations of cichlid fishes in Lake Tanganyika. The most parsimonious ancestral foraging strategy among the Tropheini is relatively unselective substrate browsing of aufwuchs. Several lineages evolved more specialized foraging strategies, such as selective combing of microscopic diatoms or picking of macro‐invertebrates. We found that representatives of these specialized lineages bear reduced infection with food‐web‐transmitted acanthocephalan helminths, but not with parasites with a direct life cycle. Possibly, the evolution of selective foraging strategies entailed reduced ingestion of intermediate invertebrate hosts of acanthocephalans. We conclude that some species belonging to the Tropheini virtually escape acanthocephalan infection as a by‐product of trophic specialization.     

Phylogenomics of trophically diverse cichlids disentangles processes driving adaptive radiation and repeated trophic transitions

Cichlid fishes of the tribe Tropheini are a striking case of adaptive radiation, exemplifying multiple trophic transitions between herbivory and carnivory occurring in sympatry with other established cichlid lineages. Tropheini evolved highly specialized eco‐morphologies to exploit similar trophic niches in different ways repeatedly and rapidly. To better understand the evolutionary history and trophic adaptations of this lineage, we generated a dataset of 532 targeted loci from 21 out of the 22 described Tropheini species. We resolved the Tropheini into seven monophyletic genera and discovered one to be polyphyletic. The polyphyletic genus, Petrochromis, represents three convergent origins of the algae grazing trophic specialization. This repeated evolution of grazing may have been facilitated by adaptive introgression as we found evidence for gene flow among algae grazing genera. We also found evidence of gene flow among algae browsing genera, but gene flow was restricted between herbivorous and carnivorous genera. Furthermore, we observed no evidence supporting a hybrid origin of this radiation. Our molecular evolutionary analyses suggest that opsin genes likely evolved in response to selection pressures associated with trophic ecology in the Tropheini. We found surprisingly little evidence of positive selection in coding regions of jaw‐shaping genes in this trophically diverse lineage. This suggests low degrees of freedom for further change in these genes, and possibly a larger role for regulatory variation in driving jaw adaptations. Our study emphasizes Tropheini cichlids as an important model for studying the evolution of trophic specialization and its role in speciation.     

Mitochondrial phylogeny of the endemic mouthbrooding lineages of cichlid fishes from Lake Tanganyika in eastern Africa

Of the three cichlid species flocks in eastern Africa, Lake Tanganyika harbors the oldest species assemblage, which is also the most diverse morphologically and behaviorally. For 12 species (20 individuals) of 12 genera of the tribe Ectodini, 852 bp from two segments (cytochrome b and control region) of the mitochondrial genome were sequenced. In addition, orthologous sequences were obtained from eight species (11 individuals) representing other mouthbrooding lineages from Lake Tanganyika. Comparisons of sequence divergences revealed that the single Tanganyikan tribe Ectodini appears to be approximately five times older than the whole Lake Malawi cichlid species flock, suggesting that the radiation of the Tanganyikan mouthbrooding lineages took place long before the species flocks of Lakes Malawi and Victoria evolved. Seven of nine surveyed tribes of Tanganyikan cichlids appear to be approximately equally divergent, and this seems to corroborate the hypothesis of a rapid simultaneous formation of lineages at an early stage in the history of the Lake Tanganyika species flock. The close genetic relationship between the endemic Tropheus lineage and a nonendemic "Haplochromine," Astatotilapia burtoni, indicates that members of the tribe Tropheini may be the sister group of the cichlid flocks of Lakes Malawi and Victoria. The phylogenetic analyses demonstrate the monophyly of the Ectodini and identify the Cyprichromini as their sister group among the Tanganyikan cichlids. Within the tribe Ectodini the molecular data suggest both a branching pattern different than that previously proposed and a subdivision of the Ectodini into four clades, instead of the two originally described. The previously postulated model of morphological transformations believed to be responsible for the drastically different types of ecological specialization found among the Ectodini might therefore be in need of reinterpretation. Characters immediately related to foraging and nutrition seem to be particularly prone to homoplasy, even among members of a single lineage of cichlid fishes.      

Why close relatives make bad neighbours: phylogenetic conservatism in niche preferences and dispersal disproves Darwin's naturalization hypothesis in the thistle tribe

The number of exotic plant species that have been introduced into the United States far exceeds that of other groups of organisms, and many of these have become invasive. As in many regions of the globe, invasive members of the thistle tribe, Cardueae, are highly problematic in the California Floristic Province, an established biodiversity hotspot. While Darwin's naturalization hypothesis posits that plant invaders closely related to native species would be at a disadvantage, evidence has been found that introduced thistles more closely related to native species are more likely to become invasive. To elucidate the mechanisms behind this pattern, we modelled the ecological niches of thistle species present in the province and compared niche similarity between taxa and their evolutionary relatedness, using fossil‐calibrated molecular phylogenies of the tribe. The predicted niches of invasive species were found to have higher degrees of overlap with native species than noninvasive introduced species do, and pairwise niche distance was significantly correlated with phylogenetic distance, suggesting phylogenetic niche conservatism. Invasive thistles also displayed superior dispersal capabilities compared to noninvasive introduced species, and these capabilities exhibited a phylogenetic signal. By analysing the modelled ecological niches and dispersal capabilities of over a hundred thistle species, we demonstrate that exapted preferences to the invaded environment may explain why close exotic relatives may make bad neighbours in the thistle tribe.     

Species-Specific Population Structure in Rock-Specialized Sympatric Cichlid Species in Lake Tanganyika,East Africa

Species richness and geographical phenotypic variation in East African lacustrine cichlids are often correlated with ecological specializations and limited dispersal. This study compares mitochondrial and microsatellite genetic diversity and structure among three sympatric rock-dwelling cichlids of Lake Tanganyika, Eretmodus cyanostictus, Tropheus moorii, and Ophthalmotilapia ventralis. The species represent three endemic, phylogenetically distinct tribes (Eretmodini, Tropheini, and Ectodini), and display divergent ecomorphological and behavioral specialization. Sample locations span both continuous, rocky shoreline and a potential dispersal barrier in the form of a muddy bay. High genetic diversity and population differentiation were detected in T. moorii and E. cyanostictus, whereas much lower variation and structure were found in O. ventralis. In particular, while a 7-km-wide muddy bay curtails dispersal in all three species to a similar extent, gene flow along mostly continuous habitat appeared to be controlled by distance in E. cyanostictus, further restricted by site philopatry and/or minor habitat discontinuities in T. moorii, and unrestrained in O. ventralis. In contrast to the general pattern of high gene flow along continuous shorelines in rock-dwelling cichlids of Lake Malawi, our study identifies differences in population structure among stenotopic Lake Tanganyika species. The amount of genetic differentiation among populations was not related to the degree of geographical variation of body color, especially since more phenotypic variation is observed in O. ventralis than in the genetically highly structured E. cyanostictus. [Reviewing Editor: Dr. Rafael Zardoya]     

Australian Snakes Do Not Follow Bergmann’s Rule

Bergmann’s Rule (i.e., the tendency of body size to increase with decreasing environmental temperature) was originally explained by a mechanism that is unique to endotherms. Nevertheless, geographic variation of body size of ectotherms, including snakes, is increasingly studied, and some claim that the rule should apply to ectotherms, or to thermoregulating ectotherms. Such studies usually focus on assemblages or on species in a region, but mostly ignore species’ ecological and biological traits when seeking biogeographic patterns. We examined the relationship between environmental temperatures and body size of 146 Australian snake species. We examined this relationship while considering the effects of ecological traits (activity time and habitat use), climatic variables which are thought to influence snake body size, and shared ancestry. Our finding suggest that Bergmann’s Rule is not a valid generalization across species of Australian snakes. Furthermore, ecological traits greatly influence the relationship between snake body size and environmental temperature. Body size of fossorial species decreases with environmental temperature, whereas body size of nocturnal, surface active species increases. Body size of diurnal, surface active species is not related to environmental temperature. Our results indicate that lumping all species in a clade together is misleading, and that ecological traits profoundly affect the geographic variation of snake body size. Though environmental temperature generally does not exert a strong selective force on snake body size, this relationship differs for taxa exhibiting different ecological traits.     

Mycoparasitism between Squamanita paradoxa and Cystoderma amianthinum (Cystodermateae, Agaricales)

Circumstantial evidence, mostly morphological and ecological, points to ten different mushroom host species for up to fifteen species of the mycoparasitic genus Squamanita. Here, molecular evidence confirms Cystoderma amianthinum as the host for S. paradoxa, a sporadically occurring and rarely collected mycoparasite with extreme host specificity. This is only the second study to use molecular techniques to reveal or confirm the identity of a cecidiocarp of Squamanita species. Phylogenetic analysis of combined nuclear ribosomal RNA genes suggests the monophyly of Squamanita, Cystoderma, and Phaeolepiota, a clade referred to as the tribe Cystodermateae. If true, S. paradoxa and C. amianthinum would represent a relatively closely related species pair involved in a mycoparasitic symbiosis.     

The ecological drivers of nuptial color evolution in darters (Percidae: Etheostomatinae)

Closely related animal lineages often vary in male coloration, and ecological selection is hypothesized to shape this variation. The role of ecological selection in inhibiting male color has been documented extensively at the population level, but relatively few studies have investigated the evolution of male coloration across a clade of closely related species. Darters are a diverse group of fishes that vary in the presence of elaborate male nuptial coloration, with some species exhibiting vivid color patterns and others mostly or entirely achromatic. We used phylogenetic logistic regression to test for correlations between the presence/absence of color traits across darter species and the ecological conditions in which these species occur. Environmental variables were correlated with the presence of nuptial color in darters with colorful species tending to inhabit environments that would support fewer predators and potentially transmit a broader spectrum of natural light compared to species lacking male coloration. We also tested the color preferences of a common darter predator, largemouth bass, and found that it exhibits a strong preference for red, providing further evidence of predation as a source of selection on color evolution in darters. Ecological selection therefore appears to be an important factor in dictating the presence or absence of male coloration in this group of fishes.     

The Systematic and Ecological Significance of Anatomical Characters of Leaves in the Tribe Thermopsideae (Fabaceae)

Anatomical characters of leaves of Chinese species of the tribe Thermopsideae, including 3 genera and 13 species, were compared. The results show that the differentiation of mesophyll cells, stomatal types, stomatal distribution and density, shape of epidermal cell, trichome shape and density, cuticular membrane and waxy ornamentation have important systematic and ecological significance among the genera of the tribe. With regard to leaf architecture, pollen morphology and external morphology studies, we suggest that Ammopiptanthus represents an isolated and primitive genus in the tribe and Piptanthus is closely related to Thermopsis. Although different vein-endings, stomatal types and trichomes indicate systematic differences, other characters, such as mesophyll type, stomatal distribution, trichome density etc. are the result of ecological adaptation among the genera. Results of the present study confirm our previous conclusion that two evolutionary trends exist in the tribe, the first towards xerophilization and the second towards cold-adaptation They are caused by the desertification of the central Asian area and the heave of intermountainous plateau. Ecological isolation may be a main evolutary motivation among the taxa in the tribe. We have obs'erved that stomatal density and size in the xeromorphic genus, Ammopipianthus, are not simply a result of the general trend in xerophytes to have high stomatal density correlating with the relatively small size. On the contrary, stomata in this genus are larger than those in the mesophytic genus, Piptanthus, and the density on the lower surface is much less than in the latter. However, stomatal density on the upper surface follows the trend in concordance with increasing aridity and altitute.     

Patterns in the geographic ranges of the world's woodpeckers

We used data on the world's woodpeckers to test for patterns in the geographic distributions of a single group of closely related species. The frequency distribution of woodpecker geographic range sizes is approximately lognormal. Most variation in range sizes is explained by differences between species within genera; that is, range size seems to be an evolutionarily labile trait. The largest woodpecker ranges are found in Eurasia, both when absolute differences are compared and when range size is measured as a proportion of estimated available habitat. Notably, there is a negative relationship between the mean range sizes attained by species in a genus or tribe in South America and the mean ranges attained by species in the same tribe or genus in North America. Large-bodied species tend to be more widely distributed and to live at higher latitudes, but both tendencies disappear if the taxonomic relatedness of species is controlled for. Species living at high latitudes also tend to be more widely distributed. This relationship seems largely due to the effect of North American woodpeckers, which show it even when the taxonomic relatedness of species is controlled. Small continents generally have more woodpecker species than do large ones. Woodpecker geographic range sizes are smaller the more woodpecker species inhabit an area. Species show less overlap in their geographic ranges with species of similar than with species of dissimilar body size. The implications of these results for our understanding of patterns in geographic range sizes are discussed.     

CHROMOSOME NUMBERS IN COMPOSITAE VII: ASTEREAE III

Reports of 129 new chromosome counts are made for the tribe Astereae of Compositae. They are mostly based on determinations of meiotic material, including first counts for one genus and 43 species or subspecies. Counts are now available for more than 63 of the 100-120 genera and 564 of the approximately 2,000 species in the tribe. Three of every four genera with more than one species counted show more than one chromosome number; 15 genera have species with populations with different numbers. Such variation is very high and indicates the need for more detailed cytotaxonomical study in the group.     

Climatic adaptation and ecological divergence between two closely related pine species in Southeast China

Climate is one of the most important drivers for adaptive evolution in forest trees. Climatic selection contributes greatly to local adaptation and intraspecific differentiation, but this kind of selection could also have promoted interspecific divergence through ecological speciation. To test this hypothesis, we examined intra‐ and interspecific genetic variation at 25 climate‐related candidate genes and 12 reference loci in two closely related pine species, Pinus massoniana Lamb. and Pinus hwangshanensis Hisa, using population genetic and landscape genetic approaches. These two species occur in Southeast China but have contrasting ecological preferences in terms of several environmental variables, notably altitude, although hybrids form where their distributions overlap. One or more robust tests detected signals of recent and/or ancient selection at two‐thirds (17) of the 25 candidate genes, at varying evolutionary timescales, but only three of the 12 reference loci. The signals of recent selection were species specific, but signals of ancient selection were mostly shared by the two species likely because of the shared evolutionary history. F_ST outlier analysis identified six SNPs in five climate‐related candidate genes under divergent selection between the two species. In addition, a total of 24 candidate SNPs representing nine candidate genes showed significant correlation with altitudinal divergence in the two species based on the covariance matrix of population history derived from reference SNPs. Genetic differentiation between these two species was higher at the candidate genes than at the reference loci. Moreover, analysis using the isolation‐with‐migration model indicated that gene flow between the species has been more restricted for climate‐related candidate genes than the reference loci, in both directions. Taken together, our results suggest that species‐specific and divergent climatic selection at the candidate genes might have counteracted interspecific gene flow and played a key role in the ecological divergence of these two closely related pine species.     

Étude des réactions croisées entre les lectines de légumineuses: In systématique et phylogénétique

Seed extracts from 56 legume species were examined for immunological relatedness using 11 different lectin antisera. The immunological cross-reactions observed indicate that only lectins from species belonging to the tribe Vicieae share common antigenic determinants and are evolutionary closely related. These results suggest that the immunochemical study of the Vicieae lectins may provide a powerful tool for determining the systematic and phylogenetic relationships within the tribe.     

Exine micromorphology and ultrastructure in Neottieae (Epidendroideae,Orchidaceae)

The diverse epidendroid orchid tribe Neottieae is characterized by multiple transitions between autotrophy and mycoheterotrophy, allogamous and autogamous mating systems, pollen released as tetrads or monads, and pollen exine tectate or semitectate. We use transmission and scanning electron microscopy on pollen of ten species of Neottieae to investigate whether the differences in pollen aggregation and exine micromorphology and ultrastructure reflect phylogenetic relationships, or whether this variation is subject to ecological constraints. Our results showed that differences in exine micromorphology are mostly concordant with phylogenetic relationships in Neottieae, i.e. an ascending tendency of pollen ornamentation from tectate (Cephalanthera) to semitectate (e.g. Neottia). In contrast, pollen aggregation, when plotted on the most recent phylogeny, shows repeated transitions between monads and tetrads that could be related to ecological constraints. Tetrads are present in species that are nectar rewarding, whereas monads are common in deceptive species. Cephalanthera is characterized by recalcitrant pollen, including the frequent occurrence of collapsed pollen. In this genus, the observed shifts from allogamous to autogamous or cleistogamous mating systems could help to reduce pollen damage caused by exposure to dry habitats.     

Beyond trophic morphology: stable isotopes reveal ubiquitous versatility in marine turtle trophic ecology

The idea that interspecific variation in trophic morphology among closely related species effectively permits resource partitioning has driven research on ecological radiation since Darwin first described variation in beak morphology among Geospiza. Marine turtles comprise an ecological radiation in which interspecific differences in trophic morphology have similarly been implicated as a pathway to ecopartition the marine realm, in both extant and extinct species. Because marine turtles are charismatic flagship species of conservation concern, their trophic ecology has been studied intensively using stable isotope analyses to gain insights into habitat use and diet, principally to inform conservation management. This legion of studies provides an unparalleled opportunity to examine ecological partitioning across numerous hierarchical levels that heretofore has not been applied to any other ecological radiation. Our contribution aims to provide a quantitative analysis of interspecific variation and a comprehensive review of intraspecific variation in trophic ecology across different hierarchical levels marshalling insights about realised trophic ecology derived from stable isotopes. We reviewed 113 stable isotope studies, mostly involving single species, and conducted a meta‐analysis of data from adults to elucidate differences in trophic ecology among species. Our study reveals a more intricate hierarchy of ecopartitioning by marine turtles than previously recognised based on trophic morphology and dietary analyses. We found strong statistical support for interspecific partitioning, as well as a continuum of intraspecific trophic sub‐specialisation in most species across several hierarchical levels. This ubiquity of trophic specialisation across many hierarchical levels exposes a far more complex view of trophic ecology and resource‐axis exploitation than suggested by species diversity alone. Not only do species segregate along many widely understood axes such as body size, macrohabitat, and trophic morphology but the general pattern revealed by isotopic studies is one of microhabitat segregation and variation in foraging behaviour within species, within populations, and among individuals. These findings are highly relevant to conservation management because they imply ecological non‐exchangeability, which introduces a new dimension beyond that of genetic stocks which drives current conservation planning. Perhaps the most remarkable finding from our data synthesis is that four of six marine turtle species forage across several trophic levels. This pattern is unlike that seen in other large marine predators, which forage at a single trophic level according to stable isotopes. This finding affirms suggestions that marine turtles are robust sentinels of ocean health and likely stabilise marine food webs. This insight has broader significance for studies of marine food webs and trophic ecology of large marine predators. Beyond insights concerning marine turtle ecology and conservation, our findings also have broader implications for the study of ecological radiations. Particularly, the unrecognised complexity of ecopartitioning beyond that predicted by trophic morphology suggests that this dominant approach in adaptive radiation research likely underestimates the degree of resource overlap and that interspecific disparities in trophic morphology may often over‐predict the degree of realised ecopartitioning. Hence, our findings suggest that stable isotopes can profitably be applied to study other ecological radiations and may reveal trophic variation beyond that reflected by trophic morphology.     

A micromorphological study of some representative genera in the tribe Saturejeae (Lamia ceae)

HUSAIN, S. Z., MARIN, P. D., ŠILIĆ, Č., QAISER, M. & PETCOVIĆ, B., 1990. A micromorphological study of some representative genera in the tribe Saturejeae (Lamiaceae). The Old World genera in the tribe Saturejeae are usually distributed either in Europe and North Africa or in the temperate parts of Asia. The centres of distribution of investigated genera are mainly in the Mediterranean region. In taxonomic revisions very little reference is made to micromorphological characters, in particular, to nutlets and leaf indumentum, in spite of the stability of these characters. Scanning electron microscopy of nutlet surface and patterns of leaf indumentum show a wide range of variation, not only among genera, but also at lower levels of classification. In view of this, nutlet surface and leaf indumentum structure, as seen with the SEM, of representative species of eight genera in the tribe Saturejeae provides useful additional character combinations in delimiting these closely related genera. This study also supports Boissier's delimitation of sections Micromeria and Pseudomelissa.     

Variation in anther and pollen morphology in Leucaena Benth. (Leguminosae-Mimosoideae)

Variation in anther and pollen morphology is described and illustrated for species of Leucaena and a selection of species from closely related genera. This is discussed in relation to the variation found in the closely related genera of the informal Leucaena, Dichrostachys and Xylia groups of the tribe Mimoseae. A number of characters for cladistic analysis are identified and putative homologies assessed. Preliminary taxonomic conclusions about the status and relationships of L. multicapitula are discussed. Mode of anther dehiscence is also described and illustrated. The observation of persistent intact tapetal membranes after anther dehiscence in some species of Leucaena is discussed in relation to pollen unit.     

Phylogeny and evolution of butterflies of the genus Parnassius: inferences from mitochondrial 16S and ND1 sequences

Phylogenetic relationships among species of the genus Parnassius and its related taxa were analyzed by comparing nucleotide sequences of mitochondrial 16S ribosomal RNA (504 sites) and NADH-dehydrogenase subunit 1 (469 sites). In the phylogenetic trees, Parnassius was found to be most closely related to Hypermnestra helios, whereas Archon apollinus, which has been classified in the tribe Parnassiini together with Parnassius and Hypermnestra, was more closely related to members of the tribe Zerynthiini. Within the Parnassius clade, six major clades corresponding to species groups were well supported, although the phylogenetic relationships among them were not clear. Although the results of the present study were in agreement with those of a previous phylogenetic study based on mitochondrial NADH-dehydrogenase subunit 5 sequences, our study strongly supported a close relationship between Parnassius and Hypermnestra, which was not well supported in the previous study.     

Chromosomal divergence and evolutionary inferences in Rhodniini based on the chromosomal location of ribosomal genes

In this study, we used fluorescence in situ hybridisation to determine the chromosomal location of 45S rDNA clusters in 10 species of the tribe Rhodniini (Hemiptera: Reduviidae: Triatominae). The results showed striking inter and intraspecific variability, with the location of the rDNA clusters restricted to sex chromosomes with two patterns: either on one (X chromosome) or both sex chromosomes (X and Y chromosomes). This variation occurs within a genus that has an unchanging diploid chromosome number (2n = 22, including 20 autosomes and 2 sex chromosomes) and a similar chromosome size and genomic DNA content, reflecting a genome dynamic not revealed by these chromosome traits. The rDNA variation in closely related species and the intraspecific polymorphism in Rhodnius ecuadoriensis suggested that the chromosomal position of rDNA clusters might be a useful marker to identify recently diverged species or populations. We discuss the ancestral position of ribosomal genes in the tribe Rhodniini and the possible mechanisms involved in the variation of the rDNA clusters, including the loss of rDNA loci on the Y chromosome, transposition and ectopic pairing. The last two processes involve chromosomal exchanges between both sex chromosomes, in contrast to the widely accepted idea that the achiasmatic sex chromosomes of Heteroptera do not interchange sequences.