Song diversification and complexity in canaries and seedeaters (Serinus spp.)

Bird song is a composite trait that shows great variation in syntax and phonology between taxa, which hampers the precise quantification of complexity. This should be more feasible for closely-related species that share aspects of song organization. We conducted a comparative study of song and syllable characteristics in the largest fringillid genus, Serinus (canaries and seedeaters), with two objectives: (1) to describe and quantify song complexity in the genus and (2) to assess evolutionary dynamics of song, in particular whether interspecific song diversification evolved along intraspecific lines of covariation between syllable characteristics. We document a pattern of intense, very labile evolution of song in the genus. Well-known vocal constraints such as the relationship between frequency and body size did not appear to limit this evolution, and intraspecific trade-offs between different aspects of syllable complexity did not constrain interspecific diversification. Factor analysis returned an axis of variation that comprehensively expresses song complexity in the genus, and is defined by song speed, repertoire size, and also various aspects of syllable phonology. The species body size is not related to this axis of complexity, but to a minor axis that may indicate better respiratory capacity of larger birds. We discuss the pattern of song evolution in Serinus spp. in the context of the evolutionary dynamics of sexually selected traits.  © 2007 The Linnean Society of London, Biological Journal of the Linnean Society , 2007, 92 , 183–194.     

Transitions in cuticular composition across a hybrid zone: historical accident or environmental adaptation?

Two subspecies of the grasshopper Chorthippus parallelus form a hybrid zone in the Pyrenees. Transitions across the zone, including changes in mating signals and reproductive isolation have been intensively studied. Cuticular pheromones have been identified as likely mate recognition signals. Since the major role of the cuticle is in waterproofing, environmental adaptation of cuticular composition has the interesting potential to generate assortative mating as an incidental by-product. We describe the pattern of variation in cuticular hydrocarbon blend in four transects through the hybrid zone. We find no evidence for a previously observed displaced cline in one blend component. There were differences between subspecies but these varied among transects and were small compared with variation between transects. We examined environmental variation within one transect and found a correlation between vegetation and cuticular composition, suggesting that environment influences the constitution of the cuticle, and hence natural selection may interact with mating signals in this species.  © 2003 The Linnean Society of London, Biological Journal of the Linnean Society , 2003, 78 , 193−201.     

Evolution of mosaic hybrid zone between invasive and endemic species of Cyprinidae through space and time

Hybridization phenomena can be considered in different ways. Hybridization studies can be used to elucidate some aspects of speciation and adaptation. However, hybridization may be a threat to endemic species. Here, we studied bidirectional introgression between two sympatric species of Cyprinidae: Chondrostoma toxostoma toxostoma and C. nasus nasus . Analysis of morphology, allozymes and mtDNA sequences revealed that the hybridization between these two cyprinids takes different forms in the same river. The mosaic hybrid zone (so called because of the absence of a simple cline) appeared to be partitioned due to the proportions of the two species along a spatial scale. The proportion of each hybrid group in this zone was unstable over a 5-year period. We propose various protection/conservation states for the C. t. toxostoma populations based on the level of introgressive hybridization and the genetic structure of both species in 'pure' populations.  © 2005 The Linnean Society of London, Biological Journal of the Linnean Society , 2005, 85 , 135–155.     

Bilateral Song Convergence in a Passerine Hybrid Zone: Genetics Contribute in One Species Only

Hybridization can drive the convergence of territorial and sexual signals. However, non-genetic processes such as competition, environment matching, or cultural transmission, also generate this pattern. We investigated the effect of hybridization on song convergence between two interspecifically territorial warblers in a moving hybrid zone. We confirmed song convergence in each species. Using an AFLP-based genetic index, we detected an effect of genetics on song convergence in Hippolais polyglotta, the expanding species. Evidence was weaker for H. icterina, the receding species. In moving zones, introgression is expected to be larger in the expanding species than in the receding. Thus, the asymmetric contribution of the genetic index to convergence was consistent with expectations for genetically determined traits in moving hybrid zones, and the observed introgression pattern of AFLP markers. However, the geographical location of individuals had an effect on song variation too when genetics was accounted for, suggesting that convergence also has non-genetic explanations. We examine the possible role of alternative processes to that of hybridization and discuss their conflicting effects on reinforcement and hybrid zone dynamics.     

Hybrids between Chorthippus brunneus and C. jacobsi (Orthoptera: Acrididae) do not show endogenous postzygotic isolation

Studies of prezygotic and postzygotic isolation in Drosophila have shown in general that species in sympatry tend to evolve prezygotic barriers earlier than do species in allopatry. However, postzygotic barriers tend to evolve at the same evolutionary rate in both sympatric and allopatric species. In contrast to these observations, the grasshoppers Chorthippus parallelus parallelus and C. p. erythropus show complete hybrid male sterility but only limited prezygotic isolation after an estimated 0.5 millions years of divergence. Like their congeners, C. brunneus and C. jacobsi form a hybrid zone where their ranges meet in northern Spain. However, the hybrid zone is mosaic and bimodal and, in contrast to the high levels of postzygotic isolation between C. parallelus subspecies, these two species showed no significant reduction in hybrid fitness in F₁ or backcross generations relative to the parental generations. The level of prezygotic isolation in laboratory tests was comparable to that between C. parallelus subspecies. These results suggest that endogenous postzygotic isolation does not play an important role in the reproductive isolation between C. brunneus and C. jacobsi , or in determining the structure of the hybrid zone. Exogenous postzygotic isolation may be present and should be tested in future studies.  © 2005 The Linnean Society of London, Biological Journal of the Linnean Society , 2005, 84 , 195–203.     

A new species of Valantia (Rubiaceae) from Spain

A new species of Valantia , V. lainzii Devesa & Ortega-Olivencia, endemic to the coastal zone of Granada (southern Spain) is described. The species recalls V. muralis L. in its general aspect, and V. deltoidea Brullo in the type of fructiferous body.  © 2003 The Linnean Society of London, Botanical Journal of the Linnean Society , 2003, 143 , 331–335.     

Historical influences on genital morphology among sympatric species: gonopod evolution and reproductive isolation in the crayfish genus Orconectes (Cambaridae)

Freshwater crayfishes represent an incredibly diverse component of temperate aquatic ecosystems. The genus Orconectes (Cambaridae) comprises approximately 17% of total global crayfish diversity and is native to central and eastern North America. Using both Bayesian and parsimony-based phylogenetic hypotheses from complete cytochrome c oxidase subunit I (COI) mtDNA sequence data (1545 bp) along with information on male gonopod shape and species distribution patterns, we examined the evolution of male genital morphology and its potential influence on species co-occurrence. We found that sympatric species exhibited similar male genital morphology more often than expected based on the frequency of genital shapes within the genus. When evolutionary history was incorporated into this analysis, the result was no longer significant, suggesting that species co-occurrence and the associated genital shapes of sympatric congeners cannot be explained solely by interspecific ecological interactions. Our results also do not support the current subgeneric classifications within Orconectes or the monophyly of the genus, both of which have been based previously on male genital morphology.  © 2006 The Linnean Society of London, Biological Journal of the Linnean Society , 2006, 89 , 1–12.     

Genetic and morphological differentiation patterns between sister species: the case of Onthophagus taurus and Onthophagus illyricus (Coleoptera, Scarabaeidae)

The present study investigated the morphological and genetic differentiation pattern between two sympatric dung beetle sister species, Onthophagus taurus and Onthophagus illyricus . The geometric morphometric approach coupled with the use of molecular markers allowed examination of the nature of interspecific relationships and an outline of the evolutionary and geographical processes that might have led to interspecific differentiation and the present-day partial sympatric and syntopic pattern of distribution. Geometric morphometrics failed to discrimininate the two species on the ground of external morphological traits, but revealed interspecific differences when the shape of the primary sexual traits was taken into account. The use of two different molecular markers (cytochrome oxidase subunit I gene and amplified fragment-length polymorphism) demonstrates that the two species are genetically well differentiated, forming two distinguishable lineages probably diverged during the Pliocene by allopatric speciation. No evidence of past or recent introgression and no support for hybridization were found, suggesting that sympatry was established subsequently, when speciation was accomplished. Both molecular markers and genital shape indicate that phenotypically intermediate individuals, despite their ambiguous appearance, are members of O. illyricus species rather than hybrids.  © 2006 The Linnean Society of London, Biological Journal of the Linnean Society , 2006, 89 , 197–211.     

Genome organization in diploid hybrid species of Argyranthemum (Asteraceae) in the Canary Islands

Two diploid species of hybrid origin, Argyranthemum lemsii and A. sundingii , have been described from different valleys in the Anaga peninsula, north-east Tenerife. They have previously been shown to originate from hybridization between the same parental species, the montane A. broussonetii and the coastal A. frutescens , A. broussonetii being the chloroplast donor in one valley and A. frutescens in the other. The specific status of the two hybrid species has been questioned. In this study we used karyotype analysis, FISH, and GISH to address the question of multiple diploid hybrid speciation. GISH did not discriminate clearly between the parental genomes, but differential labelling was observed in separate hybrid populations, indicating different chromosomal rearrangements in different valleys. Small karyotype differences and local loss of rDNA were also observed. Thus separate origins of the same hybrid combination in different valleys in Tenerife have been verified. Our results add some support to the recognition of two species of hybrid origin, but the case serves to illustrate some of the many problems connected with the species concept in plants.  © 2003 The Linnean Society of London, Botanical Journal of the Linnean Society , 2003, 141 , 491–501.     

A narrow hybrid zone between the grasshoppers Stenobothrus clavatus and Stenobothrus rubicundus: courtship song analysis

The closely related grasshopper species Stenobothrus rubicundus and Stenobothrus clavatus are known to hybridize in a very narrow contact zone on Mt. Tomaros in northern Greece. These species produce very different and complex courtship songs accompanied with visual display. We analyzed the courtship songs and underlying stridulatory movements of the hind legs in natural hybrids from Mount Tomaros. The two species were also hybridized in the laboratory and their songs were compared with the songs of the natural hybrids. Some hybrid songs were shown to have intermediate features between parental songs, whereas other hybrid songs comprised completely new elements. The clavatus‐like song elements were found to dominate in hybrid songs. These song features may influence the mating success of hybrid males in the contact zone. A comparison of hybrid songs with the song pattern of the north European S. rubicundus populations allowed us to suggest a scenario of S. rubicundus and S. clavatus origin. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, ?? , ??–??.     

A new species of Hybanthus (Violaceae) from north-eastern Brazil

A new species of Hybanthus (Violaceae) from north-eastern Brazil is described and illustrated. Hybanthus verrucosus Paula-Souza is endemic to 'caatingas' of Bahia and Alagoas states and is very distinct from all other Brazilian species of the genus, for its anterior petal, which is covered with small wart-like outgrowths. Other distinctive characteristics of the new species and its relationship with a closely related species ( H. atropurpureus (A.St.-Hil.) Taub.) are discussed.  © 2003 The Linnean Society of London, Botanical Journal of the Linnean Society , 2003, 141 , 503–506.     

Multiple barriers to gene exchange in a field cricket hybrid zone

Data on patterns of variation within hybrid zones, combined with studies of life history, mate choice, and hybrid performance, allow estimates of the contribution of different pre-zygotic and post-zygotic barriers to reproductive isolation. We examine the role of behavioural barriers to gene exchange in the maintenance of a hybrid zone between North American field crickets Gryllus firmus and Gryllus pennsylvanicus . We consider these barriers in the context of previous studies that documented temporal and ecological isolation and a one-way post-mating incompatibility (i.e. G. firmus females do not produce offspring when they mate only with heterospecific males). Based on no-choice mating experiments in the laboratory, we demonstrate strong behavioural pre-mating barriers between the two species, but no apparent fecundity or fertility costs for G. firmus females when they mate with both conspecific and heterospecific males. Furthermore, we show that G. firmus females do not discriminate between hybrids and conspecifics, whereas G. pennsylvanicus females do. This observation could explain the asymmetric allele introgression observed in the hybrid zone. We also document a failure of heterospecific males to induce normal oviposition in G. firmus females, which may be due to rapid evolution of accessory gland proteins and may serve as an additional barrier to gene exchange.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 97 , 390–402.     

Hybridization patterns and the evolution of reproductive isolation in ducks

Much of our knowledge of the evolution of reproductive isolation comes from studies of Drosophila . This body of work has revealed the following patterns: (1) reproductive isolation increases with phylogenetic distance between hybridizing species; (2) reproductive isolation is greater between sympatric than allopatric species with the same level of divergence; and (3) hybrid crosses conform to Haldane's rule. We tested for the existence of these patterns in ducks (subfamily Anatinae, sensu Livezey, 1997b ) based on 1037 hybrids of known parentage. Our analyses of the number of interspecific crosses in relation to phylogenetic distance found a significant deviation between the observed and expected distribution of crosses controlling for the topology of the Anatinae phylogeny. In particular, we found both an excess of hybrid crosses among closely related species and a scarcity among distantly related species. The number of hybrid males also decreased with increasing phylogenetic distance between parental species, although the number of hybrid females remained low and constant. Sympatric species produced higher numbers of hybrid males than allopatric ones, despite no difference in phylogenetic distance among parental species in compared groups. The number of hybrid males exceeded the number of hybrid females, consistent with Haldane's rule. This was evident even though the analysis was restricted to a reduced set of phylogenetically independent crosses. However, the pattern was no longer significant after correction for the number of hybrid males by the male-biased sex ratio of adult ducks.  © 2002 The Linnean Society of London, Biological Journal of the Linnean Society , 2002, 77 , 193−200.     

Karyotype evolution in the genus Boronia (Rutaceae)

New chromosome counts are reported for Boronia clavata 2 n  = 14, B. heterophylla 'Near White' 2 n  = 15, B.  'Carousel' 2 n  = 16, B. deanei 2 n  = 22, B. chartacea 2 n  = 32, B. keysii 2 n  = 32, B. pilosa 2 n  = 44, B. anethifolia 2 n  = 36 and B. citriodora 2 n  = 108. Studies in 20 genotypes of 18 species and one interspecific hybrid revealed that they are highly complex in terms of chromosome number, ploidy level, chromosomal length, karyotype constitution and asymmetry. Karyotype analysis indicated that Boronia taxa with high chromosome numbers are primitive and those with lower numbers are derived. The basic chromosome number for this genus is suggested to be x = 18. Analysis of chromosome number, variations of total chromosome length (TCL) and average chromosome length (ACL), Nombre Fondamental (NF) and karyotype asymmetry suggest that dysploid reduction is the major mechanism in Boronia karyotype evolution. Chromosomal rearrangements might also have been involved. Origin, chromosome number changes and spread of Boronia are discussed in relation to the species divergence and the geological and climatic changes of the Australian continent.  © 2003 The Linnean Society of London, Botanical Journal of the Linnean Society , 2003, 142 , 309–320.     

A new diploid Hieracium (Asteraceae: Lactuceae) from Bulgaria

Hieracium kittanae Vladimirov, a new diploid (2 n  = 18) species restricted to a few gorges in the Central Rhodope Mountains, south Bulgaria, is described and illustrated. It occurs in crevices of limestone rock in shady to sunny places with high atmospheric humidity. Morphologically, it to some extent resembles taxa from the H. glaucinum and H. schmidtii groups, but is otherwise very distinct, without close relatives in the Rhodope and neighbouring mountains.  © 2003 The Linnean Society of London, Botanical Journal of the Linnean Society, 2003 , 143 , 213−218.     

Courtship song and immune function in the field cricket Gryllus bimaculatus

It has been assumed that sexual ornaments have evolved to reveal males' health and vigour for females. Choosy females may indirectly use ornaments as an indicator of the presence and effectiveness of genes for resistance against parasites. In this study we tested whether females of the Mediterranean field cricket, Gryllus bimaculatus, can use courtship song as a cue for choosing males with high immunocompetence, measured as encapsulation rate of nylon implants and lytic activity of haemolymph. We found that female crickets preferred courtship songs from males with a high encapsulation rate. Female crickets also had a tendency to prefer courtship songs with high tick rate and long high-frequency tick duration. These preferred song components were positively correlated with encapsulation rate, but negatively correlated with lytic activity of the male. In contrast to previous studies of crickets, there was no correlation between male weight and encapsulation rate or lytic activity. There is some evidence in another cricket species that the ability to encapsulate pathogens is heritable. Thus, in light of this study it seems possible that by preferring males according to their courtship song, females might benefit by increasing the parasite resistance of their offspring.  © 2003 The Linnean Society of London, Biological Journal of the Linnean Society , 2003, 79 , 503–510.     

Voice variance may signify ongoing divergence among black-legged kittiwake populations

Acoustic features are important for individual and species recognition. However, while dialectal variations in song characteristics have been described in many songbirds, geographical divergence in vocal features across populations has seldom been studied in birds that are not thought to have song-learning abilities. Here, we document marked differences in the vocal structure of calls of two populations of black-legged kittiwakes ( Rissa tridactyla ), a seabird whose call is considered as not being learned from other individuals. We found that calls vary both within and between populations. Within-population variation may convey individual identity, whereas the marked differences in frequency and temporal parameters observed between the two populations may reveal ongoing divergence among kittiwake populations. Moreover, we were unable to detect any sex signature in adult calls in a Pacific population (Middleton, Alaska), while these were detected in an Atlantic population (Hornøya, Norway), potentially affecting sexual behaviours. Despite the fact that these calls seemed to change over the reproductive season and across years, the individual signature remained fairly stable. Such vocal differences suggest that Pacific and Atlantic populations may be undergoing behavioural divergences that may reveal early stages of speciation, as is suggested by molecular data.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 97 , 289–297.     

Hybrid origin of the imperial pheasant Lophura imperialis (Delacour and Jabouille, 1924) demonstrated by morphology, hybrid experiments, and DNA analyses

The imperial pheasant Lophura imperialis was described in 1924 from a captive pair that was obtained in Vietnam, and that became the sole founders of a captive line in France. Always considered a highly endangered and mysterious species, and despite concerted searches, L. imperialis was not found again in the wild until one was trapped in 1990, and the captive population gradually died out. Its status as a distinct species was unquestioned until the late 1990s when the possibility of a hybrid origin was raised. To elucidate the taxonomic status of L. imperialis , we studied all the existing museum specimens, carried out captive hybridization experiments, and analysed mitochondrial DNA and microsatellites. All these lines of evidence demonstrate congruently and conclusively that L. imperialis is an occasional hybrid between silver pheasant L. nycthemera and Edwards's pheasant L. edwardsi, with the 1990 bird probably being a hybrid between L. nycthemera and Vietnamese pheasant L. hatinhensis. Thus L. imperialis has no taxonomic standing and should be removed from lists of species of conservation concern. However, hybridization with L. nycthemera may pose a further threat to the survival in the wild of the endangered L. edwardsi and L. hatinhensis .  © 2003 The Linnean Society of London, Biological Journal of the Linnean Society , 2003, 80 , 573–600.     

Evidence for ecological speciation in the sister species Candidula unifasciata (Poiret, 1801) and C. rugosiuscula (Michaud, 1831) (Helicellinae, Gastropoda)

This paper studies the speciation of two land snail species in south-east France. By using two mitochondrial and one nuclear gene, the species are shown to be sister species within the larger clade of western European Candidula species. The species never occurred syntopically, but a narrow contact zone population was identified. Recent range expansions inferred from phylogeographical methods indicated that the present day distributions are not limited by the colonization capacity of the species. Analysis of environmental variables suggested distribution along an ecotone. The gastropod community is also correlated to this gradient. Morphometric shell analysis showed that the divergence between the sister species is a result of desiccation-resistant shell characters. We therefore conclude that the cause for the ecological speciation is most probably the establishment of the Mediterranean climate ≈ 3.2 mya. A model of ecological speciation with a moving ecotone is put forward.  © 2003 The Linnean Society of London, Biological Journal of the Linnean Society , 2003, 79, 611–628.