Male advertisement call characters as phylogeographical indicators in European water frogs

Anuran vocalizations are valuable in the determination of species identity and have the potential to discriminate intraspecific variation. We developed novel bioacoustic sampling techniques, based on Fast Fourier Transforms, to increase the precision and sensitivity of male advertisement call analysis and applied the method to European water frogs. This approach robustly separated the three types of north European water frogs (Rana ridibunda, R. lessonae and their viable, fertile hybrid R. esculenta) by their call subunit characteristics. The hybrid frog exhibited a high frequency call component absent from both the parental species. Furthermore, call analysis demonstrated significant intraspecific differences among populations of all three frog types. Call characters of R. ridibunda changed systematically as a function of longitude. This trend may reflect either clinal variation in selection pressures across Europe, or the consequences of drift following postglacial colonization from eastern refugia. High resolution vocalization analysis therefore provides a potentially useful method for investigating intraspecific differentiation and the phylogeographical origins of anuran distributions. © 2002 The Linnean Society of London. Biological Journal of the Linnean Society, 2002, 77 , 355?365.     

Molecular systematics of mantelline frogs from Madagascar and the evolution of their femoral glands

Several genera of frogs from Madagascar, classified in the family Mantellidae, subfamily Mantellinae, possess structures commonly called 'femoral glands' on the ventral side of their shanks. The question arises as to the origin and phylogenetic significance of these glands. A molecular phylogeny based on 3601 nucleotide DNA sequences of three mitochondrial and two nuclear genes of 30 mantellid species provided strong support for monophyly of the included mantellines, all characterized by enlarged femoral gland clusters, as well as for those with gland clusters of coordinated central arrangement of secretion ducts. However, the phylogeny also strongly supported the hypothesis of convergent evolution of structurally similar glands in unrelated frogs ( Indirana , Petropedetes ), and several trends of convergent evolution of gland structure within mantellines. We studied the light microscopic structure of the femoral glands in a representative array of 18 mantellid species. Males of all species of the subfamily Mantellinae were characterized by clusters of distinct single glands. Each was structurally similar to an enlarged granular gland and secreted independently, probably through a single duct. By contrast, the largely semi-aquatic frogs in the genus Mantidactylus had a specialized cluster of glands, in which the secretion ducts led into a macroscopically recognizable central depression. In Boophis opisthodon , a mantellid species of the subfamily Boophinae without externally recognizable femoral glands, we observed a large number of enlarged granular glands of various sizes in the ventral skin of the shank. This observation is consistent with the hypothesis that the large and more uniform organs of mantellines are derived granular glands.  © 2007 The Linnean Society of London, Biological Journal of the Linnean Society , 2007, 92 , 529–539.     

Phylogenetic patterns of diversification in a clade of Neotropical frogs (Anura: Aromobatidae: Mannophryne)

We used partial sequences of mitochondrial 16S and cytochrome oxidase I genes to perform a phylogenetic study of collared frogs (Anura: Aromobatidae: Mannophryne ), a genus endemic to Venezuela and the islands of Trinidad and Tobago. We analysed 1.2 kb from 13 of the 15 described species of Mannophryne . Maximum parsimony, maximum likelihood and Bayesian analyses support the monophyly of Mannophryne . Mannophryne consists of three deeply differentiated clades that split from each other in a relatively short period of time. The diversification of Mannophryne occurred well before the glacial-interglacial periods of the Quaternary. Our data support the taxonomic validity of M. olmonae , a species endemic to Tobago Island. Mannophryne olmonae is more closely related to the continental species Mannophryne riveroi than to the Trinidad island endemic Mannophryne trinitatis . As in most tropical clades of frogs, molecular evidence indicates that species richness in Mannophryne is largely underestimated and, consequently, current priorities for conservation are inadequate.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 97 , 185–199.     

Glass frogs (Centrolenidae) of Yanayacu Biological Station, Ecuador, with the description of a new species and comments on centrolenid systematics

We describe Centrolene bacatum , C. buckleyi , Cochranella posadae , and a new species of Cochranella from Yanayacu Biological Station on the Amazonian slopes of the Ecuadorian Andes. The new species differs from other species in Centrolenidae by a combination of characters, including reduced webbing between Fingers III and IV, and kidneys covered with white peritoneum. We summarize the current generic and infrageneric classification in Centrolenidae and discuss some of its problems. A phylogenetic analysis of morphological and behavioural data shows that the genera Centrolene and Cochranella might not be monophyletic; the genus Hyalinobatrachium and, in particular, the group H. fleischmanni seem to be monophyletic. However, an analysis with many more characters is needed to resolve the relationships of glass frogs.  © 2006 The Linnean Society of London, Zoological Journal of the Linnean Society , 2006, 147 , 489–513.     

Multiple origins of invasive and ‘native’ water frogs (Pelophylax spp.) in Switzerland

The marsh frog (Pelophylax ridibundus) has been introduced in many areas in Central and Western Europe as a result of commercial trade with Eastern Europe, and is rapidly replacing the native pool frog (P. lessonae). A large number of Pelophylax species are distributed in Eastern Europe and the strong phenotypic similarity between these species is rendering their identification hazardous. Consequently, alien populations of Pelophylax might not strictly be composed of P. ridibundus as previously suspected. In the present study, we analysed the cytochrome‐b and NADH dehydrogenase subunit 3 genes of introduced and native Pelophylax species from Switzerland (299 individuals) in order to properly identify the source populations of the invaders and the genetic status of the native species. Our study highlighted the occurrence of several genetic lineages of invasive frogs in western Switzerland. Unexpectedly, we also showed that several populations of the native pool frog (P. lessonae) cluster with the Italian pool frog P. bergeri from central Italy (considered by some authors as a subspecies of P. lessonae). Hence, these populations are probably also the result of introductions, meaning that the number of native P. lessonae populations is fewer than expected in Switzerland. These findings have important implications concerning the conservation of the endemic pool frog populations, as the presence of multiple alien species could strongly affect their long‐term subsistence. © 2014 The Linnean Society of London, Biological Journal of the Linnean Society, 2014, 112 , 442–449.     

Microanatomy of the amniote femur and inference of lifestyle in limbed vertebrates

The femoral microanatomy of 155 species of extant amniotes (57 species of mammals, 15 species of turtles, 56 species of lepidosaurs, and 27 species of birds) of known lifestyle is studied to demonstrate a possible link between some basic parameters of bone structure and specific lifestyles, as well as phylogenetic relationships between taxa. Squared change parsimony with random taxon reshuffling and pairwise comparisons reveal that most compactness and size parameters exhibit both phylogenetic and ecological signals. A discriminant analysis produces several inference models, including a ternary model (aquatic, amphibious, terrestrial) that yield the correct lifestyle in 88% of the cases. These models are used to infer the lifestyle of three extinct Permian temnospondyls: Eryops megacephalus, Acheloma dunni, and Trimerorhachis insignis. © 2013 The Linnean Society of London, Biological Journal of the Linnean Society, 2013, 109 , 644–655.     

The vocal sac of Hylodidae (Amphibia,Anura): Phylogenetic and functional implications of a unique morphology

Anuran vocal sacs are elastic chambers that recycle exhaled air during vocalizations and are present in males of most species of frogs. Most knowledge of the diversity of vocal sacs relates to external morphology; detailed information on internal anatomy is available for few groups of frogs. Frogs of the family Hylodidae, which is endemic to the Atlantic Forest of Brazil and adjacent Argentina and Paraguay, have three patterns of vocal sac morphology—that is, single, subgular; paired, lateral; and absent. The submandibular musculature and structure of the vocal sac mucosa (the internal wall of the vocal sac) of exemplar species of this family and relatives were studied. In contrast to previous accounts, we found that all species of Crossodactylus and Hylodes possess paired, lateral vocal sacs, with the internal mucosa of each sac being separate from the contralateral one. Unlike all other frogs for which data are available, the mucosa of the vocal sacs in these genera is not supported externally by the mm. intermandibularis and interhyoideus . Rather, the vocal sac mucosa projects through the musculature and is free in the submandibular lymphatic sac. The presence of paired, lateral vocal sacs, the internal separation of the sac mucosae, and their projection through the m. interhyoideus are synapomorphies of the family. Furthermore, the specific configuration of the m. interhyoideus allows asymmetric inflation of paired vocal sacs, a feature only reported in species of these diurnal, stream‐dwelling frogs.     

Ultraviolet and carotenoid‐based coloration in the viviparous lizard Zootoca vivipara (Squamata: Lacertidae) in relation to age,sex, and morphology

Lizards display structural and pigment‐based colorations, and their visual system is sensitive to wavelengths of 300–700 nm. However, few studies in squamate reptiles have quantified interindividual colour variation that includes the structural ultraviolet (UV) component (300–400 nm). In the present study, we investigated variability of a ventral UV/yellow–red ornamentation in the common lizard Zootoca vivipara, including an analysis of spatial distribution, as well as sex and age differences. We also investigated whether the expression of coloration is related to body size and condition. Our analyses revealed two distinct patches: a gular patch with a strong UV reflectance and a belly patch with a dominant yellow–red reflectance. Males displayed a less saturated throat coloration with higher UV chroma and UV hue, and had a redder but duller belly coloration than females. Yearlings had less elaborate ornaments than adults, although they already displayed a yellow–red sexual dichromatism on the belly. UV sexual dichromatism was only apparent in adults as a result of a weaker UV reflectance in females, suggesting potential fitness costs of a bright UV coloration in that sex. Different colour traits were related to body size in both sexes, as well as to body condition in males. We discuss the potential evolutionary scenarios leading to the maintenance of this ornament in common lizards. © 2013 The Linnean Society of London, Biological Journal of the Linnean Society, 2013, 110 , 128–141.     

Female frog auditory sensitivity,male calls,and background noise: potential influences on the evolution of a peculiar matched filter

The matched filter hypothesis proposes that the tuning of females' auditory sensitivity matches the spectral energy distribution of males' signals. Such correspondence is expected to arise over evolutionary time, as it promotes conspecific information transfer and reduces interference from other sound sources. Our main objective was to determine the correspondence between the acoustic sensitivity of female frogs of Eupsophus roseus and the spectral characteristics of advertisement vocalizations produced by conspecific males. We also aimed to determine how auditory sensitivity is related to the characteristics of background noise. We analysed data on the auditory sensitivity of E. roseus females, and recordings of conspecific male vocalizations and of the acoustic environment during the breeding period of this species. Our results indicate a concordance between the auditory sensitivity of females and call spectra that would provide an appropriate detection of these signals. In addition, this matching is large relative to the correspondence between auditory sensitivity with the spectra of the abiotic and biotic background noise, with the last component being associated with calls of the related species Eupsophus vertebralis. This may be an adaptation of receivers confronting sound interference, which improves the capability of E. roseus to communicate sexually by means of acoustic signals. © 2013 The Linnean Society of London, Biological Journal of the Linnean Society, 2013, 110 , 814–827.     

Dispersal,fragmentation, and isolation shape the phylogeography of the European lineages of Polyommatus (Agrodiaetus) ripartii (Lepidoptera: Lycaenidae)

Polyommatus ripartii is a biogeographically and taxonomically poorly understood species of butterfly with a scattered distribution in Europe. Recently, it has been shown that this species includes several European endemic and localized taxa (galloi, exuberans, agenjoi) that were previously considered species and even protected, a result that poses further questions about the processes that led to its current distribution. We analysed mitochondrial DNA and the morphology of P. ripartii specimens to study the phylogeography of European populations. Three genetically differentiated but apparently synmorphic lineages occur in Europe that could be considered evolutionarily significant units for conservation. Their strongly fragmented and counterintuitive distribution seems to be the result of multiple range expansions and contractions along Pleistocene climatic oscillations. Remarkably, based on the 79 specimens studied, these genetic lineages do not seem to extensively coexist in the distributional mosaic, a phenomenon most evident in the Iberian Peninsula. One of the important gaps in the European distribution of P. ripartii is reduced by the discovery of new Croatian populations, which also facilitate a better understanding of the biogeography of the species. © 2013 The Linnean Society of London, Biological Journal of the Linnean Society, 2013, 109 , 817‐829.     

Rampant polyphyly indicates cryptic diversity in a clade of Neotropical flycatchers (Aves: Tyrannidae)

Polyphyletic arrangements in DNA phylogenies are often indicators of cryptic species diversity masked by erroneous taxonomic treatments that are frequently based on morphological data. Although mitochondrial (mt)DNA polyphyly is detected relatively rarely in phylogenetic studies, it has recently been found in a variety of tyrant‐flycatcher (Tyrannidae) groups. In the present study, we provide a DNA phylogeny for a mitochondrial and a nuclear locus with a complete species sampling in Zimmerius flycatchers, showing that the genus is characterized by multiple mtDNA polyphyly. Based on phylogenetic and life‐history information, we suggest the elevation of a number of taxa to species status, leading to a doubling of Zimmerius species‐level diversity compared to taxonomic treatments conducted before 2001. © 2013 The Linnean Society of London, Biological Journal of the Linnean Society, 2013, ●●, ●●–●●.     

Phylogenetic systematics of the Rana signata complex of Philippine and Bornean stream frogs: reconsideration of Huxley's modification of Wallace's Line at the Oriental–Australian faunal zone interface

We addressed the evolutionary relationships and biogeographical patterns of a model organism of low relative dispersal ability by electrophoretically assaying the products of 42 presumptive gene loci in Philippine and Bornean members of the Rana signata complex of SE Asian stream frogs. Utilizing three distantly related species of ranid frogs to deeply root trees consisting of five more closely‐related species and six in‐group species of the Rana signata complex, we conducted phylogenetic analyses that produced concordant topologies, regardless of the data coding strategy employed. All analyses support the hypothesis of monophyly for the Rana signata complex on the whole, but none provides support for the monophyly of its Philippine members. Our analyses of morphometric and allozyme data (along with biogeographical information) indicate that (1) most previously‐recognized Philippine and Bornean subspecies of the Rana signata complex should be recognized as full species in appreciation of their status as independent evolutionary lineages; (2) Rana picturata Boulenger (until very recently included in the synonymy of Rana signata signata) is deserving of specific rank; (3) the Mindoro Isl. (Philippine) population, previously confused with Rana signata similis of Luzon Isl. is a new species; (4) two major clades (((R. signata (R. grandocula + R. similis)) + (R. picturata (R. mangyanum + R. moellendorffi))) of Bornean + Philippine lineages are recognized, corresponding to two separate faunal exchanges between the Philippines and the edge of the Sunda Shelf; (5) invasions of the oceanic portions of the Philippine islands from the Sunda Shelf have occurred along both the eastern (Sulus–Mindanao–Samar–Leyte–Luzon) arc and the western (Palawan–Busuanga–Mindoro) island arcs; (6) northern reaches of Wallace’s Line (as modified by Huxley) include exceptions to an otherwise discrete faunal separation. These results suggest the need for revision of this biogeographical barrier, increased recognition of temporal patterns of island connectedness and geographical proximity, and/or a greater appreciation of dispersal abilities of ranid frogs. © 2002 The Linnean Society of London, Biological Journal of the Linnean Society, 2002, 76 , 393–461.     

Structure and function of defensive glands in soldiers of Glossotermes oculatus (Isoptera: Serritermitidae)

The soldier caste represents the most conspicuous realization of termite eusociality, characterized by an extreme anatomical, behavioural, and physiological specialization. Numerous strategies have evolved in soldiers, including extreme adaptations such as self‐sacrifice by autothysis. In the present study, we investigated the structure and function of defensive glands in Glossotermes oculatus soldiers aiming to understand their use in combat. Three glands are involved in defence: labral, frontal, and labial glands. Mandibles are used to bite the enemy, whereas the secretions of labral and labial glands are discharged into the wound. A striking characteristic of G. oculatus is the lack of the frontal pore; the secretion of the frontal gland is discharged by a rupture of the body wall. We hypothesized that this self‐sacrifice is an efficient way of blocking a gallery under attack. A similar development of the frontal gland occurs in Serritermes serrifer, which supports the close relationship between the two genera inferred from morphological and genetic analyses. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 99 , 839–848.     

Circulating nucleic acids: possible inherited effects

The presence of circulating nucleic acids in man, animals and plants is well documented. It is clear that such nucleic acids can not only circulate freely within an organism, but can also enter cells when their biology may be changed either epigenetically or genetically. Evidence is presented concerning a possible influence of these nucleic acid fragments on the genetics of the F1 generation of man, animals and plants. The data presented also offer a mechanism by which the incorporation of horizontally transferred genes between organisms may be achieved. The role that circulating nucleic acids might play in modifying the F1 generation and possibly the evolutionary process is considered. © 2013 The Linnean Society of London, Biological Journal of the Linnean Society, 2013, 110 , 931–948.     

Evolution of venom delivery structures in madtom catfishes (Siluriformes: Ictaluridae)

The use of venom to subdue prey or deter predators has evolved multiple times in numerous animal lineages. Catfishes represent one of the most easily recognized, but least studied groups of venomous fishes. Venom glands surround spines on the dorsal and pectoral fins that serve as venom delivery structures. Species of madtom catfishes in the genus Noturus were found to each have one of four venom delivery morphologies: (1) smooth spine with no venom gland; (2) smooth spine with venom gland associated with shaft of spine; (3) serrated spine with venom gland associated with shaft of spine; and (4) serrated spine with venom gland associated with shaft of spine and posterior serrations. Analyses accounting for the phylogenetic history of Noturus species suggest that a serrated pectoral spine with a venom gland is the ancestral condition for the genus. The presence of serrations and a venom gland have been largely conserved among Noturus species, but sting morphology has changed at least five times within the genus. Four of these changes have resulted in a loss of morphological complexity, including the loss of posterior serrations, loss of venom glands associated with the posterior serrations, and one complete loss of the venom gland. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 102 , 115–129.     

Evolutionary significance of oral morphology in the carnivorous tadpoles of tiger frogs, genus Hoplobatrachus (Ranidae)

We describe the larval stages of three species of the Asian-African tiger frogs Hoplobatrachus chinensis, H. occipitalis and H. tigerinus . The tadpoles of all three species are very similar, with peculiar oral features: (1) double rows of needle-like labial teeth, (2) strong emarginations on the large jaw sheaths and (3) keratinized spurs on the buccal floor. Characters 1 and 2 (and perhaps 3) are probably related to the carnivorous habits of these tadpoles. A molecular phylogeny based on 2430 base pairs of two nuclear and four mitochondrial genes corroborated monophyly of Asian and African Hoplobatrachus, and identified Euphlyctis as their sister group. Tadpoles of the latter genus lack buccal spurs and double labial tooth rows but share large jaw sheaths, the upper with a medial projection. Therefore, the common ancestor of Euphlyctis and Hoplobatrachus probably was also characterized by this state, and may have been facultatively carnivorous. Further carnivorous specializations in Hoplobatrachus could explain why tiger frogs have been so successful in populating arid environments where ponds are at high risk of desiccation. Larval morphology may prove to be the key innovation which enabled them to disperse, in the Late Cenozoic, into their current very wide distribution area in Asia and Africa.  © 2004 The Linnean Society of London, Biological Journal of the Linnean Society , 2004, 81 , 171–181.     

The interplay between multiple predators and prey colour divergence

Evolutionary divergence in the coloration of toxic prey is expected when geographic variation in predator composition and behavior favours shifts in prey conspicuousness. A fundamental prediction of predator‐driven colour divergence is that the local coloration should experience lower predation risk than novel prey phenotypes. The dorsal coloration of the granular poison frog varies gradually from populations of conspicuous bright red frogs to populations of dull green and relatively cryptic frogs. We conducted experiments with clay models in four populations to examine the geographic patterns of taxon‐specific predation. Birds avoided the local phenotype while lizards consistently selected for decreased conspicuousness and crab predation did not depend on frog coloration. Importantly, birds and lizards favoured low conspicuousness in populations where relatively cryptic green morphs have evolved. This study provides evidence for the interplay among distinct selective pressures, from multiple‐predator taxa, acting on the divergence in protective coloration of prey species. © 2014 The Linnean Society of London, Biological Journal of the Linnean Society, 2014, 113 , 580–589.