Charting the course of reed‐warblers across the Pacific islands

Aim Deciphering the complex colonization history of island archipelagos is greatly facilitated by comprehensive phylogenies. In this study we investigate the phylogeny and biogeography of the insular reed‐warblers (genus Acrocephalus) of the tropical Pacific Ocean, from Australia to eastern Polynesia. Location Oceania. Methods We used sequences of mitochondrial DNA (cytochrome b, ND2 and ATP8 genes) to infer the colonization patterns of reed‐warblers endemic to Pacific islands and Australia. We sampled all known taxa of Acrocephalus in the Pacific except A. luscinius nijoi, for which no sample was available. Most taxa were represented by toe‐pad samples from museum specimens collected in the 19th and 20th centuries. With a few exceptions, several specimens per taxon were sequenced independently in two institutions (Smithsonian Institution and Natural History Museum of Geneva). Results Our data indicate that Pacific reed‐warblers do not form a monophyletic group, because A. luscinius luscinius from Guam falls outside the main Pacific radiation. The remaining Pacific taxa are divided into two clades: one clade includes all the reed‐warblers from Micronesia (except Guam) and Australia, and two Polynesian taxa from the Line Islands and the southern Marquesas; the other clade includes all remaining Polynesian taxa. The taxa endemic to three archipelagos (Mariana, Marquesas and Society islands) are polyphyletic, suggesting several independent colonizations. Main conclusions Our results provide evidence for a complex pattern of colonization of the Pacific by reed‐warblers. Calibration analyses suggest that reed‐warbler lineages are much younger than the ages of the islands they occupy. Several remote archipelagos were colonized independently more than once. Consequently, we infer that the colonization of reed‐warblers in the Pacific did not follow a regular, stepping‐stone‐like pattern. The phylogeny also suggests a previously undetected case of reverse colonization (from island to continent) for the Australian lineage and indicates that A. luscinius, as currently defined, is not monophyletic. We discuss the supertramp strategy of reed‐warblers in the Pacific and show that, although Pacific reed‐warblers meet some of the supertramp criteria in their aptitude for colonizing remote archipelagos, their life history characteristics do not fit the model.     

Confirmation of Acrocephalus scirpaceus avicenniae (Aves: Acrocephalidae) from mangroves on the Red Sea coast near Jazan,southwest Saudi Arabia

Members of the Eurasian (or Common) Reed Warbler complex, Acrocephalus scirpaceus, are widespread across much of Europe, Africa, the Middle East and Central Asia. With its relatively complex taxonomy, the identity of several local (and sometimes remote) populations remains somewhat unresolved. In Saudi Arabia, populations of reed warblers were first identified in mangroves at Yanbu’ on the Red Sea coast in 1984, with several subsequent records up to 900 km further south toward the Yemen border. We took morphological data from 51 individuals and genetic material from three individuals captured near Jazan in southwest Saudi Arabia. Both genetic and morphometric data confirmed that these birds belong to the taxon A. scirpaceus avicenniae, sometimes referred to as the Mangrove or Red Sea Reed Warbler.     

The phytogeographical affinities of the Pitcairn Islands – a model for south‐eastern Polynesia?

Aim To identify how the Pitcairn group relates biogeographically to the south‐eastern Polynesian region and if, as a subset of the regions flora, it can then be used as a model for biogeographical analyses. Location The Pitcairn group (25°4′ S, 130°06′ W) comprises four islands: Pitcairn, a relatively young, high volcanic Island; Henderson, an uplifted atoll, the uplift caused by the eruption of Pitcairn; and two atolls, Ducie and Oeno. The remote location, young age and range of island types found in the Pitcairn Island group makes the group ideal for the study of island biogeography and evolution. Methods A detailed literature survey was carried out and several data sets were compiled. Dispersal method, propagule number and range data were collected for each of the 114 species that occurs in the Pitcairn group, and environmental data was also gathered for islands in Polynesia. Analyses were carried out using non‐metric multidimensional scaling and clustering techniques. Results The flora of the Pitcairn Islands is derived from the flora of other island groups in the south‐eastern Polynesian region, notably those of the Austral, Society and Cook Islands. Species with a Pacific‐wide distribution dominate the overall Pitcairn group flora. However, each of the islands show different patterns; Pitcairn is dominated by species with Pacific, Polynesian and endemic distributions, with anemochory as the dominant dispersal method (39.5%); Henderson is also dominated by species with Pacific, Polynesian and endemic distributions, but zoochory is the dominant dispersal method (59.4); Oeno and Ducie are dominated by Pantropic species with hydrochory as the most common dispersal method (52.9% and 100%, respectively). Main conclusions ? Habitat availability is the most significant factor determining the composition and size of the flora. ? South‐east Polynesia is a valid biogeographical unit, and should include the Cook, Austral, Society, Marquesas, Gambier, Tuamotu and Pitcairn Islands with Rapa, but should exclude Easter Island, Tonga and Samoa. ? Regionalization schemes should take island type into consideration. ? The Pitcairn Island group can serve as a useful model for Pacific biogeographical analyses.     

New insights into the systematics of the enigmatic Polynesian sandpipers Aechmorhynchus parvirostris and Prosobonia leucoptera

With only a single extant representative, endemic to the Tuamotu Archipelago, the Polynesian sandpipers (Aechmorhynchus and Prosobonia) may have had a larger distribution in Eastern Polynesia in the past, with four endemic taxa. Although these aberrant sandpipers' membership to the Scolocapidae has been well supported, finding their closest living taxa has proved difficult and the phylogenetic relationships of these taxa have remained unresolved. We present the first molecular analysis of the Polynesian sandpipers, including sampling of the only known specimen of the extinct Prosobonia leucoptera, collected in 1773. Based on mitochondrial and nuclear gene sequence data, the phylogenetic analyses demonstrate that the Polynesian sandpipers were sister taxa and belonged to the clade that included the other sandpipers (Calidris and allies) and turnstones (Arenaria), although without a close relative among extant genera. Divergence time estimates suggested that the lineage leading to Prosobonia diverged from the other extant sandpipers during the Oligocene and that either the Line Islands or the Tuamotu Archipelago were probably the first archipelagos colonized by the Prosobonia lineage. On the basis of these results, we suggest that Aechmorhynchus parvirostris and Prosobonia leucoptera be regarded as related species within the same genus, and thus that the senior name Prosobonia be used for both taxa.     

Territorial spacing and interspecific competition in three species of reed warblers

Summary Ecological and temporal aspects of the occurrence and dispersion patterns in three reed warbler species were examined. This provided insight into their intra-and interspecific territoriality. Seasonal changes in the occurrence of territorial males in a reed bed along a gradient from the shore landward were studied. The results demonstrate that intra- and interspecific spacing depends on interspecific dominance and/or ecological constraints. In this case it was shown that there is competition for common habitats between reed warblers and great reed warblers, whereas the moustached warbler is ecologically segregated. The reasons for these dispersal patterns are discussed. They do not appear to enhance the availability of food resources. It seems more likely that territorial spacing might function to reduce nest predation.     

Sex ratio variation among broods of great reed warblers Acrocephalus arundinaceus

The sex of 746 great reed warbler fledglings (from 175 broods) was determined by the use of single primer polymerase chain reaction. The reliability of the technique was confirmed as 104 of the fledglings were subsequently recorded as adults of known sex. The overall sex ratio did not differ from unity. Variation in sex ratios between broods was larger than expected from a binomial distribution. Female identity explained some of the variation of brood sex ratio indicating that certain females consistently produced sex ratios that departed from the average value in the population. The theory of sex allocation predicts that parents should adjust the sex ratio of their brood to the relative value of sons and daughters and this may vary in relation to the quality of the parents or to the time of breeding. In the great reed warbler, the proportion of sons was not related to time of breeding, or to any of five female variables. Of five male variables, males with early arrival date tended to produce more daughters. The sex ratio of fledglings that were a result of extra-pair fertilizations did not differ from that of legitimate fledglings. Hence, there is currently no evidence of that female great reed warblers invest in a higher proportion of sons when mated with attractive males.     

Uniform phenotype conceals double colonization by reed-warblers of a remote Pacific archipelago

Aim Remote oceanic islands often provide good illustrations of adaptive radiations, but phylogenetic studies have also demonstrated unexpected multiple colonization events for a given archipelago. In this study we investigate the relationships among endemic populations of the Marquesas reed‐warbler, Acrocephalus mendanae Tristram, 1883, which have colonized nearly all islands of this remote Polynesian archipelago, and which exhibit a very uniform plumage pattern. We study the phylogeny and morphology of all subspecies in the Marquesas, providing an examination of the position of the Marquesas lineages in relation to reed‐warblers distributed across multiple Polynesian archipelagos. Location This study focused on all the main islands of the Marquesas archipelago, along with samples from other Polynesian archipelagos (Society, Tuamotu, Austral, Cook, Kiribati) and Australia. Methods We used mitochondrial DNA markers (cytochrome b and ND2 genes) to develop a phylogeny of the main eastern Polynesian taxa. All subspecies for the Marquesas were investigated, including multiple individuals per island. Phylogenetic analyses using maximum‐likelihood and Bayesian approaches were employed to infer relationships among A. mendanae populations and between the main Polynesian archipelagos. Morphometric analyses based on 110 specimens from museum collections were performed on external characters to investigate the differences between islands, and these results were compared to the phylogeny. Results Our data indicate that the Marquesas reed‐warbler is in fact a polyphyletic taxon including two independent lineages: the northern Marquesas reed‐warbler, closely related to the Tuamotu reed‐warbler, and the southern Marquesas reed‐warbler, sister taxon to that endemic to the Kiribati. Analyses of morphological characters show that the size and shape features of the Marquesas reed‐warblers exhibit high plasticity linked to adaptation to ecological factors, particularly habitat richness (the diversity of vegetation structure that provides suitable resources and habitat for reed‐warblers, simplified here as the number of indigenous plant species). Main conclusions Our results suggest that reed‐warblers have successfully colonized the Marquesas archipelago, one of the most remote groups of islands in the Pacific Ocean, at least twice. Both events occurred more or less simultaneously at ca. 0.6 Ma, and are more recent than the islands' formation. We outline the taxonomic consequences of our phylogeny and discuss the supertramp strategy of reed‐warblers in the Pacific.     

Phylogeography of skinks (Chalcides) in the Canary Islands inferred from mitochondrial DNA sequences

Mitochondrial DNA (mtDNA) evolution was investigated in skinks of the genus Chalcides found in the Canary Islands ( Ch. sexlineatus, Ch. viridanus and Ch. simonyi ), together with some North African congenerics ( Ch. polylepis and Ch. mionecton ). Several sites were included within islands to cover areas of known within-island geographical variation in morphology. Skinks from the islands of El Hierro and La Gomera appear to be sister taxa. The relationships between this clade and the Tenerife and Gran Canarian skinks were not fully resolved, although the best working hypothesis indicated monophyly with the former, with the latter forming a closely related outgroup. Ch. simonyi from Fuerteventura was more distantly related to the Western Canary Island skinks and did not show close relationships with the North African species Ch. mionecton and Ch . polylepis . Possible colonization sequences for the four most Western Canary Islands were considered. El Hierro appears to have been colonized relatively recently from La Gomera, commensurate with the recent origin of this island, while dispersal between La Gomera and Tenerife and between Gran Canaria and Tenerife or La Gomera appears to have taken place considerably earlier. Substantial within-island haplotype divergence was found in Gran Canaria and Tenerife. This may be a result of recent periods of intense volcanic activity found within these two islands. Lower levels of within-island differentiation are found in La Gomera and El Hierro and may be explained by lower levels of volcanic activity during recent geological history and a more recent colonization, respectively.     

A spatiotemporal analysis of acoustic interactions between great reed warblers (Acrocephalus arundinaceus) using microphone arrays and robot audition software HARK

Acoustic interactions are important for understanding intra‐ and interspecific communication in songbird communities from the viewpoint of soundscape ecology. It has been suggested that birds may divide up sound space to increase communication efficiency in such a manner that they tend to avoid overlap with other birds when they sing. We are interested in clarifying the dynamics underlying the process as an example of complex systems based on short‐term behavioral plasticity. However, it is very problematic to manually collect spatiotemporal patterns of acoustic events in natural habitats using data derived from a standard single‐channel recording of several species singing simultaneously. Our purpose here was to investigate fine‐scale spatiotemporal acoustic interactions of the great reed warbler. We surveyed spatial and temporal patterns of several vocalizing color‐banded great reed warblers (Acrocephalus arundinaceus) using an open‐source software for robot audition HARK (Honda Research Institute Japan Audition for Robots with Kyoto University) and three new 16‐channel, stand‐alone, and water‐resistant microphone arrays, named DACHO spread out in the bird's habitat. We first show that our system estimated the location of two color‐banded individuals’ song posts with mean error distance of 5.5 ± 4.5 m from the location of observed song posts. We then evaluated the temporal localization accuracy of the songs by comparing the duration of localized songs around the song posts with those annotated by human observers, with an accuracy score of average 0.89 for one bird that stayed at one song post. We further found significant temporal overlap avoidance and an asymmetric relationship between songs of the two singing individuals, using transfer entropy. We believe that our system and analytical approach contribute to a better understanding of fine‐scale acoustic interactions in time and space in bird communities.     

Unexpected evolutionary diversity in a recently extinct Caribbean mammal radiation

Identifying general patterns of colonization and radiation in island faunas is often hindered by past human-caused extinctions. The insular Caribbean is one of the only complex oceanic-type island systems colonized by land mammals, but has witnessed the globally highest level of mammalian extinction during the Holocene. Using ancient DNA analysis, we reconstruct the evolutionary history of one of the Caribbean''s now-extinct major mammal groups, the insular radiation of oryzomyine rice rats. Despite the significant problems of recovering DNA from prehistoric tropical archaeological material, it was possible to identify two discrete Late Miocene colonizations of the main Lesser Antillean island chain from mainland South America by oryzomyine lineages that were only distantly related. A high level of phylogenetic diversification was observed within oryzomyines across the Lesser Antilles, even between allopatric populations on the same island bank. The timing of oryzomyine colonization is closely similar to the age of several other Caribbean vertebrate taxa, suggesting that geomorphological conditions during the Late Miocene facilitated broadly simultaneous overwater waif dispersal of many South American lineages to the Lesser Antilles. These data provide an important baseline by which to further develop the Caribbean as a unique workshop for studying island evolution.     

The flora of the Pitcairn Islands: a review

vascular plant flora of the Pitcairn Islands, south-central Pacific Ocean, is described based on extensive new collections made in 1991 and previously published records. Two vascular plants occur on Ducie Atoll; one (Pemphis acidula) is a new record. Sixty-three native vascular plants occur on Henderson, of which nine are endemic; Canavalia rosea, Operculina lurpethum, Psilotum nudum and Solanum americanum are new records for the island. Oeno Atoll has 16 native vascular plants; the single endemic (Bidens hendersonensis var. oenoensis) was not found in 1991 despite careful searches. Triumfetta procumbens was new for Oeno. Sixty-six native vascular plants have now been recorded from Pitcairn Island, there are two endemic ferns and seven endemic angiosperms in this number. A number of non-native taxa were new to Pitcairn. Some of the previously described taxa could not be found on Pitcairn, probably because they are very rare and only a small amount of time was spent collecting on Pitcairn. Many of the Pitcairn taxa are threatened by the spread of introduced species, especially Syzygium jambos.     

Evolution and biogeography of the genus Tarentola (Sauria: Gekkonidae) in the Canary Islands,inferred from mitochondrial DNA sequences

Sequences from fragments of the 12S ribosomal RNA and cytochrome b mitochondrial genes were used to analyze phylogenetic relationships among geckos of genus Tarentola from the Canary Islands. A surprisingly high level of within island differentiation was found in T. delalandii in Tenerife and T. boettgeri in Gran Canaria. Molecular differentiation between populations of T. angustimentalis on Lanzarote and Fuerteventura, and between Moroccan and Iberian Peninsula T. mauritanica, also indicate that at least two subspecies should be recognized within each of them. Phylogenetic relationships among these species reveals a higher level of differentiation and a more complex colonization pattern than those found for the endemic genus Gallotia. Lack of evidence for the presence of T. boettgeri bischoffi on the island of Madeira does not seem to support the origin of T. delalandii, T. gomerensis and the canarian subspecies of T. boettgeri from this island, whereas molecular data confirms that T. angustimentalis is a sister species of the continental T. mauritanica. Several independent colonization events from the continent and the extinction of some species are probably responsible for the current distribution of Tarentola in the Canary Islands.     

Parallel evolution in molar outline of murine rodents: the case of the extinct Malpaisomys insularis (Eastern Canary Islands)

The lava mouse Malpaisomys was part of Pleistocene and Holocene faunas of the eastern Canary Islands; it became extinct during historical times. In order to evaluate the evolutionary processes of this endemic species, we set out to identify its relationships with possible mainland relatives. Its dental morphology was compared to a set of fossil and modern murine rodents from various phylogenetic groups, characterized by different diets and dental patterns, using a quantitative method based on a Fourier analysis of the outline of the first upper and lower molars. This morphometric analysis identified different evolutionary grades that are independent of the phylogenetic group. The first cluster is associated with primitive, asymmetric dental outlines, the second with intermediate forms linked to a more herbivorous diet. Highly differentiated forms diverge not only from the second cluster but also from each other. Our investigations reveal Malpaisomys to have had an intermediate dental pattern, a result which confirms previous palaeoecological interpretations of this taxon. However, conclusions about its closest mainland relative remain tentative. Based on a comparison of dental size and shape, as well as geographical considerations, a possible mainland ancestor could be either Paraethomys (North-African Pliocene) or Occitanomys (South-western European Pliocene). Such results support the hypothesis of a Pliocene colonization event, in which case Malpaisomys would display a normal evolutionary rate in dental size and shape, in spite of the insular context. In contrast, a late colonization by a modern representative of the African fauna would imply exceptionally high evolutionary rates compared to the background morphological evolution and a decrease in size that is unlikely under insular conditions.  © 2004 The Linnean Society of London, Zoological Journal of the Linnean Society , 2004, 142 , 555–572.     

The first precinctive Carabidae from Moorea,Society Islands: new Mecyclothorax spp. (Coleoptera) from the summit of Mont Tohiea

Seven species of Mecyclothorax Sharp from Moorea, Society Islands are newly described; Mecyclothorax perraulti sp. n., Mecyclothorax pahere sp. n., Mecyclothorax menemene sp. n., Mecyclothorax mahatahi sp. n., Mecyclothorax popotioaoa sp. n., Mecyclothorax mapo sp. n., and Mecyclothorax fatata sp. n. These constitute the first Mecyclothorax species described from Moorea, and the first carabid beetle species shown to be geographically restricted to that island. Each of the newly described species is most similar to a different species on the island of Tahiti, suggesting that none of the seven Moorean taxa are evolutionary end-products of autochthonous speciation within Moorea. The occurrence of precinctive Mecyclothorax species on both Moorea and Tahiti demonstrates that radiation of Mecyclothorax in the Society Islands has been facilitated by speciation events implicating both islands. Whether this speciation has been preceded by vicariance or dispersal is discussed, with the generality of a dispersal hypothesis tested using information from Society Island Nabidae (Hemiptera). Salient morphological characters for taxa in the Society and Hawaiian Islands are compared to those representing a broad survey of southwest Pacific Mecyclothorax spp. This comparison supports the independent founding of each radiation in the Societies and Hawaii from an Australian ancestral propagule, likely drawn from the ecologically general, geographically widespread Mecyclothorax punctipennis (Macleay).     

Molecular phylogeny and the historical biogeography of the warblers of the genus Sylvia (Aves)

A molecular phylogeny based on DNA/DNA hybridization revealed that the Sylvia-Parisoma complex is monophyletic and includes three main groups of species, the “mid-European” warblers, the genus Parisoma, and the “eu-Mediterranean” Sylvia species sensu stricto. The latter can be assigned to three main clusters, a “West-Mediterranean” group, a “Central-Mediterranean group”, and an “East-Mediterranean” group. The radiation of the whole complex is much more ancient than formerly believed. It started ca 12–13 Ma ago and the ancestors of the main extant groups differentiated during the Pliocene. Only speciation events within the “eu-Mediterranean” lineages occurred during the Pleistocene. The paleoclimatical and paleoecological history of the Mediterranean region is too complicated to provide any evidence for direct relationships between past events and evolutionary steps of these taxa which did not leave any reliable fossil record. However, some major speciation events may be related to well documented climatical crises as well as paleobotanical data. The largely man-induced extension of matorrals over several millenia presumably extended the range of several species that were formerly much more restricted, which complicates reconstruction of the spatio-temporal course of speciation.     

Taxonomic status of the extinct Canary Islands Oystercatcher Haematopus meadewaldoi

Mitochondrial genes were sequenced from four specimens of the extinct Canary Islands Oystercatcher Haematopus meadewaldoi and compared with African Oystercatcher Haematopus moquini, Eurasian Oystercatcher Haematopus ostralegus and an old unidentified extralimital ‘black’ oystercatcher specimen from The Gambia. At these loci, H. meadewaldoi was approximately 99.65% identical to multiple Eurasian Oystercatcher samples and in phylogenetic trees fell within the range of genetic variation observed in that species. The mystery Gambian bird was resolved as an extralimital H. moquini. We conclude that H. meadewaldoi was most likely a recently diverged melanistic morph or subspecies of H. ostralegus, although further genomic studies will be required to determine whether there has been a period of isolation followed by introgression.     

Patterns of colonization, evolution and gene flow in species of the genus Patella in the Macaronesian Islands

The study of phylogeographical patterns may contribute to a better understanding of factors affecting the dispersal of organisms in ecological and historical times. For intertidal organisms, islands are particularly suitable models allowing the test of predictions related to the efficacy of pelagic larvae dispersal. Here, we study the phylogeographical patterns and gene flow within three groups of species of the genus Patella present in the Macaronesian Islands that have been previously shown to be monophyletic. The genetic variability of around 600 bp of the mitochondrial gene cytochrome c oxidase subunit I was studied by single strand conformation polymorphism and/or sequencing for seven species of limpets. A total of 420 samples were analysed from the Macaronesian archipelagos, North Africa, and Atlantic and Mediterranean shores of the Iberian Peninsula. No clear geographical pattern or temporal congruence was found between the three groups of species, pointing to independent histories and colonization events. However, for the three groups, the split between the Macaronesian and the mainland forms most probably occurred before 3.9 million years ago, predating the establishment of the current circulation patterns. The presence of pelagic larvae in these species is shown to be insufficient to ensure gene flow between continental and Macaronesian populations and between the Macaronesian archipelagos. In the endangered Azorean populations of Patella candei, there is restricted gene flow to Flores and Graciosa.     

Systematics and evolution of the libyan jird based on molecular and morphometric data

The libyan jird is one of the most widely distributed species among wild rodents, with its range extending from Morocco to China. Fifteen subspecies were described but their validity and the phylogenetic relationships among them are uncertain. Based on a comprehensive sampling, this study aims to define subspecies limits within Meriones libycus and to discuss the factors driving subspecific diversification. We used an integrative approach combining molecular (Cytochrome b and Cytochrome Oxidase Subunit 1 genes) and geometric morphometric data. Genetic data allowed us to identify three allopatric lineages within M. libycus: Western lineage in North Africa, Central lineage in Saudi Arabia, Jordan, and Syria, and Eastern lineage in Iran, Afghanistan, and China. These three lineages can also be differentiated based on skull morphology. Our results support the existence of at least three subspecies within the libyan jird: Meriones libycus libycus, M. l. syrius, and Meriones libycus erythrourus. Based on our divergence time estimates, all divergence events within M. libycus probably occurred during the Pleistocene, after 1.597 Ma. Quaternary climate fluctuations in the Sinai Peninsula explain the differentiation between the African M. l. libycus and the Levant-Arabian M. l. syrius. The differentiation of M. l. syrius with respect to the eastern M. l. erythrourus is putatively linked to the climatic fluctuations and tectonic activity of the Zagros Mountains and/or the Mesopotamia Plain of Iraq during the Pleistocene.