A survey for the Critically Endangered Liben Lark Heteromirafra archeri in Somaliland,north-western Somalia

The Critically Endangered Archer's Lark (now Liben Lark) Heteromirafra archeri was formerly considered to be endemic to north-western Somalia and known only from the Tog Wajaale Plain, where 18 specimens were collected between 1918 and 1922. Fifteen visits between 1970 and 2008 failed to relocate the species there, although popula- tions are now known from adjacent Ethiopia. We conducted three days of intensive surveys on the Tog Wajaale Plain in May 2010. Despite the three other lark species present being in full display, and H. archeri being recorded to have bred in early June, no Liben Larks were found. Vegetation structure surveys indicated that the plain has a taller and denser growth of grass than either of the other known localities for Liben Lark (the Liben and Jijiga Plains) making Tog Wajaale Plain seem superficially more suitable for the species, which prefers areas of taller grass elsewhere. However, previous large-scale agricultural activities may have altered the composition of grass species and precipitated the observed invasion of exotic weeds, notably Parthenium hysterophorus. Importantly, the Tog Wajaale Plain has a greater density of bushes than either the Liben or Jijiga Plains, possibly making ground-nesting birds more susceptible to predation by perch hunters.     

Densely sampled phylogenetic analyses of the Lesser Short-toed Lark (Alaudala rufescens) — Sand Lark (A. raytal) species complex (Aves,Passeriformes) reveal cryptic diversity

The taxonomy of the Lesser/Asian Short-toed Lark Alaudala rufescens–cheleensis complex has been debated for decades, mainly because of minor morphological differentiation among the taxa within the complex, and different interpretations of the geographical pattern of morphological characters among different authors. In addition, there have been few studies based on non-morphological traits. It has recently been suggested based on a molecular study of the lark family Alaudidae that the Sand Lark A. raytal is nested within this complex. We here analysed mitochondrial cytochrome b (cyt b) from 130 individuals across the range of this complex (hereafter called Alaudala rufescens–raytal complex), representing all except two of the 18 currently recognized subspecies. We also analysed 11 nuclear markers from a subsample of these individuals, representing all of the clades found in the cyt b tree. Five primary clades were recovered, which confirmed that A. raytal is nested within this complex. Divergence time estimates among these five clades ranged from 1.76 to 3.16 million years (my; 95% highest posterior density [HPD] 1.0–4.51 my) or 1.99–2.53 my (95% HPD 0.96–4.3 my) in different analyses. Only four of the currently recognized subspecies were recovered as monophyletic in the cyt b tree. Our results call for a taxonomic revision, and we tentatively suggest that at least four species should be recognized, although we stress the need for an approach integrating molecular, morphological and other data that are not yet available.     

The Orange River avifauna: abundance,richness and comparisons

Like many other southern African rivers, the avifaunal richness and abundance of the Orange River, southern Africa's longest, remains virtually unknown. Given that the Orange River runs through hyper-arid regions in its lower reaches, its linear oasis effect and its role as a reservoir for Red Data bird species is likely but has not been quantified. Two surveys either side of the border town of Noordoewer, in the rainy and dry seasons of 1996 and 1997 deter-mined that species richness was higher in riverine habitat (71 species) than in the surrounding Nama Karoo (46 species) confirming the linear oasis effect. Surveys revealed five Namibian Red Data species. Peregrine Falcons, Falco peregrinus, and African Fish Eagles, Haliaeetus vocifer, were relatively common while the endangered Cape Eagle Owl, Bubo capensis, occurred sporadically. Among wetland Red Data bird species, Black Storks, Ciconia nigra, occurred in less disturbed areas, and White-backed Night Herons, Gorsachius leuconotus, were an unexpected species at this locality. The overall species richness of 103 birds and a density of riparian birds of 31 -3 4 birds/10 km, makes the lower Orange River avifauna similar to the Cunene River but tenfold less abundant than Namibia's other perennial rivers. Despite the low diversity, the arid habitats of the lower Orange River support greater diversity than more easterly regions when all current surveys of richness and abundance from Lesotho to the mouth on the Namib coast are combined. This east-west trend may be explained by the slower flow and warmer water of the lower Orange River but natural turbidity and regulation may explain the generally low diversity and abundance indices. Major rivers in southern Africa appear to have been over-looked as wetland habitats, and further avifaunal surveys of all of them are recommended.     

No evidence for observer effects on Lark Sparrow nest survival

ABSTRACT Methods for monitoring bird nests might influence rates of nest predation, but the effects of various methods (e.g., visual markers and observer visitation rates) are often separately investigated among disparate avian taxa and geographic regions. Few investigators have explored the potential effects observers might have on nest success of grassland birds, despite concerns regarding population declines of these species in North America. We examined the possible effects of three monitoring techniques on daily nest survival of Lark Sparrows (Chondestes grammacus): (1) presence or absence of visible markers near nests, (2) observer visitation frequency, and (3) presence or absence of data loggers in nests. We monitored 113 Lark Sparrow nests during the 2009 breeding season. Of these nests, 88.5% failed due to predation, abandonment, weather, or unknown causes, yielding an overall nest success estimate of 9.8% based on daily survival estimation. Main effects of each monitoring technique appeared in top (ΔAICc <2) logistic exposure models. However, 95% confidence intervals around parameter estimates for each technique included zero, indicating no significant effects on daily nest survival. Our results suggest that the nest‐monitoring techniques we used had no effect on Lark Sparrow nest success and, if true, nest survival of other songbirds in arid grasslands of the Great Plains may also be unaffected by cautious nest monitoring. However, we cannot rule out the possibility that any effects of the various techniques in our study were masked by locally intense nest predation. Therefore, additional study is needed to determine if there may be observable variation in nest survival among various nest‐monitoring treatments in other areas of the southern Great Plains where nest predation is less frequent.     

Effect of Sky Lark plots and additional tramlines on territory densities of the Sky Lark Alauda arvensis in an intensively managed agricultural landscape

Capsule: Sky Lark plots and additional tramlines increase Sky Lark Alauda arvensis territory densities in winter crops.

Aims: To analyse the effects on Sky Lark territory density of Sky Lark plots and additional tramlines in winter cereals and oilseed rape.

Methods: We mapped Sky Lark territories on fields with Sky Lark plots or additional tramlines as well as on adjacent control sites, from 2010 to 2013 in Saxony, Germany, where agricultural land use is intensive, dominated by winter-sown crops and takes place in large fields.

Results: In test sites with Sky Lark plots, 5.6 and 3.1 territories per 10?ha were found in the early (April/May) and late periods (June/July) respectively, compared to 3.3 and 1.4 territories per 10?ha in control sites. Sky Lark territory densities in fields with additional tramlines were 1.6 times higher in the early period (4.2 versus 2.6 territories per 10?ha) and 2.2 times higher in the late period (3.6 versus 1.6 territories per 10?ha).

Conclusion: Sky Lark plots and additional tramlines improve large fields for the Sky Lark and have the potential to increase the Sky Lark population.     

THE WATTLED CRANE BUGERANUS CARUNCULATUS (GMELIN)

The southern African endemic Short-clawed Lark Certhilauda chuana has two geographically isolated populations. Little is known about the smaller eastern population, which is restricted to the Polokwane Plateau, South Africa. To provide input for a conservation strategy for the eastern population, this study evaluated its habitat preference. Territories previously supporting Short-clawed Lark (1995–2005) were visited in the 2008/09 breeding season. Biophysical habitat parameters were measured at each site to determine the differences between active territories and those no longer active. Active territories had significantly less grass cover, more bare ground, shorter grass, smaller trees and had burned more recently. Bush encroachment and a reduction in the fire frequency were identified as serious threats to the Short-clawed Lark. However, overgrazing and harvesting of wood in rural areas may benefit the species.     

Molecular divergence in the eastern Asia– eastern North America disjunct section Rytidospermum of Magnolia (Magnoliaceae)

Molecular divergence in the eastern Asia—eastern North American disjunct section Rytidospermum of Magnolia was investigated by allozyme electrophoresis, chloroplast DNA (cpDNA) restriction site analysis, and gene sequencing. We calculated Nei's genetic identities between two Asian species, M. officinalis var. biloba and M. hypoleuca, and three American species, M. tripetala, M. fraseri var. fraseri, and M. macrophylla var. macrophylla, by using gene frequency data from 17 nuclear-encoded allozyme loci in 67 populations. We then estimated cpDNA sequence divergence between the five species by examining restriction site variation for ten endonucleases over the entire genome. Finally, nucleotide sequences of the chloroplast gene rbcL were compared between M. hypoleuca, M. tripetala, and M. macrophylla var. macrophylla. All three methods consistently yielded low divergence values between the American species M. tripetala and its Asian sister taxa, M. officinalis var. biloba and M. hypoleuca (Nei's I = 0.712 and 0.809, respectively; D-cpDNA = 0.083% for both pairs; D-rbcL = 0.000% between M. tripetala and M. hypoleuca). The other two American species, M. fraseri var. fraseri and M. macrophylla var. macrophylla, neither of which is sister to the Asian taxa, exhibited much higher divergence from the Asian taxa. We interpreted the low divergence between M. tripetala and its Asian sister taxa as a result of recent separation (the late Miocene to early Pliocene), based on time estimates from molecular data as well as geological and paleoclimatic evidence. A comparison of our results with those of the earlier studies revealed a diverse array of levels of divergence between several eastern Asian and eastern North American species pairs. Though different extinction patterns and variation in molecular evolutionary rates may be partly responsible, this heterogeneous pattern of divergence is best explained by different times of disjunction in different taxa, which in turn suggests that the floristic similarity between the two continents was most likely attained by multiple migrations via both Bering and North Atlantic land bridges, or possibly even with involvement of dispersal.     

Nest exclosures do not improve Streaked Horned Lark nest success

ABSTRACT Improving the conservation status of rare and declining species often requires multiple strategies targeted at several vital rates. We report on one of several ongoing management actions intended to benefit the declining population of Streaked Horned Larks (Eremophila alpestris strigata). To improve Streaked Horned Lark fecundity, we employed predator exclosures (wire cages) around nests (N= 33 exclosed and 32 not exclosed) in 2009 and 2010 at two sites in Oregon and two in Washington with the goal of excluding larger birds, the primary lark nest predators. We found no statistically significant effect of exclosures on nest success. For exclosed nests, lower rates of nest predation (exclosed = 12%, unexclosed = 48%) were offset by higher rates of nest abandonment (exclosed = 27%, unexclosed = 0%). Nest abandonment was likely caused by a variety of factors including American Kestrels (Falco sparverius) perching on exclosures, and predation of adults associated with exclosed nests. Our results suggest that the current exclosure design does not improve Streaked Horned Lark fecundity and may negatively affect adult survival. To improve exclosure effectiveness, we recommend modifications that prevent kestrels from perching on exclosures and deny their access to the nest. We also recommend that modifications be applied in an adaptive management framework that includes close monitoring to assess their effectiveness, and subsequent adaptation that might include continued structural modification of exclosures or discontinued use on some or all sites.     

Breeding biology of the endangered Dupont’s Lark Chersophilus duponti in two separate Spanish shrub-steppes

Capsule: The Dupont’s Lark Chersophilus duponti in Iberia has relatively high breeding success in both core and fragmented habitats, so population declines are more likely to be the result of low juvenile or adult survival.

Aims: To measure important aspects of the reproductive biology of one of the most endangered and least known larks: the Dupont’s Lark C. duponti.

Methods: We monitored 36 nests in 2 Spanish shrub-steppes, one holding one of the largest European populations (250 pairs) and one composed by fragmented habitat patches holding a smaller population (50 pairs).

Results: The breeding season went from late-March to early July. Overall mean (±sd) clutch size was 3.47?±?0.56, and the number of fledglings per successful nest was 3.0?±?1.15. Mean nestling period was short (8.2 days). Nests showed similar daily survival rate during the incubation period (0.9750?±?0.0110) as during the nestling period (0.9545?±?0.0168), with a mean breeding success of 50%. Predation was the main cause of complete nest failure (83.3% of failed nests in both localities).

Conclusion: Breeding parameters did show no significant variation between populations. Breeding success in both sites was generally higher than recorded in previous studies of this and most other lark species, which suggests that breeding success does not compromise long-term viability of these populations. The decline of the studied populations should be explained by other causes, such as a general decrease in habitat quality, habitat loss or habitat fragmentation.     

Genetic structure among Charadrius plovers on the African mainland and islands of Madagascar and St Helena

Colonization of islands by long-distance dispersers has great impact on genetic diversification among populations and may spearhead speciation events. We investigated intra- and interspecific divergence in Charadrius plovers with populations on mainland Africa, Madagascar and St Helena. We analysed microsatellite loci and sequence data from four nuclear and two mitochondrial gene regions. Charadrius plovers are shorebirds with high dispersal and mobility. Our results confirmed genetic differentiation between Madagascar and mainland populations of three plover species (White-fronted Plover Charadrius marginatus, Kittlitz's Plover Charadrius pecuarius and, based on sequence data only, Three-banded Plover Charadrius tricollaris) but highlight substantial variation in levels of intraspecific divergence among the three species. Namely, the Kittlitz's Plover, a dispersive habitat generalist with a polygamous mating system, exhibited lower island–mainland differentiation (0.05% COI sequence divergence) compared with the two monogamous species, the White-fronted Plover (0.6% COI divergence) and Three-banded Plover (1.6% COI divergence). In addition, past colonization of the islands of St Helena and Madagascar by ancestors of today's Kittlitz's Plover has led to the evolution of two endemic island species, the Madagascar Plover Charadrius thoracicus and the more closely-related St Helena Plover Charadrius sanctaehelenae. We discuss the factors driving species differences in island–mainland divergence and highlight the importance of conserving genetically unique island populations and island habitats to safeguard future evolutionary potential.     

Assessment of species limits in African ‘brown buntings’ (Emberiza,Passeriformes) based on mitochondrial and nuclear sequence data

We estimated a phylogeny for 10 taxa currently placed in four polytypic species that collectively encompass the African ‘brown buntings’: Cape Bunting Emberiza capensis, Cinnamon‐breasted Bunting Emberiza tahapisi, Lark‐like Bunting Emberiza impetuani and House Bunting Emberiza striolata. We made use of the mitochondrial cytochrome b gene and the nuclear introns 6–7 of ornithine decarboxylase (ODC), and intron 2 of myoglobin. There was substantial cytochrome b sequence divergence between taxa currently treated as conspecific: sahari vs. striolata (2.6–3.1% (uncorrected‐p); 3.0–3.6% (HKY + I)), and goslingi vs. tahapisi (4.4–4.7% (uncorrected‐p); 5.4–5.9% (HKY + I)). The degree of divergence of the nuclear loci among taxa was limited, and these loci lacked reciprocal monophyly, most likely as a consequence of incomplete lineage sorting. A single representative of the taxon septemstriata, generally treated as a member of the dark‐throated tahapisi group, here appears to be genetically consistent with the grey‐throated goslingi, and may be of hybrid origin. All other taxa allocated to E. striolata and E. tahapisi make up four reciprocally monophyletic groups consistent with sahari, striolata, tahapisi and goslingi, respectively. The extent of genetic evidence suggests that these taxa have been evolving as separate evolutionary lineages for a long time. This is further manifested in several morphological and vocal characteristics described previously, and we propose that these divergent taxa be treated as separate species: Cinnamon‐breasted Bunting Emberiza tahapisi, Gosling's Bunting Emberiza goslingi, Striolated Bunting Emberiza striolata and House Bunting Emberiza sahari. We do not propose any taxonomic changes regarding Emberiza impetuani or Emberiza capensis.     

NOTE ON THE OCCURRENCE OF A VENTRAL SUCKER-LIKE MOUTH FORM IN LABEO UMBRATUS (PISCES,CYPRINIDAE)

SUMMARY

The occurrence of a ventral sucker-like mouth form in certain populations of Labeo umbratus, which are free from competition by L. aapensis, is reported. It is proposed that this might be an adaptation to feeding off stones which enhances the fish's changes of survival. If this is so, the absense of mouth form variation in Orange River populations illustrates the force of interspecific competition with L. capensis.     

GENETIC RELATIONSHIPS AND PATTERNS OF ALLOZYMIC DIVERGENCE IN THE IPOMOPSIS AGGREGATA COMPLEX AND RELATED SPECIES (POLEMONIACEAE)

The Ipomopsis aggregata complex consists of diploid, outcrossing, perennial herbs. The group is highly variable morphologically and is treated as three species: I. aggregata, I. tenuituba, and I. arizonica. Geographic races of I. aggregata and I. tenuituba are recognized as subspecies. Enzyme electrophoresis was used to examine genetic relationships among populations and taxa in the Ipomopsis aggregata complex and some related species. Genetic data for 23 allozyme loci from 60 populations were also used to determine how genetic variation is distributed geographically. Populations in the southwestern United States were more variable than those in the northwest: the center of genetic diversity corresponded to the center of species diversity. Allozymic data provided no evidence of loss of genetic variability associated with recent and rapid divergence. Genetic relationships based on Nei's genetic identity did not correspond to taxonomic relationships. For example, populations of both I. arizonica and I. tenuituba clustered within I. aggregata. Despite relatively high levels of genetic diversity among populations, diversity among taxa was low. Results indicated that floral divergence and concomitant speciation have occurred recently in the Ipomopsis aggregata complex. Allozymic patterns also reflected convergent evolution for floral morphology and possible introgression. Despite morphological differences among species, insufficient evolutionary time has elapsed for allelic fixation at neutral or near-neutral allozyme loci.     

4. A FIELD NATURALIST'S PEREGRINATIONS. BIRD NOTES FROM A NORTHERN SECTOR OF SOUTHERN RHODESIA—Part III

Boyer, H. J. 1988. Breeding biology of the Dune Lark. Ostrich 59:30-37.

The peak of the breeding season of the Dune Lark Mirafra erythrochlamys occurred in January and February and was not dependent on rainfall. Most nests were domed, although one undomed nest was recorded. Ninety-one percent of clutches were of two eggs (mean = 1,9; range 1–2; n = 11). The eggs are described and measurements given. Incubation, by the female only, began with the laying of the second egg, and hatching occurred after 13–14 days. Growth and development of nestlings are described. The young left the nest after 12–14 days, and post-nestling parental care lasted for approximately one month. Sixty-one percent of eggs hatched. and 28% produced young which successfully left the nest. Most losses of eggs and young were the result of predation.     

LIFE-HISTORY VARIATION AND COMPARATIVE PHYLOGEOGRAPHY OF DARTERS (PISCES: PERCIDAE) FROM THE NORTH AMERICAN CENTRAL HIGHLANDS

Phylogeography seeks to evaluate the relationship between genetic variation and geographic distribution of a species to examine the influence of historical events on divergence. Congruent phylogeographic patterns in codistributed species indicate historical association of the taxa being compared, and the uniform action of biogeographic events in shaping genetic variation. We sought to evaluate the congruence of patterns of genetic variation of five closely related fish species across a well-defined biogeographic boundary. We gathered allozymic and mitochondrial DNA (mtDNA) sequence data for five species of darters (Percidae: Etheostomatini) from populations distributed among biogeographic regions in the Ozark and Ouachita highlands of the south-central United States. Comparisons among species revealed noncongruence in the magnitude of genetic divergence in both allozymes and mtDNA sequences. We hypothesized that noncongruence resulted, in part, from differences in life histories of the species in our comparison. To address this hypothesis, we evaluated the association between gene flow (measured by N_em) and variation in body size and fecundity variables because they have been shown to influence gene flow in fishes. Correlation analysis revealed an association between gene flow and fecundity (r = 0.88), but not with body size (r = 0.36) or reproductive investment per individual (r = –0.23). The result was similar when independent contrasts of the original variables were used in correlation analyses. Phylogeographic analysis of mtDNA sequence data indicated the importance of history, evident in gene trees of Percina nasuta and Percina phoxocephala. Divergence rates between these two taxa may differ because of historically persistent differences in population sizes, reflected in present-day abundance and fecundity differences. Conversely, Percina caprodes showed little evidence of divergence in mtDNA sequences and yielded the highest mean N_em values from allozyme data. Comparisons among closely related, codistributed taxa may help discriminate among the possibilities for noncongruence in biogeographic studies by allowing evaluation of the influence of life history on patterns of gene flow.     

Contrasting intra versus interspecies DNA sequence variation for representatives of the Batrachospermales (Rhodophyta): Insights from a DNA barcoding approach

Sixty‐five accessions of the species‐rich freshwater red algal order Batrachospermales were characterized through DNA sequencing of two regions: the mitochondrial cox1 gene (664 bp), which is proposed as the DNA barcode for red algae, and the UPA (universal plastid amplicon) marker (370 bp), which has been recently identified as a universally amplifying region of the plastid genome. upgma phenograms of both markers were consistent in their species‐level relationships, although levels of sequence divergence were very different. Intraspecific variation of morphologically identified accessions for the cox1 gene ranged from 0 to 67 bp (divergences were highest for the two taxa with the greatest number of accessions; Batrachospermum helminthosum and Batrachospermum macrosporum); while in contrast, the more conserved universal plastid amplicon exhibited much lower intraspecific variation (generally 0–3 bp). Comparisons to previously published mitochondrial cox2–3 spacer sequences for B. helminthosum indicated that the cox1 gene and cox2–3 spacer were characterized by similar levels of sequence divergence, and phylogeographic patterns based on these two markers were consistent. The two taxa represented by the largest numbers of specimens (B. helminthosum and B. macrosporum) have cox1 intraspecific divergence values that are substantially higher than previously reported, but no morphological differences can be discerned at this time among the intraspecific groups revealed in the analyses. DNA barcode data, which are based on a short fragment of an organellar genome, need to be interpreted in conjunction with other taxonomic characters, and additional batrachospermalean taxa need to be analyzed in detail to be able to draw generalities regarding intraspecific variation in this order. Nevertheless, these analyses reveal a number of batrachospermalean taxa worthy of more detailed DNA barcode study, and it is predicted that such research will have a substantial effect on the taxonomy of species within the Batrachospermales in the future.     

Behaviour of Dinosaurian Track-Makers in the Winton Formation (Cretaceous,Albian–Cenomanian) at Lark Quarry,Western Queensland,Australia: Running or Swimming?

A remarkable assemblage of dinosaur trackways in the Winton Formation (Albian–Cenomanian) at Lark Quarry, in western Queensland, Australia, has long been regarded as evidence of a stampede involving small theropods, whose tracks were classified in the ichnogenus Skartopus, and small ornithopods, whose tracks represented a second ichnogenus, Wintonopus. However, one recently-published study has claimed that existing interpretation of Lark Quarry is incorrect: it maintains that all the track-makers were ornithopods, that the ichnogenus Skartopus is a variant form of Wintonopus, and that most of the Lark Quarry track-makers were not running but more probably swimming downstream in a current of water. Those iconoclastic claims are investigated here and shown to be untenable. They derive from overgeneralized interpretation of the Lark Quarry track assemblage and invoke ad hoc auxiliary hypotheses which are either untestable or demonstrably incorrect. Closer inspection of the evidence underpinning those claims corroborates the existing interpretation of Lark Quarry as the site of a dinosaurian stampede and confirms the validity of the original distinction between theropod tracks (Skartopus) and ornithopod tracks (Wintonopus).     

The <Emphasis Type="Italic">Drosophila</Emphasis> RNA-binding protein Lark is required for localization of Dmoesin to the oocyte cortex during oogenesis

The RNA-binding protein Lark has an essential maternal role during Drosophila oogenesis. Elimination of maternal expression results in defects in cytoplasmic dumping and actin cytoskeletal organization in nurse cells. The function of this protein is dependent on the activity of one or more N-terminal RNA-binding domains. Here, we report the identification of Dmoesin (Dmoe) as a candidate RNA target of Lark during oogenesis. In addition to actin defects in the nurse cells of lark mutant ovaries, we observed mislocalization of posteriorly localized mRNAs including oskar and germ cell less in the developing oocyte. Anteriorly and dorsally localized mRNAs were not affected. In addition, we observed displacement of the actin cytoskeleton from the oocyte plasma membrane. These phenotypes are reminiscent of mutations in Dmoe and suggested that this RNA maybe a potential target of Lark. We observed a significant decrease in Dmoe protein associated with the membrane of the developing oocyte with no changes in expression or localization within the nurse cells. Evidence for an association between Lark protein and moe RNA during oogenesis comes from results of a microarray-based Ribonomics approach to identify Lark RNA targets. Thus, our results provide evidence that Dmoe RNA is a target of Lark during oogenesis and that it likely regulates either the splicing or translation of this RNA. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Ring species as demonstrations of the continuum of species formation

In the mid‐20th century, Ernst Mayr (1942) and Theodosius Dobzhansky (1958) championed the significance of ‘circular overlaps’ or ‘ring species’ as the perfect demonstration of the gradual nature of species formation. As an ancestral species expands its range, wrapping around a geographic barrier, derived taxa within the ring display interactions typical of populations, such as genetic and morphological intergradation, while overlapping taxa at the terminus of the ring behave largely as sympatric, reproductively isolated species. Are ring species extremely rare or are they just difficult to detect? What conditions favour their formation? Modelling studies have attempted to address these knowledge gaps by estimating the biological parameters that result in stable ring species (Martins et al. 2013), and determining the necessary topographic parameters of the barriers encircled (Monahan et al. 2012). However, any generalization is undermined by a major limitation: only a handful of ring species are known to exist in nature. In addition, many of them have been broken into multiple species presumed to be evolving independently, usually obscuring the evolutionary dynamics that generate diversity. A paper in this issue of Molecular Ecology by Fuchs et al. (2015), focused on the entire genealogy of a bulbul (Alophoixus) species complex, offers key insights into the evolutionary processes underlying diversification of this Indo‐Malayan bird. Their findings fulfil most of the criteria that can be expected for ring species (Fig.  1 ): an ancestor has colonized the mainland from Sundaland, expanded along the forested habitat wrapping around Thailand's lowlands, adjacent taxa intergrade around the ring distribution, and terminal taxa overlap at the ring closure. Although it remains unclear whether ring divergence has resulted in restrictive gene flow relative to that observed around the ring, their results suggest that circular overlaps might be more common in nature than currently recognized in the literature. Most importantly, this work shows that the continuum of species formation that Mayr and Dobzhansky praised in circular overlaps is found in biological systems currently described as ‘rings of species’, in addition to the idealized ‘ring species’.     

Divergent selection and drift shape the genomes of two avian sister species spanning a saline–freshwater ecotone

The role of species divergence due to ecologically based divergent selection—or ecological speciation—in generating and maintaining biodiversity is a central question in evolutionary biology. Comparison of the genomes of phylogenetically related taxa spanning a selective habitat gradient enables discovery of divergent signatures of selection and thereby provides valuable insight into the role of divergent ecological selection in speciation. Tidal marsh ecosystems provide tractable opportunities for studying organisms' adaptations to selective pressures that underlie ecological divergence. Sharp environmental gradients across the saline–freshwater ecotone within tidal marshes present extreme adaptive challenges to terrestrial vertebrates. Here, we sequence 20 whole genomes of two avian sister species endemic to tidal marshes—the saltmarsh sparrow (Ammospiza caudacutus) and Nelson's sparrow (A. nelsoni)—to evaluate the influence of selective and demographic processes in shaping genome‐wide patterns of divergence. Genome‐wide divergence between these two recently diverged sister species was notably high (genome‐wide F_ST = 0.32). Against a background of high genome‐wide divergence, regions of elevated divergence were widespread throughout the genome, as opposed to focused within islands of differentiation. These patterns may be the result of genetic drift resulting from past tidal march colonization events in conjunction with divergent selection to different environments. We identified several candidate genes that exhibited elevated divergence between saltmarsh and Nelson's sparrows, including genes linked to osmotic regulation, circadian rhythm, and plumage melanism—all putative candidates linked to adaptation to tidal marsh environments. These findings provide new insights into the roles of divergent selection and genetic drift in generating and maintaining biodiversity.