Both natural selection and isolation by distance explain phenotypic divergence in bill size and body mass between South Australian little penguin colonies

Morphological variation between populations of the same species can arise as a response to genetic variation, local environmental conditions, or a combination of both. In this study, I examined small‐scale geographic variation in bill size and body mass in little penguins (Eudyptula minor) across five breeding colonies in South Australia separated by <150 km. To help understand patterns driving the differences, I investigated these variations in relation to environmental parameters (air temperature, sea surface temperature, and water depth) and geographic distances between the colonies. I found substantial morphological variation among the colonies for body mass and bill measurements (except bill length). Colonies further located from each other showed greater morphological divergence overall than adjacent colonies. In addition, phenotypic traits were somewhat correlated to environmental parameters. Birds at colonies surrounded by hotter sea surface temperatures were heavier with longer and larger bills. Birds with larger and longer bills were also found at colonies surrounded by shallower waters. Overall, the results suggest that both environmental factors (natural selection) and interpopulation distances (isolation by distance) are causes of phenotypic differentiation between South Australian little penguin colonies.     

Biogeographical basis of recent phenotypic divergence among birds: a global study of subspecies richness

Theory predicts that biogeographic factors should play a central role in promoting population divergence and speciation. Previous empirical studies into biogeography and diversification have been relatively restricted in terms of the geographical area, phylogenetic scope, and the range of biogeographic factors considered. Here we present a global analysis of allopatric phenotypic divergence (measured as subspecies richness) across more than 9600 bird species. The main aim of this study was to examine the extent to which biogeographical factors can explain patterns of phenotypic divergence. Analysis of the taxonomic distribution of subspecies among species suggests that subspecies formation and extinction have occurred at a considerably faster rate than has species formation. However, the observed distribution departs from the expectation under a random birth-death model of diversification. Across 19 phylogenetic trees, we find no significant linear relationship between species age and subspecies richness, implying that species age is a poor predictor of subspecies richness. Both subspecies richness and subspecies diversification rate are found to exhibit low phylogenetic signal, meaning that closely related species do not tend to possess similar numbers of subspecies. As predicted by theory, high subspecies richness was associated with large breeding range size, island dwelling, inhabitation of montane regions, habitat heterogeneity, and low latitude. Of these factors, breeding range size was the variable that explained the most variation. Unravelling whether species that have invaded previously glacial areas have more or fewer subspecies than expected proves to be complicated due to a covariation between the postglacial colonization, latitude, geographic range size, and subspecies richness. However, the effect of postglacial colonization on subspecies richness appears to be small. Mapping the distribution of species' subspecies richness globally reveals geographical patterns that correspond to many of the predictions of the statistical models, but may also reflect geographical variation in taxonomic practice. Overall, we demonstrate that biogeographic models can explain about 30% of the global variation in subspecies richness in birds.     

Weak large‐scale population genetic structure in a philopatric seabird,the European Shag Phalacrocorax aristotelis

Quantifying population genetic structure is fundamental to testing hypotheses regarding gene flow, population divergence and dynamics across large spatial scales. In species with highly mobile life‐history stages, where it is unclear whether such movements translate into effective dispersal among discrete philopatric breeding populations, this approach can be particularly effective. We used seven nuclear microsatellite loci and mitochondrial DNA (ND2) markers to quantify population genetic structure and variation across 20 populations (447 individuals) of one such species, the European Shag, spanning a large geographical range. Despite high breeding philopatry, rare cross‐sea movements and recognized subspecies, population genetic structure was weak across both microsatellites and mitochondrial markers. Furthermore, although isolation‐by‐distance was detected, microsatellite variation provided no evidence that open sea formed a complete barrier to effective dispersal. These data suggest that occasional long‐distance, cross‐sea movements translate into gene flow across a large spatial scale. Historical factors may also have shaped contemporary genetic structure: cluster analyses of microsatellite data identified three groups, comprising colonies at southern, mid‐ and northern latitudes, and similar structure was observed at mitochondrial loci. Only one private mitochondrial haplotype was found among subspecies, suggesting that this current taxonomic subdivision may not be mirrored by genetic isolation.     

Genetic variation and bill size dimorphism in a passerine bird, the reed bunting Emberiza schoeniclus

In passerine birds morphological differentiation in bill size within species is not commonly observed. Bill size is usually associated with a trophic niche, and strong differences in it may reflect the process of genetic differentiation and, possibly, speciation. We used both mitochondrial DNA (mtDNA) and nuclear microsatellites to study genetic variation between two subspecies of reed bunting, Emberiza schoeniclus schoeniclus and E.s. intermedia , along their distributional boundary in western Europe. These two subspecies are characterized by a high dimorphism in bill size and, although breeding populations of the two subspecies are found very close to each other in northern Italy, apparently no interbreeding occurs. The observed morphological pattern between the two subspecies may be maintained by geographically varying selective forces or, alternatively, may be the result of a long geographical separation followed by a secondary contact. MtDNA sequences of cytochrome b and ND5 (515 bp) showed little variation and did not discriminate between the two subspecies, indicating a divergence time of less than 500 000 years. The analysis of four microsatellite loci suggested a clear, although weak, degree of genetic differentiation in the large- and small-billed populations, as indicated by F _ST and R _ST values and genetic distances. The correlation between bill size and genetic distance between populations remained significant after accounting for the geographical distances between sampling localities. Altogether, these results indicate a very recent genetic differentiation between the two bill morphs and suggest that a strong selection for large bills in the southern part of the breeding range is probably involved in maintaining the geographical differentiation of this species.     

Local and large-scale climatic variables as predictors of the breeding numbers of endangered Lesser Black-backed Gulls on the Norwegian Coast

In the 1970s and 1980s, the nominate subspecies of the Lesser Black-backed Gull (Larus fuscus fuscus) showed a dramatic drop in breeding numbers on the Norwegian Coast, and in 2000, the population in some colonies was only 10–20% of the population in 1980. This decline has been attributed to the collapse in the stock of Norwegian spring spawning herring (Clupea harengus). In this study, we examined whether local climate (sea and air temperatures), winter NAO (North Atlantic Oscilliation), and the year-class strength and size of 0-group herring could predict the relative changes in breeding numbers between years, mainly after the population collapse. Breeding birds were counted in 19 of the years between 1980 and 2007 in an archipelago on the coast of Helgeland, northern Norway. The best model predicting changes in breeding numbers for the period between 1980 and 2005 (for which data on 0-group herring was available) included mean local air temperature in winter (January–March) and winter NAO, explaining 57% of the variation between years, while the other factors had little effect. When also adding the years 2006–2007 (no herring data), the best model included only mean air temperature in winter, explaining 41% of the variation. In conclusion, the high positive correlation between breeding numbers and climatic factors probably resulted from a higher availability of important fish prey after mild winters, for which 0-group herring presently may only account for a limited proportion. However, this prey might have been of much more importance prior to the population decline.     

Spatial predictors of genomic and phenotypic variation differ in a lowland Middle American bird (Icterus gularis)

Spatial patterns of intraspecific variation are shaped by geographical distance among populations, historical changes in gene flow and interactions with local environments. Although these factors are not mutually exclusive and operate on both genomic and phenotypic variation, it is unclear how they affect these two axes of variation. We address this question by exploring the predictors of genomic and phenotypic divergence in Icterus gularis, a broadly distributed Middle American bird that exhibits marked geographical variation in body size across its range. We combined a comprehensive single nucleotide polymorphism and phenotypic data set to test whether genome‐wide genetic and phenotypic differentiation are best explained by (i) isolation by distance, (ii) isolation by history or (iii) isolation by environment. We find that the pronounced genetic and phenotypic variation in I. gularis are only partially correlated and differ regarding spatial predictors. Whereas genomic variation is largely explained by historical barriers to gene flow, phenotypic diversity can be best predicted by contemporary environmental heterogeneity. Our genomic analyses reveal strong phylogeographical structure coinciding with the Chivela Pass at the Isthmus of Tehuantepec that was formed during the Pleistocene, when populations were isolated in north–south refugia. In contrast, we found a strong association between body size and environmental variables, such as temperature and precipitation. The relationship between body size and local climate is consistent with a pattern produced by either natural selection or environmental plasticity. Overall, these results provide empirical evidence for why phenotypic and genomic data are often in conflict in taxonomic and phylogeographical studies.     

Body condition and feather molt of a migratory shorebird during the non‐breeding season

Migratory shorebirds have some of the highest fat loads among birds, especially species which migrate long distances. The upland sandpiper Bartramia longicauda makes long‐distance migrations twice a year, but variation in body condition or timing of feather molt during the non‐breeding season has not been studied. Molt is an important part of the annual cycle of migratory birds because feather condition determines flight performance during migration, and long‐distance movements are energetically costly. However, variation in body condition during molt has been poorly studied. The objective of our field study was to examine the timing and patterns of feather molt of a long distance migratory shorebird during the non‐breeding season and test for relationships with body size, fat depots, mass, and sex. Field work was conducted at four ranches in the Northern Campos of Uruguay (Paysandú and Salto Departments). We captured and marked 62 sandpipers in a 2‐month period (Nov–Jan) during four non‐breeding seasons (2008–2012). Sex was determined by genetic analyses of blood samples taken at capture. Molt was measured in captured birds using rank scores based on published standards. Body mass and tarsus length measurements showed female‐biased sexual size dimorphism with males smaller than females. Size‐corrected body mass (body condition) showed a U‐shaped relationship with the day of the season, indicating that birds arrived at non‐breeding grounds in relatively good condition. Arriving in good body condition at non‐breeding grounds is probably important because of the energetic demands due to physiological adjustments after migration and the costs of feather molt.     

Growth of Stonechats Saxicola torquata from Africa and Europe: an analysis of genetic and environmental components

The postnatal growth of the European subspecies of the Stonechat Saxicola torquata rubicola and the African subspecies Saxicola torquata axillaris and their hybrids was studied to investigate differences in growth rates. Comparisons were made between subspecies, between genetically related chicks within subspecies and between individuals within sibling groups. Genetic and environmental effects on variation and their impact on the growth curves were evaluated. We compared posthatching increase in body mass and wing-length between chicks raised under identical conditions and between chicks raised under different photoperiod and rearing conditions. Significant differences in body mass increase between the two subspecies were found for all parameters except for the inflection point of the growth curve. Differences in wing-length occurred during the late posthatching development and were significant only for asymptotic size. However, interactions between independent variables masked the effects on some parameters. Furthermore, our data showed that the components of the total genetic variability are differently attributed to each level of comparison: we found the form of the growth curve and asymptotic size significantly different between subspecies; size at a given day and asymptotic size were genetically different between sibling groups within subspecies; age at the inflection point of the growth curve showed most variability between individuals within groups of related siblings. Cross-breeding experiments showed that the growth parameters in the hybrids were intermediate, providing further evidence for a genetic basis of the observed differences. In this particular case, the between-subspecies variability reflects genetically adjusted adaptation rather than phenotypic responses to different ecological conditions. The phenotypic responses of growth to differences in “environment” have been investigated in groups of birds reared under different conditions (hand-reared v parent-reared chicks; European v equatorial photoperiod). The rearing method affected only size parameters and not the form of the growth curve of body mass. Significant differences were found for maximum body mass, final body mass and asymptote. Photoperiod had no effect on growth.     

Status of the endangered ivory gull, <Emphasis Type="Italic">Pagophila eburnea</Emphasis>, in Greenland

The ivory gull, a rare high-Arctic species whose main habitat throughout the year is sea ice, is currently listed in Greenland as ‘Vulnerable’, and as ‘Endangered’ in Canada, where the population declined by 80% in 20 years. Despite this great concern, the status of the species in Greenland has been largely unknown as it breeds in remote areas and in colonies for which population data has rarely, if at all, been collected. Combining bibliographical research, land surveys, aerial surveys and satellite tracking, we were able to identify 35 breeding sites, including 20 new ones, in North and East Greenland. Most colonies are found in North Greenland and the largest are located on islands and lowlands. The current best estimate for the size of the Greenland population is approx. 1,800 breeding birds, but the real figure is probably >4,000 adult birds (i.e. >2,000 pairs) since all colonies have not yet been discovered and since only 50% or less of the breeding birds are usually present in the colonies at the time the censuses take place. Although this estimate is four to eight times higher than that previously arrived at, the species seems to be declining in the south of its Greenland breeding range, while in North Greenland the trends are unclear and unpredictable, calling for increased monitoring efforts.     

When Bergmann's rule fails: evidences of environmental selection pressures shaping phenotypic diversification in a widespread seabird

Organisms tend to exhibit phenotypes that can be shaped by climate, commonly demonstrating clinal variations along latitudinal gradients. In vertebrates, air temperature plays a major role in shaping body size in both ectothermic and endothermic animals. However, additional small‐scale environmental factors can also act as selection pressures in the marine ecosystem (e.g. primary productivity), evidencing multi‐scale processes acting on marine organisms. In this study, we tested Bergmann's rule in a widely distributed seabird, the brown booby Sula leucogaster, in addition to evaluating the relationship of sea surface temperature and chlorophyll α with phenotypes. We used traits from a morphometric dataset (culmen, wing chord, and tarsus length) and body mass of 276 brown boobies distributed on six breeding sites along a latitudinal gradient in the South Atlantic Ocean (0–27°S). We found significant differentiation among colonies, but phenotypic similarities were observed between colonies located at the extremes of the latitudinal gradient. As the colony nearest to the Equator, Saint Peter and Saint Paul archipelago, had the largest and heaviest individuals, the model containing only air temperature explained < 5% of the allometric variation, providing no substantial support for Bergmann's rule. However, when we added the interaction of chlorophyll α and sea surface temperature the deviance explained rose to over 80%. Primary productivity and sea surface temperature do not follow a latitudinal gradient in the ocean and, therefore, the role of small‐scale oceanographic processes in shaping body size and the importance of considering additional environmental variables when testing Bergmann's rule in marine organisms are evident.     

Weak population genetic differentiation in the most numerous Arctic seabird,the little auk

Quantifying patterns of genetic diversity and differentiation among populations of Arctic birds is fundamental for understanding past and ongoing population processes in the Arctic. However, the genetic differentiation of many important Arctic species remains uninvestigated. Here, phylogeography and population genetics were examined in the worldwide population of a small seabird, the little auk (dovekie, Alle alle)—the most numerous avian species of the Arctic ecosystem. Blood samples or feathers were collected from 328 little auks (325 from the nominate subspecies and 3 from the A. a. polaris) in nine main breeding aggregations in the northern Atlantic and one location from the Pacific Ocean. The mtDNA haplotypes of the two subspecies were not segregated into separate groups. Also, no genetic structure was found within the nominate race based on microsatellite markers. The level of genetic differentiation among populations was low yet significant (mean F _ST = 0.005). Some pairwise F _ST comparisons revealed significant differences, including those involving the most distant Pacific colony as well as among some Atlantic populations. Weak population differentiation following the model of isolation by distance in the little auk is similar to the patterns reported in other high-Arctic bird species, indicating that a lack of distinct genetic structure is a common phenomenon in the Arctic avifauna.     

Speciation through temporal segregation of Madeiran storm petrel (Oceanodroma castro) populations in the Azores?

Madeiran storm petrels Oceanodroma castro breed on three small islets in the Azores: Vila, off Santa Maria, and Praia and Baixo, off Graciosa. Analysis of data on brood patch, incubation periods, chick body size and recaptures of adults provides evidence of the existence of two distinct populations (hot- and cool-season) breeding annually on Baixo and Praia, out of phase by four to five months and overlapping in colony attendance during August and early September; on Vila only the cool-season population is present. Analyses of adult morphology indicate highly significant phenotypic differentiation between the sympatric hot- and cool-season breeders, whereas an almost complete phenotypic uniformity characterizes allopatric breeders within the same season. The hot-season birds are 10% smaller in egg and body mass but have longer wings and tails than cool-season birds. The two groups were readily separated by discriminant analysis. The preference to breed in the cooler season is interpreted as a consequence of greater food availability in that period. Morphological differentiation between seasonal populations is interpreted as an adaptative response to different environmental conditions in the two seasons. The hypothesis is given that the hot-season population has evolved from the cool-season population owing to density-dependent constraints on crowded colonies, forcing birds to time-share nest sites. These populations may represent a case of sympatric speciation through temporal partitioning of reproduction and may be better treated as sibling species.     

Cold tolerance,and not earlier arrival on breeding grounds,explains why males winter further north in an Arctic‐breeding songbird

Sex biases in distributions of migratory birds during the non‐breeding season are widespread; however, the proximate mechanisms contributing to broad‐scale sex‐ratio variation are not well understood. We analyzed a long‐term winter‐banding dataset in combination with spring migration data from individuals tracked by using geolocators to test three hypotheses for observed variation in sex‐ratios in wintering flocks of snow buntings Plectrophenax nivalis. We quantified relevant weather conditions in winter (temperature, snowfall and snow depth) at each banding site each year and measured body size and condition (fat scores) of individual birds (n > 5500). We also directly measured spring migration distance for 17 individuals by using light‐level geolocators. If the distribution pattern of birds in winter is related to interactions between individual body size and thermoregulation, then larger bodied birds (males) should be found in colder sites (body size hypothesis). Males may also winter closer to breeding grounds to reduce migration distance for early arrival at breeding sites (arrival timing hypothesis). Finally, males may be socially dominant over females, and thus exclude females from high‐quality wintering sites (social dominance hypothesis). We found support for the body size hypothesis, in that colder and snowier weather predicted both larger body size and higher proportions of males banded. Direct tracking revealed that males did not winter significantly closer to their breeding site, despite being slightly further north on average than females from the same breeding population. We found some evidence for social dominance, in that females tended to carry more fat than males, potentially indicating lower habitat quality for females. Global climatic warming may reduce temperature constraints on females and smaller‐bodied males, resulting in broad‐scale changes in distributional patterns. Whether this has repercussions for individual fitness, and therefore population demography, is an important area of future research.     

Dynamics of antarctic penguin populations in relation to inter-annual variability in sea ice distribution

To investigate the role of sea ice cover on penguin populations we used principal component analysis to compare population variables of Adélie (Pygoscelis adeliae) and chinstrap (Pygoscelis antarctica) penguins breeding on Signy Island, South Orkney Islands with local (from direct observations) and regional (from remote sensing data) sea ice variables. Throughout the study period, the Adélie penguin population size remained stable, whereas that of chinstrap penguins decreased slightly. For neither species were there significant relationships between population size and breeding success, except for an apparent inverse density-dependent relationship between the number of Adélie breeding pairs and the number of eggs hatching. For both species, no general relationship was found between either population size or breeding success and the local sea ice conditions. However, the regional sea ice extent at a particular time prior to the start of the breeding season was related to the number of birds that arrived to breed. For both species, this period occurred before the sea ice reached its maximum extent and was slightly earlier for Adélie than for chinstrap penguins. These results suggest that sea ice conditions outside the breeding season may play an important role in penguin population processes.     

Pelagic distribution and numbers of the Antarctic petrelThalassoica antarctica in the Weddell Sea during spring

Estimates of the population size of the Antarctic petrelThalassoica antarctica are hampered by its breeding locations in remote nunatak areas of the Antarctic continent. Studies at sea can provide additional information on numbers. Seabird censuses during spring 1992 showed few Antarctic petrels in the western part of the Weddell Sea, but high numbers east of the South Sandwich Islands. The vast majority of the birds occurred in a band of 300 km north to 150 km south of the outer ice edge. Numbers at sea fluctuated in agreement with synchronized colony attendance patterns in the pre-breeding phase and peaked shortly before egglaying when colonies are completely deserted. In this period, densities of Antarctic petrels in the marginal ice zone suggest that at least 2.7±0.5 million individuals occur in the Weddell Sea. It is likely that similar numbers occur immediately east of the study area. The distribution of the petrels matches the location, but not the size, of known breeding colonies in Dronning Maud Land and Coats Land, which suggests that important colonies in this area remain to be discovered. The observations imply that Antarctic petrels in the Weddell Sea commute over ice a surprisingly large distance of at least 2,000 km for a single pre-breeding visit to the colonies.     

Genetic differences and phenotypic plasticity in body size between high- and low-altitude populations of the ground beetle Carabus tosanus

The body size of a univoltine carabid beetle Carabus tosanus on Shikoku Island, Japan, was clearly smaller in higher‐altitude populations (subspecies), which possibly represents incipient speciation. To explore the determinants of altitudinal differences in body size in this species, we studied the degree of phenotypic plasticity by conducting rearing experiments at two constant temperatures and examined genetic differences through interpopulation crosses. At 15 °C, C. tosanus had a longer developmental period and a shorter adult body than at 20 °C. Nevertheless, variation in body size due to temperature effects (phenotypic plasticity) was small compared to the interpopulation differences, which suggests substantial genetic differences between populations (subspecies) at different altitudes. In F₁ offspring from crosses between a low‐altitude (subspecies tosanus) and a high‐altitude population (subspecies ishizuchianus), adult body length was affected by the genotypes of both parents, with an interaction effect of parental genotype and offspring sex. Further analyses revealed that adult body length was affected by sex‐linked factors in addition to autosomal factors. These genetic differences in body size may have resulted from adaptations to different altitudes and may be important for the process of incipient speciation because body size differences could contribute to premating reproductive isolation.     

Gene flow on the ice: genetic differentiation among Adélie penguin colonies around Antarctica

Each summer Adélie penguins breed in large disjunct colonies on ice-free areas around the Antarctic continent. Comprising > 10 million birds, this species represents a dominant feature of the Antarctic ecosystem. The patchy distribution within a large geographical range, natal philopatry and a probable history of refugia, suggest that this species is likely to exhibit significant genetic differentiation within and among colonies. We present data from seven microsatellite DNA loci for 442 individuals from 13 locations around the Antarctic continent. With the exception of one locus, there was no significant genic or genotypic heterogeneity across populations. Pairwise FST values were low with no value > 0.02. When all colonies were compared in a single analysis, the overall FST value was 0.0007. Moreover, assignment tests were relatively ineffective at correctly placing individuals into their respective collection sites. These data reveal a lack of genetic differentiation between Adélie penguin colonies around the Antarctic continent, despite substantial levels of genetic variation. We consider this homogeneity in terms of the dispersal of individuals among colonies and the size of breeding groups and discuss our results in terms of the glacial history of Antarctica.     

Patterns of genetic,phenotypic, and acoustic variation across a chiffchaff (Phylloscopus collybita abietinus/tristis) hybrid zone

Characterizing patterns of evolution of genetic and phenotypic divergence between incipient species is essential to understand how evolution of reproductive isolation proceeds. Hybrid zones are excellent for studying such processes, as they provide opportunities to assess trait variation in individuals with mixed genetic background and to quantify gene flow across different genomic regions. Here, we combine plumage, song, mtDNA and whole‐genome sequence data and analyze variation across a sympatric zone between the European and the Siberian chiffchaff (Phylloscopus collybita abietinus/tristis) to study how gene exchange between the lineages affects trait variation. Our results show that chiffchaff within the sympatric region show more extensive trait variation than allopatric birds, with a large proportion of individuals exhibiting intermediate phenotypic characters. The genomic differentiation between the subspecies is lower in sympatry than in allopatry and sympatric birds have a mix of genetic ancestry indicating extensive ongoing and past gene flow. Patterns of phenotypic and genetic variation also vary between regions within the hybrid zone, potentially reflecting differences in population densities, age of secondary contact, or differences in mate recognition or mate preference. The genomic data support the presence of two distinct genetic clades corresponding to allopatric abietinus and tristis and that genetic admixture is the force underlying trait variation in the sympatric region—the previously described subspecies (“fulvescens”) from the region is therefore not likely a distinct taxon. In addition, we conclude that subspecies identification based on appearance is uncertain as an individual with an apparently distinct phenotype can have a considerable proportion of the genome composed of mixed alleles, or even a major part of the genome introgressed from the other subspecies. Our results provide insights into the dynamics of admixture across subspecies boundaries and have implications for understanding speciation processes and for the identification of specific chiffchaff individuals based on phenotypic characters.     

Revisiting Fisher: range size drives the correlation between variability and abundance of British bird eggs

We evaluate the correlation between intraspecific variation in egg size and population size in breeding British birds. Using information on abundance, range occupancy, migration status and phylogenetic relationships among species, we show that a wider geographical distribution rather than larger population size per se best predicts egg size variability. A similar result applies to wing length variability. Results from a phylogenetic path analysis suggest that geographical variation is the most parsimonious causal explanation for high intraspecific variation in common species.     

Geographical and individual variation in echolocation calls of the intermediate leaf‐nosed bat,Hipposideros larvatus

The cause and significance of variation in echolocation call frequency within hipposiderid bats is not well understood despite an increasing number of allopatric and sympatric examples being documented. We examined variation patterns in the resting frequency (RF) of echolocation calls emitted by the intermediate leaf‐nosed bat, Hipposideros larvatus, on a broad geographical scale. Data mining technology and Kruskal–Wallis test both showed substantial variation with a longitudinal pattern in RF in H. larvatus among colonies, and this variation was associated with geographical distance and not body size. In addition, we found that a high degree of variability between individuals was hidden under the geographical variation. The results support an effect of random cultural drift, and challenge the prey detection hypothesis. Moreover, an acoustic difference among local island colonies may be indicative of a vocal dialect. We found that each colony of H. larvatus seems to maintain a ‘private bandwidth’, which could be used for colony identity and individual communication thus helping individuals and colonies to get a number of fitness benefits.