Determinants of local recruitment in a growing colony of Audouin's gull

1. Local recruitment of Audouin's gull ( Larus audouinii Payraudeau) was studied between 1988 and 1997 at the Ebro Delta colony (north-western Mediterranean). Since its establishment in 1981, the colony has dramatically grown to include, in 1997, 65% of the total world population. Several hypotheses were tested, involving the effects of a badger predatory event in 1994, and sex, age and cohort (year of birth) on recruitment.
2. Results supported the prediction that colony size influenced recruitment: the probability for any individual to have previously bred increased throughout the study, together with colony size. At the end of the study, 90% of breeders aged 4 years had already been recruited at age 3, the age of first reproduction by Audouin's gulls. As expected by the dramatic increase of breeding numbers, most local recruitment occurred at very young ages, especially when compared with other Laridae.
3. Neither food availability nor reproductive success affected recruitment. Recruitment was not affected by high nest predation by the badger, although after the event, the proportion of Ebro Delta birds nesting on the nearby Columbretes Islands tripled.
4. Probability of first reproduction depended on age: it was the highest at ages 3 and 4, and then decreased sharply with age to stabilize beyond age 6 to a value depending on the year and cohort but always very low (< 5%). Cohort and sex did not influence local recruitment.
5. Annual resighting rates ranged between 35% and 82%, and were higher for females. This may represent a sex-dependent suspension of breeding, probably as a trade-off between early recruitment and future survival.     

Natal philopatry does not lead to population genetic differentiation in Buller's albatross (Thalassarche bulleri bulleri)

Genetic variability in the only two existing populations of Buller's albatross (Thalassarche bulleri bulleri) was assessed using six polymorphic microsatellite loci. Large biological samples were obtained from both the Snares (n = 99) and the Solander Islands (n = 27). Several measures of genetic differentiation including F(ST) and R(ST) and a principal coordinates analysis (PCO) suggest a complete absence of genetic structure between three breeding colonies on the Snares Islands, and between them and one breeding colony on the Solander Islands. Mark/recapture studies of recently banded albatross chicks on the Snares found high natal philopatry in T. b. bulleri, but the microsatellite DNA data suggest that sufficient gene flow still occurs between all four breeding colonies to maintain a genetically homogeneous population overall.     

Diet of Yellow-legged Gull ( Larus michahellis ) chicks along the Spanish Western Mediterranean coast: the relevance of refuse dumps

In recent decades, the Yellow-legged Gull (Larus michahellis) has become a problematic species in many Mediterranean countries, mainly because it interferes with human interests. However, this gull also has a negative impact on several other bird species, many of which are classified as endangered. Two different European Union Action Plans are currently under development with the aim of decreasing the availability of food derived from human activities, such as garbage and fishery discards, which are considered to be the main causes of the superpopulations of this gull. Here, we describe the diet of Yellow-legged Gull chicks, with particular emphasis on establishing the dependence of each population on refuse dumps, in order to forecast changes in gull population dynamics in response to the management decisions being implemented. We sampled four colonies along the Western Mediterranean in Spain: the Medes Islands, the Ebro Delta, the Columbretes Islands, and Mazarrón Island. To elucidate their feeding ecology and to avoid obtaining a discrete estimation from a single sampling, we collected regurgitates from each colony three times throughout the chick-rearing period. Slightly differential feeding habits were observed between chick age classes. Younger chicks in all four colonies tended to be consistently provisioned with smaller prey such as invertebrates. Distinct uses of several foraging habitats among localities were observed. In particular, the use of refuse dumps was common and abundant in two of the colonies: the Medes and Mazarrón Islands. As a consequence of current management strategies, generalized reductions in Yellow-legged Gull populations and increases in the consumption of alternative food resources to those of fishery discards and refuse scraps are expected. Finally, we predict that decreased food availability will force some gulleries to increase predation on endangered species, thereby raising a conservation concern. X. Ruiz: deceased on 27 April 2008.     

Body size variation in Audouin's Gull Larus audouinii; a density-dependent effect?

A significant decrease in the body size of Audouin's Gulls Larus audouinii breeding at the Chafarinas Islands is reported. The decrease in linear measurements in the current breeding population ranged from 2.5% to 5.6% in males and from 0.61% to 4.4% in females. This was detected when assessing the reliability of a sex-discriminating function derived for the same colony 13 years earlier. When applied to the current population, this function failed to predict the sex of a large proportion of males (44%). The relative decrease in mean size was significantly greater in males than in females for culmen, nalospi and tarsus lengths, while for bill depth at culmen, wing length and body mass the relative decrease was similar in both sexes. Since the extent of differences depended on sex, these differences cannot be attributed to a systematic between-observer bias. Mean body size reduction might be either the result of a greater proportion of small breeding birds in the current population, because of increased availability of nesting sites (competition relaxation hypothesis), or an outcome of environmental factors affecting growth parameters (environmental constraint hypothesis). According to the first hypothesis, the changes observed would be associated with higher variability values. Conversely, if the second is true, the degree of variability should be similar. Since there are no significant differences in the degree of variability shown in the two data sets, our results support the second hypothesis. The environmental constraint acting via growth parameters is probably related to the increase in the number of Audouin's Gull breeding pairs while food availability was depleted. Our data suggest that changes in the duration of the growth period, rather than in the growth rates themselves, are involved in the body size differences found.     

Composition of chick meals from one of the main little auk (Alle alle) breeding colonies in Northwest Greenland

During the last decade, increasing information on little auk (Alle alle) biology, ecology and behaviour has been reported. However, only a few of these studies have focused on the breeding population in the Avanersuaq (Thule) district of Northwest Greenland, where 80 % of the global little auk population is estimated to breed. This study reports on the chick diet composition from one of the largest colonies, the Paakitsoq colony, located on the south-eastern margin of the North Water (NOW) Polynya. Results revealed the highest proportion of Calanus hyperboreus, a large lipid-rich copepod, in chick diet reported for any little auk colony. Results confirmed that the cold, highly productive waters of the NOW Polynya are favourable foraging grounds for the little auks during the breeding season. Species diversity within and between the chick meals was low, which probably reflects a high availability of a few preferred prey species. Individual chick meals were generally low in number of prey items and total energy content compared with other published results. This may be explained by a higher feeding frequency or by the samples being collected late in the breeding period (during late chick rearing), when chicks have a reduced growth rate and may require less energy than at earlier developmental stages.     

Small‐scale spatial variation in evolvability for life‐history traits in the storm petrel

The evolutionary potential in the timing of recruitment and reproduction may be crucial for the ability of populations to buffer against environmental changes, allowing them to avoid unfavourable breeding conditions. The evolution of a trait in a local population is determined by its heritability and selection. In the present study, we performed pedigree‐based quantitative genetic analyses for two life‐history traits (recruiting age and laying date) using population data of the storm petrel over an 18‐year period in two adjacent breeding colonies (only 150 m apart) that share the same environmental conditions. In both traits, natal colony effect was the main source of the phenotypic variation among individuals, and cohort variance for recruitment age and additive genetic variance for laying date were natal colony‐specific. We found significant heritability only in laying date and, more specifically, only in birds born in one of the colonies. The difference in genetic variance between the colonies was statistically significant. Interestingly, selection on earlier breeding birds was detected only in the colony in which heritable variation in laying date was found. Therefore, local evolvability for a life‐history trait may vary within a unexpectedly small spatial scale, through the diversifying natural selection and insulating gene flow. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, 106 , 439–446.     

Chinstrap penguins (Pygoscelis antarctica) nesting on Sabrina Islet, Balleny Islands, Antarctica

Although the majority of chinstrap penguins (Pygoscelis antarctica) inhabit the Scotia Arc, a minuscule population of ten breeding pairs has been reported several times from Chinstrap Islet, in the Balleny Islands, 5,000 km distant on the opposite side of Antarctica. An aerial photographic census (December 2000) reveals that the overall penguin population of the Balleny Islands has declined by 8% since the last census 16 years ago, while the combined populations of the two largest colonies, Chinstrap Islet and Sabrina Islet, have increased by 11%. A ground visit confirmed for the first time that chinstrap penguins also breed on Sabrina Islet, occupying 20-24 nests on the margins of the Adélie colony (3,790 pairs in all). The small size and anomalous location of the Balleny Islands chinstrap population clearly warrant further study and the strongest protective measures.     

Morphological and genetic variability of Snow Petrels Pagodroma nivea

Snow Petrels vary greatly in size, the largest birds being over twice as heavy as the smallest. Two morphological types have been distinguished, but the Adelieland population alone shows continuous variation spanning the complete range of the species. There is no evidence of size-related assortative mating. Size variation cannot be attributed to nest site location nor to individual age, nor is the large sexual dimorphism sufficient to account for the situation. The great individual variation in size is accompanied by an important genetic variability as revealed by a starch gel electrophoretic study.
Comparisons of wing length data from many localities around the Antarctic Continent suggest a cline, with a peak at the Balleny Islands. Taking into account the extent of size variation at breeding places only, it appears that morphological variability is low at Balleny Islands, where only large form birds are known, and at Antarctic localities inhabited exclusively by small form birds. In contrast, several localities on the continent in the vicinity of Balleny Islands, and especially Adelieland, exhibit a high degree of variability.
We propose that during the quaternary glaciations, the Antarctic continent provided no refugia for this species, but, when glaciation ended, Snow Petrels re-established colonies, coming from two refugia, a low latitude one inhabited by small birds Pagodroma nivea nivea , and one in higher latitudes characterized by large birds P. n. major. Reproductive isolation broke down, and a hybridization zone was created. The high proportion of polymorphic loci exhibited by the Adelieland population is then attributable to the mixing of two previously isolated genetic pools.     

Colony sex ratios vary with breeding system but not relatedness asymmetry in the facultatively polygynous ant Pheidole pallidula

Abstract.— We investigated sex allocation in a Mediterranean population of the facultatively polygynous (multiple queen per colony) ant Pheidole pallidula . This species shows a strong split sex ratio, with most colonies producing almost exclusively a single-sex brood. Our genetic (microsatellite) analyses reveal that P. pallidula has an unusual breeding system, with colonies being headed by a single or a few unrelated queens. As expected in such a breeding system, our results show no variation in relatedness asymmetry between monogynous (single queen per colony) and polygynous colonies. Nevertheless, sex allocation was tightly associated with the breeding structure, with monogynous colonies producing a male-biased brood and polygynous colonies almost only females. In addition, sex allocation was closely correlated with colony total sexual productivity. Overall, our data show that when colonies become more productive (and presumably larger) they shift from monogyny to polygyny and from male production to female production, a pattern that has never been reported in social insects.     

Satellite remote sensing in monitoring change of seabirds: use of Spot Image in king penguin population increase at Ile aux Cochons,Crozet Archipelago

The king penguin colony at Ile aux Cochons, Crozet Archipelago (southern Indian Ocean) is the largest in the world. The change in its surface area was investigated by means of satellite remote sensing using spot scene. The satellite picture enabled us to locate the colony and to determine that the surface area occupied by the colony has increased by 56% between 1962 and 1988. The increase in the surface area occupied by the colony was confirmed by a map realised in 1982 (by the Institut Géographique National). The high resolution system available on the spot system which allows at present at 10 m ground resolution, and 5 m in the near future, can thus be a useful tool for monitoring medium to long-term changes in marine bird populations breeding in colonies composed of birds spaced a constant distance from one another. Based on the surface area occupied by the colony and the published data on the population size on Possession Island, as well as information on the breeding frequency of the species, we estimated that the total breeding population is about 1,000,000 pairs for the whole archipelago and that about 70% of birds breed at Ile aux Cochons.     

Colony social organization and population genetic structure of an introduced population of formosan subterranean termite from New Orleans, Louisiana

The Formosan subterranean termite, Coptotermes formosanus Shiraki, is an invasive species in many parts of the world, including the U.S. mainland. The reasons for its invasive success may have to do with the flexible social and spatial organization of colonies. We investigated the population and breeding structure of 14 C. formosanus colonies in Louis Armstrong Park, New Orleans, LA. This population has been the focus of extensive study for many years, providing the opportunity to relate aspects of colony breeding structure to previous findings on colony characteristics such as body weight and number of workers, wood consumption, and intercolony aggression. Eight colonies were headed by a single pair of outbred reproductives (simple families), whereas six colonies were headed by low numbers of multiple kings and/or queens that were likely the neotenic descendants of the original colony (extended families). Within the foraging area of one large extended family colony, we found genetic differentiation among different collection sites, suggesting the presence of separate reproductive centers. No significant difference between simple family colonies and extended family colonies was found in worker body weight, soldier body weight, foraging area, population size, or wood consumption. However, level of inbreeding within colonies was negatively correlated with worker body weight and positively correlated with wood consumption. Also, genetic distance between colonies was positively correlated with aggression levels, suggesting a genetic basis to nestmate discrimination cues in this termite population. No obvious trait associated with colony reproductive structure was found that could account for the invasion success of this species.     

Status and structure of the griffon vulture (Gyps fulvus) population in Crete

Griffon vulture (Gyps fulvus) population surveys were conducted during 1996–2002 in the island of Crete (Greece) to document population status and structure. Fieldwork was carried out during the breeding period when birds could be monitored in their colonies. Total population size was estimated at 379 individuals (range = 341–417) with adult birds comprising 63%. The breeding population was estimated at 141 pairs, which were distributed on an average in 23 colonies per year (range = 16–30) while the mean number of breeding pairs that laid eggs was 98 (range= 64–126). Crete thus supports the largest insular population of the species in the world and hosts 70–80% of the breeding population of the species in Greece. Population density was estimated at 6.9 individuals/100 km², 2.6 breeding pairs/100 km² and 1.8 nesting pairs/100 km². The average home range of an occupied colony (i.e., breeding group) was estimated at ca. 204 km² producing a theoretical foraging range of 8 km radius around the breeding cliff. No trends in the total number of individuals and breeding pairs appeared to exist, although significant differences in population size of individual colonies occurred between the years. The majority of the population was concentrated in small-sized colonies, which showed a low occupancy rate. The number of abandoned sites and the colonization of new ones could represent a shift of breeding pairs to alternative colonies provoked by local food abundance and conspesific attraction.     

Variable queen number in ant colonies: no impact on queen turnover, inbreeding, and population genetic differentiation in the ant Formica selysi

Variation in queen number alters the genetic structure of social insect colonies, which in turn affects patterns of kin-selected conflict and cooperation. Theory suggests that shifts from single- to multiple-queen colonies are often associated with other changes in the breeding system, such as higher queen turnover, more local mating, and restricted dispersal. These changes may restrict gene flow between the two types of colonies and it has been suggested that this might ultimately lead to sympatric speciation. We performed a detailed microsatellite analysis of a large population of the ant Formica selysi, which revealed extensive variation in social structure, with 71 colonies headed by a single queen and 41 by multiple queens. This polymorphism in social structure appeared stable over time, since little change in the number of queens per colony was detected over a five-year period. Apart from queen number, single- and multiple-queen colonies had very similar breeding systems. Queen turnover was absent or very low in both types of colonies. Single- and multiple-queen colonies exhibited very small but significant levels of inbreeding, which indicates a slight deviation from random mating at a local scale and suggests that a small proportion of queens mate with related males. For both types of colonies, there was very little genetic structuring above the level of the nest, with no sign of isolation by distance. These similarities in the breeding systems were associated with a complete lack of genetic differentiation between single- and multiple-queen colonies, which provides no support for the hypothesis that change in queen number leads to restricted gene flow between social forms. Overall, this study suggests that the higher rates of queen turnover, local mating, and population structuring that are often associated with multiple-queen colonies do not appear when single- and multiple-queen colonies still coexist within the same population, but build up over time in populations consisting mostly of multiple-queen colonies.     

Genetic analysis of the breeding system of an invasive subterranean termite, Reticulitermes santonensis, in urban and natural habitats

Reticulitermes santonensis is a subterranean termite that invades urban areas in France and elsewhere where it causes damage to human-built structures. We investigated the breeding system, colony and population genetic structure, and mode of dispersal of two French populations of R. santonensis. Termite workers were sampled from 43 and 31 collection points, respectively, from a natural population in west-central France (in and around the island of Oleron) and an urban population (Paris). Ten to 20 workers per collection point were genotyped at nine variable microsatellite loci to determine colony identity and to infer colony breeding structure. There was a total of 26 colonies, some of which were spatially expansive, extending up to 320 linear metres. Altogether, the analysis of genotype distribution, F-statistics and relatedness coefficients suggested that all colonies were extended families headed by numerous neotenics (nonwinged precocious reproductives) probably descended from pairs of primary (winged) reproductives. Isolation by distance among collection points within two large colonies from both populations suggested spatially separated reproductive centres with restricted movement of workers and neotenics. There was a moderate level of genetic differentiation (F(ST) = 0.10) between the Oleron and Paris populations, and the number of alleles was significantly higher in Oleron than in Paris, as expected if the Paris population went through bottlenecks when it was introduced from western France. We hypothesize that the diverse and flexible breeding systems found in subterranean termites pre-adapt them to invade new or marginal habitats. Considering that R. santonensis may be an introduced population of the North American species R. flavipes, a breeding system consisting primarily of extended family colonies containing many neotenic reproductives may facilitate human-mediated spread and establishment of R. santonensis in urban areas with harsh climates.     

Contrasting bottom-up effects of warming ocean on two king penguin populations

Breeding success is often correlated with climate, but the underlying bottom-up mechanisms remain elusive—particularly in marine environments. Consequently, conservation plans of many species often consider climate change as a unilateral threat, ignoring that even nearby populations can show contradicting trends with climate. Better understanding the relationship between climate and environment at different scales can help us interpret local differences in population trends, ultimately providing better tools to evaluate the global response of a species to threats such as global warming. We studied a growing king penguin population nesting at Kerguelen island (Southern Indian Ocean), hosting one of the largest colonies in the world. We used a unique dataset of foraging, breeding success, and climate data spanning over 25 years to examine the links between climate, marine environment, and breeding success at this colony. The results were then compared to the neighboring population of Crozet, which experienced the steepest decline for this species over the past few decades. At Crozet, penguins experienced lower breeding success in warmer years due to productive currents shifting away from the colony, affecting foraging behavior during chick rearing. At Kerguelen, while chick mass and survival experienced extreme variation from year to year, the annual variation was not associated with the position of the currents, which varied very little compared to the situation in Crozet. Rather than being affected by prey distribution shifts, we found evidence that chick provisioning in Kerguelen might be influenced by prey abundance, which seem to rather increase in warmer conditions. Furthermore, warmer air temperature in winter increased chick survival rate, likely due to reduced thermoregulation cost. Investigating the mechanisms between climate and fitness allowed us to predict two different fates for these populations regarding ongoing global warming.     

Impacts of human visitors on breeding success and long-term population trends in Adélie Penguins at Casey,Antarctica

Breeding populations of Aélie Penguins Pygoscelis adeliae have been counted at two localities near Casey Station in Wilkes Land, East Antarctica since 1959/60 and 1968/69. At Whitney Point, the breeding population increased from 1122 pairs in 14 colonies in 1959/60 to 4714 pairs in 36 colonies in 1992/93. All new colonies at Whitney Point established on relict colony sites identified in 1959/60. On Shirley Island, the total breeding population has remained at 7770 pairs ±10% between 1968/69 and 1992/93, except in 1990/91 when the population peaked at 8719 pairs. An association between the age of a colony and its rate of increase was observed at Whitney Point, with new colonies (those established since 1971/72) increasing more rapidly than colonies extant in 1959/60. At Shirley Island, where most of the colonies extant in 1968/69 have decreased in population, the establishment and growth of 13 colonies has offset this decrease; these new colonies also exhibited the association between age and rate of increase. Breeding success (chicks fledged per nest) was significantly lower for Shirley Island colonies than for those at Whitney Point. Human visitors to Shirley Island from Casey station are believed to be responsible for the observed changes in the distribution and abundance of breeding pairs and for maintaining the stable population by reducing overall breeding success through the disturbance associated with visits.     

Unique and isolated: population structure has implications for management of the endangered New Zealand sea lion

Female otariids (eared seals) frequently display strong levels of philopatry, a behaviour that has the potential to influence population structure, particularly at the mitochondrial level. Conversely, male otariids often move between breeding colonies, likely facilitating nuclear gene flow between colonies. Such gender-specific movements have the potential to influence species population structure. Here we investigate the genetic population structure of the endangered New Zealand (NZ) sea lion, using nuclear (microsatellite) and mitochondrial molecular markers, with the intention to better inform conservation through identification of management units for the species. The strong levels of female philopatry in this species have potential to lead to population structure at the mitochondrial loci. In contrast, weak or no population structure is expected across nuclear loci. NZ sea lions were sampled from the main breeding areas across the species’ current distribution (three Auckland Islands sites, two Campbell Island sites, one Stewart Island site and one Otago Peninsula site). Individuals were screened for microsatellite (n?=?271; 16 loci) and mitochondrial (n?=?56; 1027 bp D-loop and 1189 bp cytb). Despite a small (c. 9880 individuals) population size, moderate levels of microsatellite variation are observed in the NZ sea lions, in contrast to low levels of mitochondrial genetic variation. Results from mitochondrial DNA analyses revealed no population structure, suggesting that the strong level of female philopatry in NZ sea lions alone is not sufficient to maintain genetic population structure. Due to the frequent male movements between breeding colonies, no population structure was detected across the nuclear loci either. The absence of genetic structure suggests that, from a genetic perspective, NZ sea lions can be considered to be a single population. Despite this, the differing impacts of threats (e.g. fisheries by-catch) to each individual breeding colony must also be taken into consideration when defining management units for this endangered species.     

The role of despotism and heritability in determining settlement patterns in the colonial lesser kestrel

Avian colony size variation is an evolutionary puzzle in terms of unequal fitness payoffs. We used a long-term marked lesser kestrel (Falco naumanni) population, where individual fitness increases with colony size, to test whether subordinates are evicted despotically from the largest colonies. Yearlings were smaller and lighter, were more attacked than expected, and lost most disputes over nest holes with older birds. Agonistic interactions increased with colony size; consequently, most first breeders recruited in colonies smaller than those at which they first tried to settle. As expected when subordination is a transient state, birds dispersed to a larger colony as they got older even after breeding successfully. The population consequences of these behavioral processes were that the relative frequency of yearlings and first breeders decreased with colony size. At the same time, breeding colony size was repeatable within individuals, so we estimated the amount of heritable variation in this trait. Estimates of heritability derived from parent-offspring and full-sib analyses were consistently high (h2=0.53) when individuals reached asymptotic morphological values and presumably overcame subordinate transient states. Age-related dominance asymmetries masked resemblance among relatives in colony size, but both phenomena seem to coexist in this population and explain a considerable proportion of colony size variation.     

An American termite in Paris: temporal colony dynamics

Termites of the genus Reticulitermes are widespread invaders, particularly in urban habitats. Their cryptic and subterranean lifestyle makes them difficult to detect, and we know little about their colony dynamics over time. In this study we examined the persistence of Reticulitermes flavipes (Kollar) colonies in the city of Paris over a period of 15 years. The aim was (1) to define the boundaries of colonies sampled within the same four areas over two sampling periods, (2) to determine whether the colonies identified during the first sampling period persisted to the second sampling period, and (3) to compare the results obtained when colonies were delineated using a standard population genetic approach versus a Bayesian clustering method that combined both spatial and genetic information. Herein, colony delineations were inferred from genetic differences at nine microsatellite loci and one mitochondrial locus. Four of the 18 identified colonies did not show significant differences in their genotype distributions between the two sampling periods. While allelic richness was low, making it hard to reliably distinguish colony family type, most colonies appeared to retain the same breeding structure over time. These large and expansive colonies showed an important ability to fuse (39% were mixed-family colonies), contained hundreds of reproductives and displayed evidence of isolation-by-distance, suggesting budding dispersal. These traits, which favor colony persistence over time, present a challenge for pest control efforts, which apply treatment locally. The other colonies showed significant differences, but we cannot exclude the possibility that their genotype distributions simply changed over time.     

Molecular insight into patterns of colony composition and paternity in the common mole-rat Cryptomys hottentotus hottentotus

We report the discovery of intraspecific variation in both colony composition and patterns of paternity in two populations of the social common mole-rat Cryptomys hottentotus hottentotus. These two populations represent the mesic and arid habitat extremes of the species' broad ecological range in South Africa. Until recently colonies of the common mole-rat were thought to consist of familial groups whereby all colony members were the offspring of a monogamous reproductive pair. The remaining colony members were thought to forego reproduction until both social and ecological conditions favoured dispersal and opportunities for independent outbreeding. Results from genetic assignment tests using microsatellite markers indicate that while colony composition is dominated by familial groups, colonies within both populations included both adult and subadult foreign conspecifics. Analysis of parentage reveals that the social organization of C. h. hottentotus is not that of strict monogamy; paternity of offspring was not assigned consistently to the largest, most dominant male within the colony. Moreover, a number of significantly smaller males were found to sire offspring, suggesting a sneak-mating strategy by subordinate within-colony males. Extra-colony extra-pair paternity (ECP) was also found to characterize C. h. hottentotus colonies, occurring with similar frequencies in both habitats. Both dominant established breeding males and subordinate males were identified as siring young in nonsource colonies. Furthermore, established breeding males were found to sire extra-colony young in the same season as siring young within their source colonies. We discuss the significance of these results within the context of the divergent ecological regimes characterizing the two sites and observe that our results revisit the accuracy of using behavioural and morphological characters, which have structured the basis of our understanding of the behavioural ecology of this species, as indicators of breeding status in mark-recapture studies.