Sex‐specific prey partitioning in breeding piscivorous birds examined via a novel,noninvasive approach

Piscivorous birds frequently display sex‐specific differences in their hunting and feeding behavior, which lead to diverging impacts on prey populations. Cormorants (Phalacrocoracidae), for example, were previously studied to examine dietary differences between the sexes and males were found to consume larger fish in coastal areas during autumn and winter. However, information on prey partitioning during breeding and generally on sex‐specific foraging in inland waters is missing. Here, we assess sex‐specific prey choice of Great Cormorants (Phalacrocorax carbo) during two subsequent breeding seasons in the Central European Alpine foreland, an area characterized by numerous stagnant and flowing waters in close proximity to each other. We developed a unique, noninvasive approach and applied it to regurgitated pellets: molecular cormorant sexing combined with molecular fish identification and fish‐length regression analysis performed on prey hard parts. Altogether, 364 pellets delivered information on both, bird sex, and consumed prey. The sexes differed significantly in their overall prey composition, even though Perca fluviatilis, Rutilus rutilus, and Coregonus spp. represented the main food source for both. Albeit prey composition did not indicate the use of different water bodies by the sexes, male diet was characterized by higher prey diversity within a pellet and the consumption of larger fish. The current findings show that female and male cormorants to some extent target the available prey spectrum at different levels. Finally, the comprehensive and noninvasive approach has great potential for application in studies of other piscivorous bird species.     

Sexual differences in the diet of great cormorants <Emphasis Type="Italic">Phalacrocorax carbo sinensis</Emphasis> wintering in Greece

Sexual differences in the diet of the great cormorant, Phalacrocorax carbo sinensis, were studied in four Greek wintering areas, the Amvrakikos Gulf, the Axios and Evros Deltas and the Messolonghi Lagoon, through the analysis of stomach contents. Great cormorants are birds sexually dimorphic in size, with males being generally larger than females. Although similar prey species were found in the stomachs of both sexes in all the studied areas, significant differences were observed with respect to the proportion of species taken. Male birds ate higher proportions of large fish species such as grey mullets, European sea bass, Dicentrarchus labrax, and Prussian carp, Carassius gibelio, while female birds took higher proportions of smaller species such as big-scale sand smelt, Atherina boyeri, and black goby, Gobius niger. As a consequence, male great cormorants were found to feed on significantly larger prey than did females by means of fish standard length and body mass. There was no significant difference between the sexes in the mass of food found in stomachs.     

Does size dimorphism reduce competition between sexes? The diet of male and female pine martens at local and wider geographical scales

Sex-specific niche segregation is often used to explain sexual size dimorphism (SSD). However, whether food niche partitioning between sexes occurs as a case of sexual size dimorphism or by other mechanisms, such as behavioural dimorphism or habitat segregation, remains poorly understood. To evaluate the nature and extent of food-niche differentiation between sexes in a solitary predator I examined variation in the diet of male and female pine martensMartes martes Linnaeus, 1758 in years of high and low rodent abundance. Small mammals were the most important prey for pine martens in years of both low and high rodent abundance (occurring in more than 49% of scats). Birds, invertebrates and plant material were relatively common food items in summer diet, whereas ungulate carcasses were often consumed in autumn—winter. In general, males consumed more ungulate carcasses, plant material, amphibians and reptiles than did females, whereas females preyed more on squirrels and birds than males. There was significant seasonally dependent, between-sex variation in the occurrence of shrews, small rodents, other mammals, birds and invertebrates in marten diet. Whereas the occurrence of bank vole, birds, carcasses and plant material changed between sexes, seasons and years with various rodent abundances, both sexes consumed larger prey and had increased food niche breadth in years of low compared with high rodent abundance. Neither prey size nor food niche breadth were significantly different between males and females. The food-niche overlap between sexes was consistently lower in spring and in years of low rodent abundance. A wider geographical comparison of different marten populations showed that the diet of males and females varied significantly between locations. Females consistently preyed on squirrels and birds, whereas males fed more often on ungulate carcasses and plant material. Local and geographical comparison of male and female diets suggest that food-niche partitioning between male and female pine martens changes across different habitat and food conditions, and is not related to sexual size dimorphism, but rather to behavioural differences between sexes.     

Body size, prey handling efficiency and choice of diet in three coniferous forest tits (Paridae)

The main purpose of this study was to link morphological differences between great tit ( Parus major ), willow tit ( P. montanus ) and coal tit ( P. ater ) and their rate of energy acquisition and choice of diet in order to explore the potential for competitive relations between them more directly. Handling times were measured in the laboratory by presenting mealworms of different sizes to the birds. Great tits were more efficient in handling large prey than were the smaller-bodied willow- and coal tits; for small prey sizes the coal tit was the least efficient species. Using the ratio of prey mass to the handling time value, a utility function for each species was constructed. These results suggests a potential for a segregation of the species on the food axis. However, results from the prey choice experiment show that despite considerable differences in functional morphology between the three species they do not differ significantly in the range of prey size exploited. My results suggest that the alleged importance of prey size partitioning is not likely to play the major role for the coexistence of these coniferous forests tits.     

Sex-specific niche partitioning and sexual size dimorphism in Australian pythons (Morelia spilota imbricata)

Sexual dimorphism is usually interpreted in terms of reproductive adaptations, but the degree of sex divergence also may be affected by sex-based niche partitioning. In gape-limited animals like snakes, the degree of sexual dimorphism in body size (SSD) or relative head size can determine the size spectrum of ingestible prey for each sex. Our studies of one mainland and four insular Western Australian populations of carpet pythons ( Morelia spilota ) reveal remarkable geographical variation in SSD, associated with differences in prey resources available to the snakes. In all five populations, females grew larger than males and had larger heads relative to body length. However, the populations differed in mean body sizes and relative head sizes, as well as in the degree of sexual dimorphism in these traits. Adult males and females also diverged strongly in dietary composition: males consumed small prey (lizards, mice and small birds), while females took larger mammals such as possums and wallabies. Geographic differences in the availability of large mammalian prey were linked to differences in mean adult body sizes of females (the larger sex) and thus contributed to sex-based resource partitioning. For example, in one population adult male snakes ate mice and adult females ate wallabies; in another, birds and lizards were important prey types for both sexes. Thus, the high degree of geographical variation among python populations in sexually dimorphic aspects of body size and shape plausibly results from geographical variation in prey availability.  © 2002 The Linnean Society of London, Biological Journal of the Linnean Society , 2002, 77 , 113–125.     

Cormorant Catch Concerns for Fishers: Estimating the Size-Selectivity of a Piscivorous Bird

Conflict arises in fisheries worldwide when piscivorous birds target fish species of commercial value. This paper presents a method for estimating size selectivity functions for piscivores and uses it to compare predation selectivities of Great Cormorants (Phalacrocorax carbo sinensis L. 1758) with that of gill-net fishing on a European perch (Perca fluviatilis L. 1758) population in the Curonian Lagoon, Lithuania. Fishers often regard cormorants as an unwanted “satellite species”, but the degree of direct competition and overlap in size-specific selectivity between fishers and cormorants is unknown. This study showed negligible overlap in selectivity between Great Cormorants and legal-sized commercial nets. The selectivity estimation method has general application potential for use in conjunction with population dynamics models to assess fish population responses to size-selective fishing from a wide range of piscivorous predators.     

Do the evolutionary interactions between moths and bats promote niche partitioning between bats and birds?

Ecological theory suggests that the coexistence of species is promoted by the partitioning of available resources, as in dietary niche partitioning where predators partition prey. Yet, the mechanisms underlying dietary niche partitioning are not always clear. We used fecal DNA metabarcoding to investigate the diets of seven nocturnal insectivorous bird and bat species. Low diet overlap (2%–22%) supported resource partitioning among all species. Differences in diet corresponded with species identity, prey detection method, and foraging behavior of predators. Insects with ultrasonic hearing capabilities were consumed significantly more often by birds than bats, consistent with an evolved avoidance of echolocating strategies. In turn, bats consumed a greater proportion of noneared insects such as spruce budworms. Overall, our results suggest that evolutionary interactions among bats and moths translate to dietary niche partitioning and coexistence among bats and nocturnal birds.     

Wing shape variation of Finnish and Central European Willow Warblers Phylloscopus trochilus and Chiffchaffs P. collybita

Wing shape variation of European Willow Warblers Phylloscopus trochilus and Chiffchaffs P. collybita was studied using indices calculated from wing formulae. Our data were from free-living local populations of P. t. acredula and P. c. abietinus from southern Finland, and P. t. trochilus and P. c. collybita from southwestern Germany.
There were no significant shape differences between the subspecies of the Willow Warbler in which sexual dimorphism was pronounced. The subspecies of the Chiffchaff were significantly different while the sexes were not. There were also significant differences between adult and immature individuals in the autumn. The intrapopulation variation must be considered in attempts to recognize different subspecies or populations in data on birds caught during migration.
In the Willow Warbler at least, the difference between age-groups was larger in males than in females. If the change in wing formula due to the complete pre-nuptial moult is similar in both sexes, juvenile males intermediate between adult females and adult males suffer disproportionately high mortality. It was earlier suggested that sexual selection increases body size in Willow Warbler males, but the present results imply additional selection pressures for increasing sexual dimorphism.     

Maximizing dietary information retrievable from carcasses of Great Cormorants Phalacrocorax carbo using a combined morphological and molecular analytical approach

Avian carcasses can provide important information on the trophic ecology of birds. Usually, the number of carcasses available for examination is limited and therefore it is important to gain as much dietary information per specimen as possible. In piscivorous birds and raptors, the stomach has been the primary source of dietary information, whereas the gut (intestine) has so far been neglected as it usually contains only a few morphologically identifiable hard parts of prey. Molecular approaches have the potential to retrieve dietary information from the gut, although this has not yet been verified. As well as identifying the prey, it is important to estimate any secondary predation to avoid food web errors in dietary analyses. The assignment of accidentally consumed prey is notoriously difficult regardless of the prey identification approach used. In the present study, morphological and molecular analyses were, for the first time, combined to maximize the dietary information retrievable from the complete digestive tract of Great Cormorants Phalacrocorax carbo sinensis. Moreover, a novel approach based on predator–prey size ratios was applied to these piscivorous birds to minimize the number of samples that might contain secondarily predated prey. The stomach contents of the examined birds were found to provide the most dietary information when morphological and molecular analyses were used in combination. However, compared with the morphological approach, the molecular analysis increased the number of fish species detected by 39%. The molecular approach also permitted the identification of fish DNA in the Cormorant guts. Predator–prey size ratios derived from morphological analysis of fish hard parts can reduce the incidence of potential confounding influence of secondarily predated prey by 80%. Our findings demonstrate that a combination of morphological and molecular approaches maximizes the trophic information retrievable from bird carcasses.     

THE FOODS OF GREAT AND SOOTY SHEARWATERS PUFFINUS GRAVIS AND P. GRISEUS IN EASTERN CANADIAN WATERS

The foods of Great and Sooty Shearwaters Puffinus gravis and P. griseus are described from birds collected off eastern Canada. There was a broad overlap in diet, but Great Shearwaters tended to take more squid and tough-bodied fish such as mackerel Scomber scombrus while Sooties took more euphausiids Meganyctiphanes norvegica and soft-bodied fish such as Herring Clupea harengus . These differences are apparently related to differences in bill structure and in the degree of adaptation to underwater swimming.
The birds appeared to feed opportunistically on whatever prey was locally available in the size-range between euphausiids and small fish and squid: Meganyctiphanes off southwest Nova Scotia, spawning and post-spawning capelin Mallotus villosus off eastern Newfoundland, and migrating squid Illex illecebrosus on the Grand Bank. Possible factors influencing prey selection are discussed. It is suggested that the timing and routes of the birds' migrations in the North Atlantic are related to the exploitation of such local concentrations.
Despite the overlap in diets, differences in the distributions of the two species rule out the possibility of significant competition for common food resources.     

Functional and ecological relevance of intraspecific variation in body size and shape in the lizard Podarcis melisellensis (Lacertidae)

Within populations, individual animals may vary considerably in morphology and ecology. The degree to which variation in morphology is related to ecological variation within a population remains largely unexplored. We investigated whether variation in body size and shape among sexes and age classes of the lizard Podarcis melisellensis translates in differential whole-animal performance (sprint speed, bite force), escape and prey attack behaviour in the field, microhabitat use and diet. Male and female adult lizards differed significantly in body size and head and limb proportions. These morphological differences were reflected in differences in bite strength, but not in sprint speed. Accordingly, field measurements of escape behaviour and prey attack speed did not differ between the sexes, but males ate larger, harder and faster prey than females. In addition to differences in body size, juveniles diverged from adults in relative limb and head dimensions. These shape differences may explain the relatively high sprint and bite capacities of juvenile lizards. Ontogenetic variation in morphology and performance is strongly reflected in the behaviour and ecology in the field, with juveniles differing from adults in aspects of their microhabitat use, escape behaviour and diet.  © 2008 The Linnean Society of London, Biological Journal of the Linnean Society , 2008, 94 , 251–264.     

FOOD PREFERENCE AND THE EXTENT OF OVERLAP IN SOME INSECTIVOROUS BIRDS, WITH SPECIAL REFERENCE TO THE TYRANNIDAE

Analysis of published records of the food of flycatchers (Tyrannidae), swallows (Hirundinidae) and vireos (Vireonidae) of North America, and terns (Laridae) of the Pacific Ocean showed that size, rather than taxonomic, differences in food appear to be the most important ones for these birds. Although the distribution of insect sizes in nature approaches a two-parameter log-normal distribution, the distributions of the sizes of food items taken by birds show no significant differences from log-normal. Birds of a given feeding type (e.g. flycatchers) show a strong correlation of average prey size with bird body weight, and a significantly less strong correlation with bill characteristics, indicating that body size is a better predictor of prey size than any single bill character. The slopes and/or intercepts of regressions of food size against body weight are different for birds of different foraging type. Values of overlap in food preference are proportional to the similarity of the two species compared, expressed as the ratio of their weights. Because insect taxa differ in size and because the amount of overlap in taxonomic composition of foods is therefore very roughly proportional to the amount of overlap in size, taxonomic differences in food may merely reflect differences in size preferences. Values for overlap in food are greater than for most cases involving spatial niche parameters, indicating space is more easily divided than food, a conclusion supported by the relative rarity of large flycatchers.     

Evolution of reversed sexual size dimorphism and role partitioning among predatory birds, with a size scaling of flight performance

To explain the adaptive significance of sex role partitioning and reversed sexual size dimorphism among raptors, owls and skuas, where females are usually larger than males, we combine several previous hypotheses with some new ideas. Owing to their structural and behavioural adaptations for prey capture, predatory birds have better prospects than other birds of defending their offspring against nest predators. This makes sex role partitioning advantageous; one parent guards the offspring while the other forages for the family. Further, among predators hunting alert prey such as vertebrates, two mates because of interference may not procur much more food than would one mate hunting alone. By contrast, two mates feeding on less alert prey may together obtain almost twice as much food as one mate hunting alone. For these reasons, partitioning of breeding labours might be adaptive only in predatory birds. An initial imbalance favours female nest guarding and male foraging: the developing eggs might be damaged if the female attacks prey; their mass might reduce her flight performance; she must visit the nest to lay; and the male feeds her before she lays (‘courtship feeding’). Increased female body size should enhance egg production, incubation, ability to tear apart prey for the young, and, in particular, offspring protection in predatory birds. Efficient foraging during the breeding period then becomes most important for the male. This imposes great demands on aerial agility in males, particularly among predators of agile prey. Flight performance decreases with increasing size in five of six aspects explored. The male must therefore not be too large in relation to the most important prey. For these reasons, he should be smaller than the female. Among predatory birds, size dimorphism increases with the proportion of birds in the diet, which may be explained as follows. Adult birds have mainly one type of predators: other predatory birds. Because almost only these specialists exploit adult birds, they carry out most of the cropping of this prey. A predator of easier prey competes with many other kinds of predators, which considerably reduce prey abundance in its territory. This is not so for predators of adult birds. Further, because birds are extremely agile, the specialized predator can hunt efficiently only within a limited size range of birds, whose flight skill it can match. Increased size dimorphism among these predators therefore should be particularly important for enlarging the combined food base of the pair. A bird specialist may consume much of the available prey in the suitable size range during the breeding period. When the predator's young are large enough to defend themselves, the female aids better by hunting than by guarding the chicks. It is advantageous among bird specialists if she hunts prey of other sizes than does the male, who has by then reduced prey abundance in his prey size class. But among predatory birds hunting easier prey the female gains little by hunting outside the male's prey spectrum, because other kinds of predators will have reduced the prey abundance outside as well as inside the male's preferred size range. Intra-pair food separation through large sexual size dimorphism therefore should be particularly advantageous among predators of birds. This may be the main reason why the degree of size dimorphism increases with the dietary proportion of birds.     

Comparative gastrointestinal morphology of the Kori bustard and secretary bird

Two secretary birds and three Kori bustards were studied to determine differences between their body size and gastrointestinal morphology. Body measurements were made on captive, live birds and gastrointestinal measurements on fresh postmortem specimens. For predator species, such as the Kori bustard and secretary bird, body size is a function of their ability to capture and destroy prey. While the secretary bird was clearly the taller of the two species, superior body weight, wing length, and therefore body size was noted for the Kori bustard. The size and length of the gastrointestinal tract varied between species. The secretary bird had the shorter, less complex digestive tract, with a foregut well adapted for consumption of large quantities of flesh. The large intestine was devoid of ceca. The gastrointestinal tract of the Kori bustard was markedly different from that of the secretary bird. The foregut was less complex and the large intestine possessed large, voluminous ceca.     

大山雀身体大小的性二态及其季节变化

动物中普遍存在雌雄个体身体大小的性二态现象。了解近缘种之间身体大小性二态现象的差异，可为深入探讨身体大小性二态现象的潜在驱动机制提供证据。国外对欧亚大山雀（Parus major）的研究发现，其喙长、跗跖长、翅长等 6 项身体大小指标存在着明显的性二态，且喙长的性二态存在季节间差异。大山雀（P. cinereus）曾被作为欧亚大山雀的一个亚种，其形态和行为与欧亚大山雀存在着诸多相似之处。为探讨大山雀是否也存在身体大小性二态及季节性差异，本研究分析了 2018 至 2020 年间在河南董寨国家级自然保护区捕捉的 226 只（雌性 96 只和雄性 130 只）大山雀的喙长、头喙长、跗跖长、翅长、尾长和体长这 6 项体征指标的两性差异及其季节变化。结果显示，大山雀上述 6 项身体大小指标均存在不同程度的性二态现象，且雄性个体仅喙长与雌性的差异不显著，其余 5 项指标均显著大于雌性。此外，身体大小指标的两性差异不随季节显著变化，但两性的跗跖长在秋季均显著短于冬季和繁殖季，尾长在繁殖季均显著长于秋季和冬季。上述结果表明，大山雀身体大小的性二态及其季节性差异与欧亚大山雀并不完全相似。无论其身体大小存在性二态和季节变化的原因，还是其与欧亚大山雀在身体大小性二态模式上的差别，都有待今后进一步的研究。     

Feeding ecology of North American gopher snakes (Pituophis catenifer, Colubridae)

Studies of food relations are important to our understanding of ecology at the individual, population and community levels. Detailed documentation of the diet of large‐bodied, widespread snakes allows us to assess size‐dependent and geographical variation in feeding preferences of gape‐limited predators. Furthermore, with knowledge of the food habits of sympatric taxa we can explore possible causes of interspecific differences in trophic niches. The feeding ecology of the North American gopher snake, Pituophis catenifer, was studied based on the stomach contents of more than 2600 preserved and free‐ranging specimens, and published and unpublished dietary records. Of 1066 items, mammals (797, 74.8%), birds (86, 8.1%), bird eggs (127, 11.9%), and lizards (35, 3.3%) were the most frequently eaten prey. Gopher snakes fed upon subterranean, nocturnal and diurnal prey. The serpents are primarily diurnal, but can also be active at night. Therefore, gopher snakes captured their victims by actively searching underground tunnel systems, retreat places and perching sites during the day, or by pursuing them or seizing them while they rested at night. Gopher snakes of all sizes preyed on mammals, but only individuals larger than 40 and 42 cm in snout–vent length took bird eggs and birds, respectively, possibly due to gape constraints in smaller serpents. Specimens that ate lizards were smaller than those that consumed mammals or birds. Gopher snakes raided nests regularly, as evidenced by the high frequency of nestling mammals and birds and avian eggs eaten. Most (332) P. catenifer contained single prey, but 95 animals contained 2–35 items. Of the 321 items for which direction of ingestion was determined, 284 (88.5%) were swallowed head‐first, 35 (10.9%) were ingested tail‐first, and two (0.6%) were taken sideways. Heavier gopher snakes took heavier prey, but heavier serpents ingested prey with smaller mass relative to snake mass, evidence that the lower limit of prey mass did not increase with snake mass. Specimens from the California Province and Arid Deserts (i.e. Mojave, Sonoran and Chihuahuan Deserts) took the largest proportion of lizards, whereas individuals from the Great Basin Desert consumed a higher percentage of mammals than serpents from other areas, and P. catenifer from the Great Plains ate a greater proportion of bird eggs. Differences in prey availability among biogeographical regions and unusual circumstances of particular gopher snake populations may account for these patterns. Gopher snakes have proportionally longer heads than broadly sympatric Rhinocheilus lecontei (long‐nosed snake), Charina bottae (rubber boa) and Lampropeltis zonata (California mountain kingsnake), which perhaps explains why, contrary to the case in P. catenifer, the smaller size classes of those three species do not eat mammals. © 2002 The Linnean Society of London, Biological Journal of the Linnean Society, 2002, 77 , 165–183.     

Home-ranges and patterns of spatial use in territorial Bonelli's Eagles Aquila fasciata

Bonelli's Eagle Aquila fasciata is one of the rarest birds of prey in Europe, where it has suffered a significant decline in recent decades. We present information on the home-ranges and spatial parameters of 18 Bonelli's Eagles radiotracked in 2002–2006 in Catalonia (northeast Spain) and describe the home-range probability kernel, distances moved, breeding area eccentricity, territorial overlap, nearest neighbour distance and breeding site fidelity, and assess the influence of sex, breeding status, season and geographical area on these parameters. Median home-range according to the minimum convex polygon (MCP) and 95% kernel were 50.3 and 36.1 km², respectively. The median breeding area eccentricity was 1477 m. There was considerable overlap in the home-range of both sexes within pairs (MCP: 71.4% and 95% kernel: 98.5%), indicating close pair bonding and similar foraging patterns. Overlap in home-ranges of up to 15% between neighbouring individuals also occurred and was positively related to breeding pair density. There was no difference in spatial parameters between sexes or with breeding status, but during the non-breeding season Eagles had larger home-ranges and stayed further from nests. The high consistency across birds suggests a pattern of spatial use that is characteristic of this species. The high level of use of breeding areas and their surroundings (50% kernel) throughout the year makes it important that these areas be protected from human disturbance. Additionally, it is necessary that heavily used areas away from nesting sites, which are used for foraging and roosting, are identified, protected and managed in a sustainable fashion.     

Species delimitation in the Hwamei Garrulax canorus

Due to the male's elaborate songs, the Hwamei Garrulax canorus is the most popular caged bird in the global Chinese community. Three allopatric Hwamei subspecies have been described: G. c. canorus in central and southern China and northern Indochina, G. c. owstoni from Hainan and G. c. taewanus from Taiwan. We sequenced the entire mitochondrial cytochrome b gene to reconstruct the molecular intraspecific phylogeny of the Hwamei. Molecular phylogenetic trees indicated that individuals of the three subspecies formed three monophyletic clades with high bootstrap support (> 95%). The basal clade was G. c. taewanus . According to a conventional molecular clock (2% divergence per million years), G. c. taewanus split from the other Hwamei taxa around 1.5 million years ago, and G. c. owstoni diverged from G. c. canorus around 0.6 million years ago. Considering the periodic connection between the Asian mainland and nearby continental islands during the glacial periods, habitat vicariance may have played a more important role than geographical vicariance in facilitating the differentiation of these taxa. Molecular diagnosability, population integrity, and concordance between the population ranges and the topology of the phylogenetic tree suggested that the Hwamei should be delimited into at least two full species: G. canorus and G. taewanus . Our work represents one of the first attempts to re-evaluate the intraspecific systematics for an eastern Asian bird species using molecular data.     

Spatial congruence between ecotones and range-restricted species: implications for conservation biogeography at the sub-continental scale

Aim To examine whether at a sub‐continental scale range‐limited species tend to occur close to areas of transition between vegetation boundaries more often than expected by chance. Location South Africa and Lesotho. Methods We examined the relationship between the distance of a grid square to ecological transition areas between vegetation types and both avian and frog range‐limited species richness in the quadrat. We used quadrats at a spatial resolution of quarter degree (15′ × 15′≈ 676 km²). Spatial congruence between areas representing range‐restricted species and those representing ecological transition zones was assessed using a random draw technique. Results Species richness and range size rarity are generally negatively correlated with distance to transition areas between vegetation communities when analysed for the whole region for both groups. Although this relationship becomes weaker after controlling for environmental energy and topographical heterogeneity, the explanatory power of distance to transition areas remains significant, and compared to the different biomes examined, accounts for most of the variation in bird richness (20%), frog richness (18%), range‐restricted bird species (17%) and range‐restricted frog species (16%) in the savanna biome. The random draw technique indicated that areas representing range‐restricted species were situated significantly closer in space to those areas representing transition areas between vegetation communities than expected by chance. Main conclusions We find that at the sub‐continental scale, when examined for South Africa, areas of transition between vegetation communities hold concentrations of range‐limited species in both birds and frogs. We find that South African endemic/range‐limited birds and frogs are located closer to ecological transition zones than endemics and non‐endemics combined. This has important implications for ongoing conservation planning in a biogeographical context.     

Genetic and morphological variation in Dunlin Calidris alpina breeding in the Palearctic tundra

The extensive overlap in morphological characters between populations of Dunlin Calidris alpina imposes problems of determining the origin of migrating and wintering birds. The morphology of the birds also varies between the sexes, and the sex of a Dunlin may often be difficult to determine. To clarify if mitochondrial DNA can be used to identify which breeding areas migrating Dunlin come from, we investigated the occurrence of different mtDNA haplotypes in Dunlin from eight breeding areas on the Russian and Siberian tundra. Four haplotypes were found and at most sites more than one haplotype occurred. The European haplotype predominated in the area west of the Taymyr Peninsula, the Siberian haplotype in central Siberia (from the Taymyr Peninsula to the Lopatka Peninsula) and the Beringian haplotype in eastern Siberia. One individual of an Alaskan haplotype, not detected previously among breeding birds outside North America, was found on Wrangel Island. The sex of each bird was identified genetically and the morphology of males and females was analysed separately. Birds with the European haplotype were generally smaller than birds with the Beringian or Alaskan haplotypes. Birds possessing the Siberian haplotype showed intermediate values in most cases. After compensating for differences between sites, males with the Siberian haplotype had significantly longer bills than males having the European haplotype. Multiple regressions indicate that mitochondrial DNA analysis improves models estimating the breeding origin of migrating Dunlin.