The phylogenetic position of the Galápagos Cormorant

The endangered Galápagos Cormorant, Phalacrocorax harrisi, is unique among the species of the Phalacrocoracidae in being flightless and sequentially polyandrous. It has had a vexed taxonomic history, variously being lumped with all the species in Phalacrocorax, being accorded its own genus, Nannopterum, or being included in Leucocarbo or Compsohalieus. Different authorities have similarly suggested a number of different species as being its closest relative. Here we use novel mitochondrial DNA sequence data to show that the Galápagos Cormorant is related to the sister pair of the mainland Americas, the Double-crested Cormorant, P. auritus, and the Neotropic Cormorant, P. brasilianus. This trio of species has high statistical support (Bayesian posterior probability of 1.00; NJ bootstrap 98%; MP bootstrap 91%). The Galápagos Cormorant is thus a relatively recent offshoot of the mainland form, which has subsequently evolved flightlessness. Until the phylogeny of the cormorants is more clearly resolved, we recommend the continued use of Phalacrocorax for all species.     

Extent of phenotypic flexibility during long‐distance flight is determined by tissue‐specific turnover rates: a new hypothesis

Phenotypic flexibility in organ size of migratory birds is typically explained in functional terms in accordance with the principal of economic design. However, proposed functional hypotheses do not adequately explain differences in phenotypic flexibility between organs during fasting and in‐flight starvation. We show that the extent of phenotypic flexibility in organ mass in five species of migratory birds during actual migration or simulated in‐flight starvation consistently ranked as follows from highest to lowest mass change: small intestine, liver, kidney, gizzard, heart, flight and leg muscle. This pattern of phenotypic flexibility in organ mass was not consistent with proposed functional hypotheses, and was almost completely explained by differences in tissue‐specific turnover rate measured in vivo using nutrients differing in their isotopic values. Thus, the fundamental process of tissue‐specific protein turnover determines extent of organ mass changes for birds during migration, this likely applies to other organisms during fasting, and no further functional explanation(s) for differences in the magnitude of phenotypic flexibility between organs is required.     

Comparing short and long‐distance dispersal: modelling and field case studies

Dispersal is a factor of great importance in determining a species spatial distribution. Short distance dispersal (SDD) and long distance dispersal (LDD) strategies yield very different spatial distributions. In this paper we compare spatial spread patterns from SDD and LDD simulations, contrast them with patterns from field data, and assess the significance of biological and population traits. Simulated SDD spread using an exponential function generates a single circular patch with a well‐defined invasion front showing a travelling‐wave structure. The invasive spread is relatively slow as it is restricted to reproductive individuals occupying the outer zone of the circular patch. As a consequence of this dispersal dynamics, spread is slower than spread generated by LDD. In contrast, the early and fast invasion of the entire habitat mediated by power law LDD not only involves a significantly greater invasion velocity, but also an entirely different habitat occupation. As newly dispersed individuals soon reach very distant portions of the habitat as well as the vicinity of the original dispersal focus, new growing patches are generated while the main patch increases its own growth absorbing the closest patches. As a consequence of both dispersal and lower density dependence, growth of the occupied area is much faster than with SDD. SDD and LDD also differ regarding pattern generation. With SDD, fractal patterns appear only in the border of the invasion front in SDD when competitive interaction with residents is included. In contrast, LDD patterns show fractality both in the spatial arrangements of patches as well as in patch borders. Moreover, values of border fractal dimension inform on the dispersal process in relation with habitat heterogeneity. The distribution of patch size is also scale‐free, showing two power laws characteristic of small and large patch sizes directly arising from the dispersal and reproductive dynamics. Ecological factors like habitat heterogeneity are relevant for dispersal, although its importance is greater for SDD, lowering the invasion velocity. Among the life history traits considered, adult mortality, the juvenile bank and mean dispersal distance are the most relevant for SDD. For LDD, habitat heterogeneity and changes in life history traits are not so relevant, causing minor changes in the values of the scale‐free parameters. Our work on short and long distance dispersal shows novel theoretical differences between SDD and LDD in invasive systems (mechanisms of pattern formation, fractal and scaling properties, relevance of different life history traits and habitat variables) that correspond closely with field examples and were not analyzed, at least in this degree of detail, by the previously existing models.     

The role of migratory ducks in the long‐distance dispersal of native plants and the spread of exotic plants in Europe

Little is known about the role of migratory waterfowl in the long‐distance dispersal (LDD) of seeds. We studied the gut contents of 42 teals Anas crecca collected in the Camargue, southern France, and found intact seeds of 16 species. There was no relationship between the probability that a given seed species was found intact in the lower gut, and the seed hardness or size. The number of seeds found in the oesophagus and gizzard (a measure of ingestion rate) was the only significant predictor of the occurrence of intact seeds in the lower gut, so studies of waterfowl diet can be used as surrogates of dispersal potential. In a literature review, we identified 223 seed species recorded in 25 diet studies of teal, pintail Anas acuta, wigeon A. penelope or mallard A. platyrhynchos in Europe. We considered whether limited species distribution reduces the chances that a seed can be carried to suitable habitat following LDD. Overall, 72% of plant species recorded in duck diets in southern Europe (36 of 50) were also recorded in the north, whereas 97% of species recorded in duck diets in the north (137 of 141) were also recorded in the south. This suggests a great potential for LDD, since most dispersed plants species occur throughout the migratory range of ducks. Migratory ducks are important vectors for both terrestrial and aquatic plant species, even those lacking the fleshy fruits or hooks typically used to identify seeds dispersed by birds. Finally, we show ducks are important vectors of exotic plant species. We identified 14 alien to Europe and 44 native to Europe but introduced to some European countries whose seeds have been recorded in duck diet.     

Inbreeding,immune defence and ectoparasite load in different mockingbird populations and species in the Galápagos Islands

Inbreeding may impair an individual's immune system, render it more susceptible to disease and hence contribute to the extinction risk of small and isolated populations, as often found on islands. So far, surprisingly few studies have assessed the effects of inbreeding on immunocompetence in wild populations. Using 26 microsatellite loci and genetic data from museum specimens and contemporary samples, we calculated short‐term and long‐term inbreeding in 13 different mockingbird populations covering the range of all 4 species in the Galápagos Islands and compared them with three different measures of innate immunity and ectoparasite load. We found no significant effect of either measure of inbreeding on natural antibody or complement enzyme titres, heterophil‐lymphocyte ratio or feather louse abundance. Hence, our results do not support a link between inbreeding and immunocompetence. However, overall statistical power and repeatabilities of antibody and complement enzyme titres were low. Nevertheless, generally, natural antibody titres were high suggesting that the mockingbirds may be equipped with a strong first line of defence, as found in other island species.     

Nematodes surfing the waves: long‐distance dispersal of soil‐borne microfauna via sea swept rhizomes

Dispersal mechanisms of soil‐borne microfauna have hitherto received little attention. Understanding dispersal mechanisms of these species is important to unravel their basic life history traits, biogeography, exchange of individuals between populations, and local adaptation. Soil‐borne nematodes and root‐feeding nematodes in particular occupy a key position in soil‐food webs and can be determinants for plant growth and vegetation structure and succession. However, their dispersal abilities have been scarcely addressed, predominantly focusing on species of agricultural importance. Still, root‐feeding nematodes are usually considered as being extremely limited and bound to the rhizosphere of plants. We investigated a mechanism for long distance dispersal of root‐feeding nematodes associated to two widespread coastal dune grasses. The nematodes are known to be crucial for the functioning of these grasses. We experimentally tested the hypothesis that root‐feeding nematodes are able to move across long distances inside rhizome fragments that are dispersed by seawater. We also tested the survival capacities of the host plants in seawater. Our study demonstrates that root‐feeding nematodes and plants are able to survive immersion in seawater, providing a mechanism for long distance dispersal of root feeding nematodes together with their host plant. Drifting rhizome fragments enable the exchange of plant material and animals between dune systems. These results provide new insights to understand the ecology of dune vegetation, the interaction with soil‐borne organisms and more importantly, re‐set the scale of spatial dynamics of a group of organisms considered extremely constrained in its dispersal capacities.     

A set of highly discriminating microsatellite loci for the Galápagos marine iguana Amblyrhynchus cristatus

We describe here the cloning of 12 (7 dinucleotide, 1 trinucleotide and 4 tetranucleotide) microsatellite loci for the Galápagos marine iguana Amblyrhynchus cristatus. When tested for individuals from five different island populations on the Galápagos archipelago, high genetic diversities (9–20 alleles per locus) and heterozygosities (0.200–0.944) were observed. All loci showed no obvious deviations from Hardy–Weinberg equilibrium. The new set of microsatellite loci was able to assign individuals reliably to their island of origin, thus being able to discriminate between residents and migrants between islands.     

Genetic evidence for high propagule pressure and long‐distance dispersal in monk parakeet (Myiopsitta monachus) invasive populations

The monk parakeet (Myiopsitta monachus) is a successful invasive species that does not exhibit life history traits typically associated with colonizing species (e.g., high reproductive rate or long‐distance dispersal capacity). To investigate this apparent paradox, we examined individual and population genetic patterns of microsatellite loci at one native and two invasive sites. More specifically, we aimed at evaluating the role of propagule pressure, sexual monogamy and long‐distance dispersal in monk parakeet invasion success. Our results indicate little loss of genetic variation at invasive sites relative to the native site. We also found strong evidence for sexual monogamy from patterns of relatedness within sites, and no definite cases of extra‐pair paternity in either the native site sample or the examined invasive site. Taken together, these patterns directly and indirectly suggest that high propagule pressure has contributed to monk parakeet invasion success. In addition, we found evidence for frequent long‐distance dispersal at an invasive site (～100 km) that sharply contrasted with previous estimates of smaller dispersal distance made in the native range (～2 km), suggesting long‐range dispersal also contributes to the species’ spread within the United States. Overall, these results add to a growing body of literature pointing to the important role of propagule pressure in determining, and thus predicting, invasion success, especially for species whose life history traits are not typically associated with invasiveness.     

Evolution of the distribution of dispersal distance under distance‐dependent cost of dispersal

Abstract We analyse the evolution of the distribution of dispersal distances in a stable and homogeneous environment in one‐ and two‐dimensional habitats. In this model, dispersal evolves to avoid the competition between relatives although some cost might be associated with this behaviour. The evolutionarily stable dispersal distribution is characterized by an equilibration of the fitness gains among all the different dispersal distances. This cost‐benefit argument has heuristic value and facilitates the comprehension of results obtained numerically. In particular, it explains why some minimal or maximal probability of dispersal may evolve at intermediate distances when the cost of dispersal function is an increasing function of distance. We also show that kin selection may favour long range dispersal even if the survival cost of dispersal is very high, provided the survival probability does not vanish at long distances.     

Large‐scale geographic variation in iridescent structural ornaments of a long‐distance migratory bird

Iridescent colours produced during moult likely play an important role in pair formation in birds. We sought to quantify geographic variation in such colouration in a duck species, Eurasian teal Anas crecca, in winter (when mating occurs) to evaluate whether this variation reflects birds’ breeding origins or differential individual migration strategies in both males and females. We combined information on feather production region and individual attributes (body size, sex and age) of Eurasian teal from 82 wintering sites in France. Feather production region (moult site or natal origin) was inferred using feather deuterium values (δD_f). We performed spectral measurements to evaluate speculum colour and brightness contrasts for 1052 teal collected over four years. Colouration differed strongly among wintering regions, with birds wintering in eastern France exhibiting higher colour contrast than those wintering in the west. Body size and colouration were positively related. There were no differences in cohort‐specific δD_f values between separate wintering regions in France, indicating that within a winter quarter teal originated from areas across the entire breeding range. Overall, patterns of spatial variation in feather colouration were related most closely to body size which was consistent with predictions of a differential migration hypothesis, with larger and more colour‐contrasting birds wintering closer to their breeding grounds. Because moult speed is also known to affect colour production, early breeders or individuals that skipped reproduction may have invested more or earlier in their feather quality to gain potential advantages in monopolizing future mates.     

Effects of long‐distance running on serum opsonic activity measured by chemiluminescence

Exhaustive exercise such as long‐distance running has been shown to increase susceptibility to infection. In order to investigate whether serum opsonic activity plays a role in such conditions, we utilized luminol‐dependent and lucigenin‐dependent chemiluminescence (LmCL and LgCL). We took serum samples from 24 male marathon runners before and after running 30 km. Neutrophils were isolated from the peripheral blood of healthy volunteers. Serum opsonic activity was examined by measuring neutrophil ROS stimulated with zymosan particles opsonized by the serum samples. Immunoglobulin and complement levels in the serum were also measured. After a 30 km run, the maximum light emission was increased and the time to reach the maximum light emission was shortened significantly (p < 0.05) in LmCL. However, there were no significant changes in the immunoglobulin and complement levels. The increase of ROS production may suggest that serum opsonic activity is accelerated after running 30 km. Thus, serum opsonic activity might not play a significant role in the susceptibility to infection after long‐distance running. Copyright © 2003 John Wiley & Sons, Ltd.     

Male reproductive success and its behavioural correlates in a polygynous mammal,the Galápagos sea lion (Zalophus wollebaeki)

Sexual selection theory predicts competitive males and choosy females. Nevertheless, since molecular marker‐based studies, paternity outside the expected mating patterns has increasingly been described. Even in highly polygynous systems, where paternity is expected to be strongly skewed towards large, dominant males, alternative mating tactics have been suggested. We examined reproductive success in the polygynous Galápagos sea lion (Zalophus wollebaeki). Semiaquatic territoriality allows females to move freely and may lower the degree of polygyny otherwise suggested by both territorial behaviour and strong sexual dimorphism. We assigned paternities with 22 microsatellites and analysed how male reproductive success was related to size, dominance status, intra‐sexual agonistic behaviour, proximity to females, and attendance in the colony. Male behaviour was consistent across two seasons for all parameters under consideration. Attendance was by far the most important determinant of paternal success. Skew in reproductive success towards large, dominant males was weak and dominance status played no role. This appears to be caused by an extremely long reproductive season lasting five or more months, making it difficult for any male to monopolize receptive females. Females seem to choose displaying males that were present in the colony for a long time rather than dominance per se. Sexual dimorphism in Galápagos sea lions may thus be more influenced by selection for fasting than fighting ability. Our data provide further evidence for alternative mating tactics, as several males gained relatively high reproductive success despite short attendance and hardly any involvement in agonistic interactions.     

Multi‐generational long‐distance migration of insects: studying the painted lady butterfly in the Western Palaearctic

Long‐range, seasonal migration is a widespread phenomenon among insects, allowing them to track and exploit abundant but ephemeral resources over vast geographical areas. However, the basic patterns of how species shift across multiple locations and seasons are unknown in most cases, even though migrant species comprise an important component of the temperate‐zone biota. The painted lady butterfly Vanessa cardui is such an example; a cosmopolitan continuously‐brooded species which migrates each year between Africa and Europe, sometimes in enormous numbers. The migration of 2009 was one of the most impressive recorded, and thousands of observations were collected through citizen science programmes and systematic entomological surveys, such as high altitude insect‐monitoring radar and ground‐based butterfly monitoring schemes. Here we use V. cardui as a model species to better understand insect migration in the Western Palaearctic, and we capitalise on the complementary data sources available for this iconic butterfly. The migratory cycle in this species involves six generations, encompassing a latitudinal shift of thousands of kilometres (up to 60 degrees of latitude). The cycle comprises an annual poleward advance of the populations in spring followed by an equatorward return movement in autumn, with returning individuals potentially flying thousands of kilometres. We show that many long‐distance migrants take advantage of favourable winds, moving downwind at high elevation (from some tens of metres from the ground to altitudes over 1000 m), pointing at strong similarities in the flight strategies used by V. cardui and other migrant Lepidoptera. Our results reveal the highly successful strategy that has evolved in these insects, and provide a useful framework for a better understanding of long‐distance seasonal migration in the temperate regions worldwide.     

Structure of a plant-pollinator network on a pahoehoe lava desert of the Galápagos Islands

Plant-pollinator interactions are important for the evolution and survival of the species involved. Plant-pollinator networks on oceanic islands are often small in size and as a consequence the connectance is high suggesting a substantial generalisation level. Further, linkage level for insular plants is shown to be lower than on mainland. The present study investigates a plant-pollinator network on the Galápagos Islands that is unique because of its very small size. We recorded pollinator visits to plant species as well as pollen grains on insect bodies. The combination of these data increased the observed number of interactions. The values for connectance and linkage level for plants were found to be consistent with similar values found in other network studies. There were no relation between the abundance of plant species and the number of pollinating species. The dominating pollinator species was the Galápagos carpenter bee Xylocopa darwini . Specimens of the shorthorned grasshopper Halmenus cuspidatus turned out to carry pollen from five plant species out of twelve and are probably functioning as pollinator. Bagging experiments revealed dependency on insect visits for a high seed set for most of the plant species, but only one species Plumbago scandens seemed to possess a pollen limited seed set. The network showed an asymmetric pattern of number of interactions per species with a few species having several interactions and many species a few. This pattern is supposed to result in a rather robust community, but is also fairly sensitive if the dominant species are threatened. The high connectance value found could, however, counteract this vulnerability.