Juan S. Monrós

Feather mites and birds: an interaction mediated by uropygial gland size?

Feather mites (Arachnida: Acari: Astigmata) feed mainly on secretions of the uropygial gland of birds. Here, we use analyses corrected for phylogeny and body size to show that there is a positive correlation between the size of this gland and mite abundance in passerine birds at an interspecific level during the breeding season, suggesting that the gland mediates interactions between mites and birds. As predicted on the basis of hypothesized waterproofing and antibiotic functions of uropygial gland secretions, riparian/marsh bird species had larger glands and higher mite loads than birds living in less mesic terrestrial environments. An unexpected pattern was a steeper relationship between mite load and gland size in migratory birds than in residents. If moderate mite loads are beneficial to a host but high loads detrimental, this could create complex selection regimes in which gland size influences mite load and vice versa. Mites may exert selective pressures on gland size of their hosts that has resulted in smaller glands among migratory bird species, suggesting that smaller glands may have evolved in these birds to attenuate a possible detrimental effect of feather mites when present in large numbers.

Patterns of nestling provisioning by a single-prey loader bird, Great Tit Parus major

Capsule Nestling provisioning rates depend on nestling age and number, and on time of season, but not on time of day. Aims To determine patterns of nestling provisioning, the effort made by the parents, and the factors which affect them. Methods Mechanical counters to determine food provisioning patterns in 229 Great Tit Parus major nests over 4 years. Results Feeding frequency per chick showed a linear increase with nestling age and total feeding frequency stabilized towards the end of the nestling period. The number of visits per nest increased linearly, while those per nestling decreased linearly with brood‐size. Feeding rates per nest declined throughout the breeding season parallel to the seasonal brood‐size decline; feeding frequencies per nestling did not vary seasonally. Adult effort was almost constant throughout the day, independently of variations in brood‐size, ambient temperature and date. Conclusion Previous studies of nestling provisioning patterns were either contradictory or poorly supported by low sample sizes. The present study involved a much larger sample and clarifies nestling provisioning patterns and the factors with which they are associated.

Laying date and clutch size of Great Tits(Parus major) in the Mediterranean region: a comparison of four habitat types

Laying data and clutch size of Great Tits were studied in four different habitats in eastern Spain: two holm oak(Quercus ilex) forests, at 500 and 900–950 m a.s.l., a zeen oak(Quercus faginea) forest, at 900–1100 m a.s.l., a pine(Pinus sylvestris) forest, at 1000–1050 m a.s.l., and orange(Citrus aurantium) plantations, at 30 m a.s.l. All sites were placed at about the same latitude (39–41°N), and all were studied during the same years (1992–95). Our results show that (1) laying date did not differ between the natural habitats at the same altitude (range of the means of yearly means 4–8 May); (2) within the same habitat type (holm oak forest) laying date was earlier at low altitude (30 Aprilvs. 8 May); (3) laying date was earlier in the orange plantations (21 April) than in natural habitats; (4) among natural habitats at the same altitude, clutch size decreased from zeen oak (mean of yearly means 7.3 eggs) to holm oak (7.0 eggs) to pine forests (6.4 eggs), though only the difference between zeen oak and pine forests was significant; (5) within the same habitat type (holm oak forest), the clutch size tended to be larger at high altitude (7.0vs. 5.9 eggs); and (6) clutch size in orange plantations (7.7 eggs) did not differ significantly from that of the zeen oak forest, but was larger than in the holm oak and pine forests. We discuss the effect of the habitat type on laying date and clutch size of Great Tits. Legedatum und Gelegegröße der Kohlmeise wurden in vier unterschiedlichen Biotopen in Ostspanien untersucht: zwei Steineichenwälder(Quercus ilex) in 500 m und 900–950 mNN, ein Bergeichenwald(Quercus faginea) in 900–1100 mNN, ein Kiefernwald(Pinus sylvestris) in 1000–1050 mNN und eine Orangenpflanzung(Citrus aurantium) in 30 mNN. Alle fünf Gebiete lagen auf etwa demselben Breitengrad (39–41°N) und wurden 1992–1995 parallel untersucht. (1) Auf gleicher Meereshöhe unterscheidet sich der Legebeginn nicht zwischen den verschiedenen Waldbiotopen (im Mittel 4.–8. Mai). (2) Innerhalb desselben Biotoptyps (Steineichenwald) war der Legebeginn auf niedrigerer Meereshöhe früher als in höheren Lagen (30. April vs. 8. Mai). (3) Im Orangenhain wurde früher mit der Eiablage begonnen (21. April) als in den Waldbiotopen. (4) Auf gleicher Meereshöhe nahm die Gelegegröße vom Bergeichenwald (Mittelwert 7.3 Eier) über die Steineichenwälder (7,0 Eier) zum Kiefernwald hin ab (6.4 Eier), jedoch ist nur der Unterschied zwischen Bergeichenwald und Kiefernwald signifikant. (5) Innerhalb der Steineichenwälder besteht die Tendenz zu größerer Gelegegröße in den höheren Lagen (7.0 vs. 5.9 Eier). (6) Im Orangenhain war die Gelegegröße mit durchschnittlich 7.7 Eier ähnlich der im Bergeichenwald, aber größer als in den Steineichen- und Kiefernwälder.

Endangered subspecies of the Reed Bunting (Emberiza schoeniclus witherbyi and E. s. lusitanica) in Iberian Peninsula have different genetic structures

In the Iberian Peninsula, populations of two subspecies of the Reed Bunting Emberiza schoeniclus have become increasingly fragmented during the last decades when suitable habitats have been lost and/or the populations have gone extinct. Presently, both subspecies are endangered. We estimated the amount of genetic variation and population structure in order to define conservation units and management practices for these populations. We found that the subspecies lusitanica has clearly reduced genetic variation in nuclear and mitochondrial markers, has a drastically small effective population size and no genetic differentiation between populations. In contrast, the subspecies witherbyi is significantly structured, but the populations still hold large amounts of variation even though the effective population sizes are smaller than in the non-endangered subspecies schoeniclus. We suggest several management units for the Iberian populations. One unit includes subspecies lusitanica as a whole; the other three units are based on genetically differentiated populations of witherbyi. The most important genetic conservation measure in the case of lusitanica is to preserve the remaining habitats in order to at least maintain the present levels of gene flow. In the case of the three management units within witherbyi, the most urgent conservation measure is to improve the habitat quality to increase the population sizes.ZusammenfassungAuf der Iberischen Halbinsel wurden die Populationen von zwei Rohrammer-Unterarten Emberiza schoeniclus in den letzten Jahrzehnten, als geeignete Habitate verloren gingen und/oder Populationen ausstarben, zunehmend fragmentiert. Derzeitig sind beide Unterarten gefährdet. Wir schätzten die Höhe der genetischen Variation und die Populationsstruktur um daraus Einheiten für den Schutz und Managementpraktiken für diese Populationen zu bestimmen. Wir fanden heraus, dass die Unterart lusitanica eine deutlich reduzierte genetische Variation in nukleären wie mitochondrialen Markern, eine dramatisch geringe effektive Populationsgröße und keine genetische Differenzierung zwischen Populationen aufweist. Im Gegensatz dazu ist die Unterart witherbyi deutlich strukturiert, aber die einzelnen Populationen beinhalten immer noch große Anteile der Gesamtvariation, wenngleich die effektiven Populationsgrößen kleiner sind als die der nicht gefährdeten Unterart schoeniclus. Wir schlagen mehrere Management-Einheiten für die Iberischen Populationen vor. Eine Einheit beinhaltet die Unterart lusitanica als Ganzes; die anderen drei Einheiten basieren auf den genetisch differenzierten Populationen der Unterart witherbyi. Die allerwichtigste Schutzmaßnahme aus genetischer Sicht im Falle der Unterart lusitanica ist die Bewahrung der verbleibenden Habitate um zumindest das derzeitige Maß an Genfluss zu erhalten. Im Falle der drei Management-Einheiten der Unterart witherbyi ist die dringlichste Schutzmaßnahme die Verbesserung der Habitate um die Populationsgrößen zu erhöhen.

Phenotypic Divergence among West European Populations of Reed Bunting Emberiza schoeniclus : The Effects of Migratory and Foraging Behaviours

Divergent selection and local adaptation are responsible for many phenotypic differences between populations, potentially leading to speciation through the evolution of reproductive barriers. Here we evaluated the morphometric divergence among west European populations of Reed Bunting in order to determine the extent of local adaptation relative to two important selection pressures often associated with speciation in birds: migration and diet. We show that, as expected by theory, migratory E. s. schoeniclus had longer and more pointed wings and a slightly smaller body mass than the resident subspecies, with the exception of E. s. lusitanica, which despite having rounder wings was the smallest of all subspecies. Tail length, however, did not vary according to the expectation (shorter tails in migrants) probably because it is strongly correlated with wing length and might take longer to evolve. E. s. witherbyi, which feed on insects hiding inside reed stems during the winter, had a very thick, stubby bill. In contrast, northern populations, which feed on seeds, had thinner bills. Despite being much smaller, the southern E. s. lusitanica had a significantly thicker, longer bill than migratory E. s. schoeniclus, whereas birds from the UK population had significantly shorter, thinner bills. Geometric morphometric analyses revealed that the southern subspecies have a more convex culmen than E. s. schoeniclus, and E. s. lusitanica differs from the nominate subspecies in bill shape to a greater extent than in linear bill measurements, especially in males. Birds with a more convex culmen are thought to exert a greater strength at the bill tip, which is in agreement with their feeding technique. Overall, the three subspecies occurring in Western Europe differ in a variety of traits following the patterns predicted from their migratory and foraging behaviours, strongly suggesting that these birds have became locally adapted through natural selection.

Phylogeography of a Habitat Specialist with High Dispersal Capability: The Savi’s Warbler Locustella luscinioides

In order to describe the influence of Pleistocene glaciations on the genetic structure and demography of a highly mobile, but specialized, passerine, the Savi’s Warbler (Locustella luscinioides), mitochondrial DNA sequences (ND2) and microsatellites were analysed in c.330 individuals of 17 breeding and two wintering populations. Phylogenetic, population genetics and coalescent methods were used to describe the genetic structure, determine the timing of the major splits and model the demography of populations. Savi’s Warblers split from its sister species c.8 million years ago and have two major haplotype groups that diverged in the early/middle Pleistocene. One of these clades originated in the Balkans and is currently widespread, showing strong evidence for population expansion; whereas the other is restricted to Iberia and remained stable. Microsatellites agreed with a genetic break around the Pyrenees, but showed considerable introgression and a weaker genetic structure. Both genetic markers showed an isolation-by-distance pattern associated with the population expansion of the eastern clade. Breeding populations seem to be segregated at the wintering sites, but results on migratory connectivity are preliminary. Savi’s Warbler is the only known migratory bird species in which Iberian birds did not expand beyond the Pyrenees after the last glaciation. Despite the long period of independent evolution of western and eastern populations, complete introgression occurred when these groups met in Iberia. Mitochondrial sequences indicated the existence of refugia-within-refugia in the Iberian Peninsula during the last glacial period, which is surprising given the high dispersal capacity of this species. Plumage differences of eastern subspecies seemed to have evolved recently through natural selection, in agreement with the glacial expansion hypothesis. This study supports the great importance of the Iberian Peninsula and its role for the conservation of genetic variation.

Invasion biology in non-free-living species: interactions between abiotic (climatic) and biotic (host availability) factors in geographical space in crayfish commensals (Ostracoda, Entocytheridae)

In invasion processes, both abiotic and biotic factors are considered essential, but the latter are usually disregarded when modeling the potential spread of exotic species. In the framework of set theory, interactions between biotic (B), abiotic (A), and movement-related (M) factors in the geographical space can be hypothesized with BAM diagrams and tested using ecological niche models (ENMs) to estimate A and B areas. The main aim of our survey was to evaluate the interactions between abiotic (climatic) and biotic (host availability) factors in geographical space for exotic symbionts (i.e., non-free-living species), using ENM techniques combined with a BAM framework and using exotic Entocytheridae (Ostracoda) found in Europe as model organisms. We carried out an extensive survey to evaluate the distribution of entocytherids hosted by crayfish in Europe by checking 94 European localities and 12 crayfish species. Both exotic entocytherid species found, Ankylocythere sinuosa and Uncinocythere occidentalis, were widely distributed in W Europe living on the exotic crayfish species Procambarus clarkii and Pacifastacus leniusculus, respectively. No entocytherids were observed in the remaining crayfish species. The suitable area for A. sinuosa was mainly restricted by its own limitations to minimum temperatures in W and N Europe and precipitation seasonality in circum-Mediterranean areas. Uncinocythere occidentalis was mostly restricted by host availability in circum-Mediterranean regions due to limitations of P. leniusculus to higher precipitation seasonality and maximum temperatures. The combination of ENMs with set theory allows studying the invasive biology of symbionts and provides clues about biogeographic barriers due to abiotic or biotic factors limiting the expansion of the symbiont in different regions of the invasive range. The relative importance of abiotic and biotic factors on geographical space can then be assessed and applied in conservation plans. This approach can also be implemented in other systems where the target species is closely interacting with other taxa.

Different Scales of Spatial Segregation of Two Species of Feather Mites on the Wings of a Passerine Bird

Abstract The “condition-specific competition hypothesis” proposes that coexistence of 2 species is possible when spatial or temporal variations in environmental conditions exist and each species responds differently to those conditions. The distribution of different species of feather mites on their hosts is known to be affected by intrinsic host factors such as structure of feathers and friction among feathers during flight, but there is also evidence that external factors such as humidity and temperature can affect mite distribution. Some feather mites have the capacity to move through the plumage rather rapidly, and within-host variation in intensity of sunlight could be one of the cues involved in these active displacements. We analyzed both the within- and between-feather spatial distribution of 2 mite species, Trouessartia bifurcata and Dolichodectes edwardsi, that coexist in flight feathers of the moustached warbler Acrocephalus melanopogon. A complex spatial segregation between the 2 species was observed at 3 spatial levels, i.e., “feather surfaces,” “between feathers,” and “within feathers.” Despite certain overlapping distribution among feathers, T. bifurcata dominated proximal and medial regions on dorsal faces, while D. edwardsi preferred disto-ventral feather areas. An experiment to check the behavioral response of T. bifurcata to sunlight showed that mites responded to light exposure by approaching the feather bases and even leaving its dorsal face. Spatial heterogeneity across the 3 analyzed levels, together with response to light and other particular species adaptations, may have played a role in the coexistence and segregation of feather mites competing for space and food in passerine birds.

A review of the Entocytheridae (Ostracoda) of the world: updated bibliographic and species checklists and global georeferenced database, with insights into host specificity and latitudinal patterns of species richness

The creation of biodiversity datasets freely available for the scientific community is a valuable task to stimulate global research on biodiversity. Among others, the Global Biodiversity Information Facility (GBIF) is a remarkable resource providing free online access to biodiversity data on many diverse taxonomic groups (including Ostracoda) from both terrestrial and marine ecosystems. More specific databases for Ostracoda georeferenced data have been built (e.g., NACODe and OMEGA), some with freely available data. However, the Entocytheridae, a family of ostracods living commensal on other crustaceans, with 220 living species, representing the third non-marine ostracod family in number of species, has been remarkably under-represented in the currently available biodiversity databases. To cover this gap, we present here a free-access world database of Entocytheridae published in GBIF and review the current knowledge of the group by updating a bibliographic and species checklists of the Entocytheridae. We also analyse the host specificity of the group and the latitudinal species richness pattern in North and Central America (including 186 spp.). The current database includes 3509 georeferenced records from 220 species, in contrast to just 44 entocytherid georeferenced records previously published in GBIF. In addition, the updated species list accounts for 43 species and 2 genera that were not included in the previous compendium on Entocytheridae published by Hart & Hart in 1974, so as 40 species not included in the 2013 Checklist provided by the Catalogue of Life. We show that the specialisation in one unique host species is not the rule in Entocytheridae, and evidence an unusual latitudinal pattern of species richness in North and Central American entocytherids, most probably related to the biogeographic history of their hosts.

Connectivity in Reed Warblers Acrocephalus scirpaceus Between Breeding Grounds in Europe and Autumn Stopover Sites in Iberia

We determined the degree of connectivity in the Eurasian Reed Warbler Acrocephalus scirpaceus between breeding regions in central and northern Europe and stopover sites in Iberia, during the autumn migration. We used both recovery data and wing length analyses to achieve this. Biometric data were obtained during 2009 at 11 sampling localities in Iberia, which we amalgamated into four major migratory areas within the region (Central, Western, Eastern and Southwestern Iberia, hereafter CI, WI, EI, SW). From the EURING database, we selected birds captured at their breeding sites in Europe and recaptured in Iberia during the autumn migration. Reed Warblers passing through WI had shorter wings than those in CI, EI and SW, suggesting that birds in WI migrated shorter distances, a fact also supported by recovery data. Although Reed Warblers showed some population overlap when passing through Iberia, we found that birds passing through EI and CI came from areas further to the east (continental Europe) than those passing through WI (mainly British Isles), thus supporting parallel migration. Reed Warblers tended to converge in southwestern Iberia, suggesting an effect of nearby geographical barriers.