The significance of subadult plumage in Darwin's finches, Geospiza fortis

I studied delayed plumage maturation, a common phenomenon inpasserine birds, in a species of Darwin's finch, Geospiza fortis,on Isla Daphne Major, Galapagos, Ecuador. To address the questionof why males advertise their youth and immaturity with subadultplumage, I determined the fitness consequences of possessingdifferent plumage states at different ages. Rate of entry intoadult plumage is both variable and heritable. In two cohorts,individuals that took longer to enter full adult plumage surviveddry conditions significantly better than those that acquiredadult plumage quickly. Behavioral observations helped revealthe reasons for this higher survival. Advantages arose fromthe avoidance of conflict that occurred by signaling subordinatestatus with subadult plumage in both the nonbreeding and breedingseasons. Although blacker males had a significantly lower survival,those black males that did survive had a reproductive advantagein their first breeding season over males with less black intheir plumage from the same cohort and raised significantlymore offspring to fledging. Thus the study demonstrates selectionon the length of the subadult plumage stage. [Behav Ecol 1990;l:161–170]     

Mechanical stress,fracture risk and beak evolution in Darwin's ground finches (Geospiza)

Darwin''s finches have radiated from a common ancestor into 14 descendent species, each specializing on distinct food resources and evolving divergent beak forms. Beak morphology in the ground finches (Geospiza) has been shown to evolve via natural selection in response to variation in food type, food availability and interspecific competition for food. From a mechanical perspective, however, beak size and shape are only indirectly related to birds'' abilities to crack seeds, and beak form is hypothesized to evolve mainly under selection for fracture avoidance. Here, we test the fracture-avoidance hypothesis using finite-element modelling. We find that across species, mechanical loading is similar and approaches reported values of bone strength, thus suggesting pervasive selection on fracture avoidance. Additionally, deep and wide beaks are better suited for dissipating stress than are more elongate beaks when scaled to common sizes and loadings. Our results illustrate that deep and wide beaks in ground finches enable reduction of areas with high stress and peak stress magnitudes, allowing birds to crack hard seeds while limiting the risk of beak failure. These results may explain strong selection on beak depth and width in natural populations of Darwin''s finches.     

Persistence of song types in Darwin's finches,Geospiza fortis,over four decades

Learned bird songs evolve via cultural evolution, with song patterns transmitted across generations by imitative learning. In Darwin''s finches of the Galápagos Islands, males learn songs from their fathers, and song types can be maintained across multiple generations. However, little is known about the time frame over which specific song types are preserved, in the face of copy errors and corresponding modifications to song structure. Here we investigate cultural evolution in songs of male Geospiza fortis, at Academy Bay, Santa Cruz Island, comparing songs recorded in 1961 by R. Bowman (20 individuals) to those recorded in 1999 by J. Podos (16 individuals). For each individual, we characterized four timing and six frequency parameters, and assessed inter-individual variation in song structure using multivariate analysis. Several 1961 song types persisted into 1999, some with remarkable fidelity. Variation among song types was extensive during both years, and we detected no changes in 10 vocal parameters across the sampling period. These results illustrate temporal continuity in a culturally acquired trait, and raise questions about mechanisms that promote stability in song structure.     

Growth and allometry of external morphology in Darwin's finches (Geospiza) on Isla Daphne Major, Galapagos

Seven widely used avian external morphological measurements were examined, mainly in Geospiza fortis and G. scandens on Isla Daphne Major, Galapagos. Geospiza grow more slowly than other finches, and smaller Geospiza species grow relatively more slowly than larger Geospiza . Weight, wing chord and tarsus grow quickly, while bill characters grow more slowly. This pattern of relative growth (dynamic allometry) is reversed in the static allometry of adults, where bill characters, particularly bill depth in G. fortis , display strong, positive allometry with respect to weight, wing and tarsus. Static allometric patterns help explain differences in the size, shape and relative variability of different characters in adults, as well as the results of multivariate analyses such as principal components or canonical variates. Several different multivariate techniques arrange five Geospiza populations in a consistent two-dimensional morphological space, with a major axis of overall body-size, and a minor axi of bill-pointedeness. An alternative analysis of dynamic and static allometry is also provided, based on multivariate techniques.     

Ecoimmunity in Darwin's Finches: Invasive Parasites Trigger Acquired Immunity in the Medium Ground Finch (Geospiza fortis)

Background

Invasive parasites are a major threat to island populations of animals. Darwin''s finches of the Galápagos Islands are under attack by introduced pox virus (Poxvirus avium) and nest flies (Philornis downsi). We developed assays for parasite-specific antibody responses in Darwin''s finches (Geospiza fortis), to test for relationships between adaptive immune responses to novel parasites and spatial-temporal variation in the occurrence of parasite pressure among G. fortis populations.

Methodology/Principal Findings

We developed enzyme-linked immunosorbent assays (ELISAs) for the presence of antibodies in the serum of Darwin''s finches specific to pox virus or Philornis proteins. We compared antibody levels between bird populations with and without evidence of pox infection (visible lesions), and among birds sampled before nesting (prior to nest-fly exposure) versus during nesting (with fly exposure). Birds from the Pox-positive population had higher levels of pox-binding antibodies. Philornis-binding antibody levels were higher in birds sampled during nesting. Female birds, which occupy the nest, had higher Philornis-binding antibody levels than males. The study was limited by an inability to confirm pox exposure independent of obvious lesions. However, the lasting effects of pox infection (e.g., scarring and lost digits) were expected to be reliable indicators of prior pox infection.

Conclusions/Significance

This is the first demonstration, to our knowledge, of parasite-specific antibody responses to multiple classes of parasites in a wild population of birds. Darwin''s finches initiated acquired immune responses to novel parasites. Our study has vital implications for invasion biology and ecological immunology. The adaptive immune response of Darwin''s finches may help combat the negative effects of parasitism. Alternatively, the physiological cost of mounting such a response could outweigh any benefits, accelerating population decline. Tests of the fitness implications of parasite-specific immune responses in Darwin''s finches are urgently needed.     

Isospora geospizae,a new coccidian parasite (Apicomplexa: Eimeriidae) from the small ground finch (Geospiza fuliginosa) and the medium ground finch (Geospiza fortis) from the Galapagos Islands

Isospora geospizae n. sp. is described from the faeces of the small ground finch Geospiza fuliginosa and the medium ground finch Geospiza fortis from Santa Cruz Island in the Galapagos archipelago. I. geospizae oocysts are spherical to subspherical, with a one-layered, smooth, colourless wall, 15.5×14.5 (13–17×12–17) m with a large polar body and no micropyle or residuum. Sporocysts are ovoid, 10×7.5 (10–12×6–9) m with small Stieda and subtieda bodies and a smoothly contoured, but irregular-shaped residuum containing coarse, non-uniform granules. Sporozoites are sausage-shaped with large refractile bodies at each end and randomly arranged in the sporocysts. Three of 34 (8.8%) G. fuliginosa and one of nine (11.1%) G. fortis were infected with I. geospizae.     

The classical case of character release: Darwin's finches (Geospiza) on Isla Daphne Major, Galápagos

The history of how Darwin's medium ground finch (Geospiza fortis) on Isla Daphne Major, Galápagos became a textbook example of character release is reviewed. Four hypotheses for the evolution of the intermediate-sized Daphne fortis are examined, including genetic drift/founder effect, hybridization with the small ground finch (G'. fuliginosa) , food supply (or local adaptation hypothesis) and character release in allopatry.
Modern data suggest that genetic drift is unlikely to have been important, due to inadequate isolation and over-riding selection and introgression on Daphne. All three remaining hypotheses have probably played a role. Hybridization with G. fuliginosa occurs, although it cannot counteract the selection pressures seen during our study. Local adaptation has also occurred, with natural selection changing the relative frequencies of fortis phenotypes in response to changes in Daphne food supplies. The selection resulted from correlations between the size of seeds available, feeding behaviour and morphology. However, recent phenotypic tracking has resulted in larger, not smaller phenotypes. There is also evidence for character release in the form of diet expansion by G. fortis during periods of food shortage, and indirect evidence for interspecific competition between fortis and the cactus ground finch (G. scandens). The Daphne fortis phenotype probably represents a balance between introgression with fuliginosa, selection for larger body size in dry years and selection for smaller body size in wet years. The simple textbook account of a character shift caused by the accidental absence of competitors should be qualified to reflect the ecological complexity of the situation.     

Isospora daphnensis n. sp. (Apicomplexa: Eimeriidae) from the medium ground finch (Geospiza fortis) from the Galapagos Islands

In March and April 1987 fecal samples from 237 nestling birds, including 199 from 85 nest sites of Geospiza fortis, 23 from 12 nest sites of Geospiza scandens, 6 from 2 nest sites of Geospiza magnirostris, and 9 from 2 nest sites of hybrids involving Geospiza fuliginosa and G. fortis, were collected from Daphne Major in the Galapagos archipelago and examined for coccidia. Only 3 of 4 nestlings from 1 nest site of G. fortis (1.5%) had oocysts in their feces. Two of the 3 infected nestlings had concurrent infections of Isospora temeraria, and all 3 nestlings were infected with a new species. Isospora daphnensis n. sp. Sporulated oocysts of I. daphnensis n. sp. are ellipsoidal, 27.3 x 23.6 (22-30 x 20-27) microns; a polar body is present, but no oocyst residuum or micropyle occurs. The oocyst wall, approximately 1.5 microns thick, is composed of a mammillated outer layer and thinner inner layer. Sporocysts are ovoid, 15.2 x 10.2 (15-16 x 9-11) microns and have a nipplelike Stieda body and a small substieda body. The sporocysts contain an irregularly shaped, smoothly contoured residuum with uniform granules and 4 sporozoites with a large refractile body at one end and lying randomly in the sporocysts.     

Pedigrees, assortative mating and speciation in Darwin's finches

Pedigree analysis is a useful tool in the study of speciation. It can reveal trans-generational influences on the choice of mates. We examined mating patterns in a population of Darwin's medium ground finches (Geospiza fortis) on Daphne Major Island to improve our understanding of how a barrier to the exchange of genes between populations arises in evolution. Body sizes of mates were weakly correlated. In one year, the smallest females were paired non-randomly with the males of similar size, and in another year the largest males were paired with the largest females. An influence of parental morphology on the choice of mates, as expected from sexual imprinting theory, was found; the body size of mates was predicted by the body sizes of both parents, and especially strongly by the father's. These associations imply that the seeds of reproductive isolation between species are present within a single variable population. The implication was subject to a natural test: two exceptionally large birds of the study species, apparently immigrants, bred with each other, as did their offspring, and not with the members of the resident population. The intense inbreeding represents incipient speciation. It parallels a similar phenomenon when another species, the large ground finch, immigrated to Daphne and established a new population without interbreeding with the resident medium ground finches.     

Rare variants, private polymorphisms, and locus heterozygosity in Amerindian populations.

The results of 21,103 electrophoretic typings distributed across 28 polypeptides in members of 12 Amerindian tribes are reported, and the accumulated results of electrophoretic studies on these same polypeptides in 21 Amerindian tribes are then analyzed. Thus far 11 'private' polymorphisms have been identified in these tribes. When the tribal samples are combined and traits achieving polymorphic proportions in the total sample excluded from consideration, the average frequency of rare variants is 2.8 per 1,000 determinations. For a subset of 23 of these polypeptides also studied in Caucasians and Japanese, variant frequencies per 1,000 determinations are: Indians, 2.2; Caucasians (British), 1.6; and Japanese, 1.5. Average locus heterogeneity for these polypeptides (based on rare variants plus polymorphisms) is: Indians, .049; Caucasians, .078; and Japanese, .077. A higher proportion of loci are monomorphic within tribes than within civilized urban populations. It is argued that for inferences concerning the forces maintaining genetic variability within populations, studies on samples from tribespeople are much more appropriate than studies on samples from civilized urban populations.     

Homeostasis,enzymatic heterozygosity and inbreeding depression in natural populations of Drosophila melanogaster

Brother-sister mating effects on offspring viability in natural populations of Drosophila melanogaster were analysed in relation to developmental homeostasis and enzymatic heterozygosity of the individuals constituting the sib pairs. High developmental homeostasis and high degree of heterozygosity are associated with a low inbreeding depression in offspring and, therefore, with a low frequency of major deleterious factors. These results are discussed in relation to maintenance of polymorphism in natural populations and the role of genetic load and founder events in speciation and adaptation. We conclude that an unusual environmental condition followed by a strong reduction in effective population size due to random elimination of individuals is a better candidate to speciation than an environmental shift leading to selection for homeostatic, highly heterozygous lethal-free individuals.     

Contrasting selection modes at the Adh1 locus in outcrossing Miscanthus sinensis vs. inbreeding Miscanthus condensatus (Poaceae)

We estimated DNA sequence variation of the Adh1 locus in the outcrossing Miscanthus sinensis (Poaceae) and its close selfing relative, M. condensatus. Tajima's test of selection is significantly negative for both overall exons and replacement sites in M. sinensis. Among its entire sample, nucleotide diversity of nonsynonymous sites is higher than that of synonymous sites. A McDonald and Kreitman test of neutrality indicates an excess of intraspecific replacement polymorphisms, suggesting possible directional selection toward advantageous mutants. However, frequent intragenic recombination suggests both purifying and positive selection is unlikely. Recent demographic expansions coupled with relaxation of purifying selection may have resulted in elevated genetic diversity at the Adh1 locus as well as the trnL-trnF intergenic spacer of cpDNA in this outcrossing species. In contrast, low levels of genetic diversity were detected at both the Adh1 locus and the cpDNA spacer in M. condensatus, consistent with bottlenecks associated with selfing in all populations. While Tajima's D and Fu and Li's F statistics did not reveal deviation from neutrality at the Adh1 locus in M. condensatus, 12 replacements vs. 10 synonymous changes were detected. Based on pairwise comparisons of the d(N)/d(S) ratio, lineages of closely related populations of the species distributed along saline habitats appeared to be under directional selection.     

Inbreeding in Pinus radiata. I. The effect of inbreeding on growth, survival and variance

 The effects of inbreeding on growth, survival and variance in a 12-year-old radiata pine trial were studied in five populations each inbred to one of five different levels: outcross (F=0), half-sib (F=0.125), full-sib (F=0.25), selfing (S₁, F=0.5), and two-generations of selfing (S₂, F=0.75). These five populations were derived from a founder population of eight clones. Inbreeding reduced diameter, growth, and survival but increased the variance for diameter. Inbreeding depression at F=0.125, 0.25, 0.5, and 0.75 was 5%, 6%, 15%, and 19% respectively for DBH; −3%, 1%, 7%, and 11% respectively, for survival. The standard deviation for diameter increased by 10%, 10%, 30%, and 25% respectively for F=0.125, 0.25, 0.5, and 0.75 and, similarly, the coefficient of variation increased by 17%, 16%, 53%, and 55% respectively. There were significant differences among the eight founder clones in their response to inbreeding. The best clone in the trial showed no inbreeding depression. Overall, inbreeding depression was found to be linearly related to the inbreeding coefficient F with no significant quadratic effects for any trait at any population level. However, two individual clones had a quadratic relationship with F for DBH and one clone had a similar relationship for survival. A significant correlation (r=0.96) between S₂ and the breeding values of founder clones was observed while the correlation (r=0.58) between S₁ and breeding values was insignificant. The low inbreeding depression in radiata pine relative to other conifers may indicate that historical purging of detrimental alleles through small geographic populations, a higher degree of population subdivision, and the relative high fecundity of inbred progenies has rendered radiata pine an ideal species to use inbreeding as a breeding tool. Received: 10 March 1998 / Accepted: 19 May 1998     

Relationship between inbreeding depression and inbreeding coefficient in maritime pine (Pinus pinaster)

The relationship between inbreeding depression and inbreeding coefficient (F) for several important traits was investigated in an 11-year trial of maritime pine (Pinus pinaster). Five levels of inbreeding (F=0; 0.125; 0.25; 0.5; 0.75) were obtained in a mating design involving ten plus-trees, or their progenies, as parents (total of 51 families). For F=0.75, the mean inbreeding depressions were 27% for height, 37% for circumference at breast height (63% for bole volume), 23% for basal straightness (better straightness of the inbred trees), and 89% for female fertility (number of cones). Large differences were observed among inbred families for the same level of inbreeding. The evolution of depression with F was more or less linear, depending on the traits. Significant differences among F-levels appeared very early for height (from 5-years of age). Inbreeding depression was much more expressed during unfavorable years than during favorable years for yearly height growth. When compared with other Pinus species, maritime pine appears to be less affected by inbreeding, especially for the percentage of filled seeds and general vigor. A reduced genetic load in maritime pine may result from the evolutionary history of the species and its scattered distribution.     

Dicentra scandens (D. Don) Walp. auch in der Flora Vietnams

The specimen 3064 of A. Petelot in 1927 from Chapa (Vietnam) is Dicentra scandens (D. Don) Walp. Anatomical characteristics of fruits and seeds are identically with such of the isotype, but morphological characteristics in the other parts are somewhat different.     

Reproductive responses to varying food supply in a population of Darwin's finches: Clutch size,growth rates and hatching synchrony

Summary I show how food shortage affects reproduction in a population of Darwin's Medium Ground Finches, Geospiza fortis. Despite the common occurrence of starvation and absence of nest predation, hatching is typically nighly synchronous and adaptive brood reductionappears to be absent. Variation in both growth rates and clutch size in association with the varying conditions is documented. This variation is interpreted as being a direct response to environmental conditions rather than adaptive phenotypic plasticity. I conclude that selection pressures to raise one or two chicks during times of food shortage, or to delay growth rates, are weak or absent.     

Gene mapping and serendipity. The locus for torticollis,keloids, cryptorchidism and renal dysplasia (31430, McKusick) is at Xq28, distal to the G6PD locus

Summary The syndrome of torticollis, keloids, cryptorchidism and renal dysplasia was described by Goeminne (1968) in a large family, and listed as X-linked incomplete dominant by McKusick (1978; No. 31430). We mapped the locus on the long arm of the X chromosome at band q28 and distal to the G6PD locus. This was achieved by the chance discovery in the literature that two females, each with a balanced X/autosome translocation involving Xq28, had partial manifestation of the syndrome as was the case with the females in the larger family of Goeminne (1968).     

Colour bands,dominance, and body mass regulation in male zebra finches (Taeniopygia guttata)

The arbitrary assignment of different coloured leg bands to zebra finches (Taeniopygia guttata) has profound effects on mate preference, reproductive success, mortality rates, parental investment and sex ratio. Choice chamber experiments indicate that the effect is mediated by altered attractiveness to members of the opposite sex. Effects on intrasexual dominance are more equivocal. We present two experiments which demonstrate significant effects of band colour on behavioural dominance (red bands are more dominant than light green bands) and the resulting diurnal pattern of gain in mass, fat, and seeds stored in the crop. Consistent with the literature on dominance and strategic regulation of body mass in other species, subordinate (green-banded) birds maintain higher fat reserves at dawn, but dominant (red-banded) birds show the highest overall daily mass gains. The lack of obvious effects of band colour on dominance in previous studies may lie in the degree to which food can be monopolized by particular individuals.