Zonotrichia Capensis Research Articles

Seasonal Space use and Territory Size of Resident Sparrows in the Central Monte Desert, Argentina

Knowledge of space use and territoriality of New World birds largely derives from studies in temperate North America and the tropics, whereas it remains poorly studied in temperate South America, particularly in arid environments. Here we studied the seasonal space use and territory size among resident individuals of three sparrows in a temperate semi-desert of Argentina: the ringed warbling-finch Poospiza torquata, the rufous-collared sparrow Zonotrichia capensis and the manycoloured Chaco-finch Saltatricula multicolor. Both territorial behaviour and use of space of the three species showed seasonal changes. The ringed warbling-finch and the rufous-collared sparrow used larger areas during the non-breeding season than during the breeding season, while the many-coloured Chaco-finch used areas of similar size in the two periods. Defence activities were absent for all three species during the non-breeding season. During the breeding season, each species defended territories that were smaller for the rufous-collared sparrow (0.79 ± 0.04 ha) than for the ringed warbling-finch (1.35 ± 0.04 ha) and the many-coloured Chaco-finch (1.21 ± 0.07 ha). Seasonal territoriality and territory size reported for resident individuals were similar to those reported for north-temperate sparrows in xeric environments.

Metabolic enzymes in seasonally acclimatized and cold acclimated rufous-collared sparrow inhabiting a Chilean Mediterranean environment

Abstract Due to the higher energy requirements of birds during winter, it is predicted that the activities of metabolic enzymes (e.g., citrate synthase, CS and cytochrome C oxidase, COX) should increase in tandem with increases in rates of energy expenditure (e.g., basal metabolic rate, BMR). However, there is mixed support for the hypothesis of enzymatic acclimatization. Furthermore, there is little information about the effect of ambient temperature on energetics and tissue enzyme activity levels in passerines inhabiting seasonal Mediterranean environments. In this study we evaluated the interplay between BMR and enzyme activities of freshly caught individuals of the passerine Zonotrichia capensis in winter and summer in a Mediterranean environment from central Chile, and also in warm (30°C) and cold (15°C) lab-acclimated birds. The results revealed a lack of seasonal variation in BMR, thermal conductance and in the activity of CS and COX. However, we found higher BMR and lower thermal conductance in cold-acclimated than in warm-acclimated birds. Also, total CS activity was higher in the flight muscles of cold-acclimated than in warm-acclimated birds. We found also a significant correlation between BMR and total CS activity in pectoral muscle. Although some authors have suggested that BMR mainly depends on the metabolic intensity and mass of internal organs, our results revealed that skeletal muscles such as the flight muscles may also affect BMR. Finally, differences in the acclimation and acclimatization responses in Rufous-collared sparrows suggest that large-scale oscillations in the physical environment might maintain the capacity for flexibility in thermogenic traits through generations.

Secondary contact followed by gene flow between divergent mitochondrial lineages of a widespread Neotropical songbird (Zonotrichia capensis)

1 * Understanding how genetic and phenotypic differences that arise in geographically isolated populations influence the outcome of secondary contact advances our knowledge of speciation. In the present study, we investigate the secondary contact between divergent lineages of a widespread Neotropical songbird, the Rufous-collared sparrow (Zonotrichia capensis). Zonotrichia capensis is morphologically and behaviourally diverse, and shows a pattern of lineage diversification produced by a Pleistocene expansion and colonization of South America from a probable Central American origin. Consistent with previous results, we find three lineages throughout the species range, showing between 1.5% and 2.5% divergence in mitochondrial control region sequences. These lineages come into secondary contact in the Dominican Republic, La Paz (Bolivia), and North-eastern Argentina. We use DNA microsatellite data to study a broad secondary contact zone in North-eastern Argentina, finding that Bayesian clustering analyses do not assign individuals to their respective mitochondrial lineages. Overall, we did not observe nuclear genetic discontinuities in the study area. We conclude that, if genetic, morphological, and/or cultural differences accumulated among lineages during isolation, they were insufficient to prevent gene flow after secondary contact. © 2014 The Linnean Society of London, Biological Journal of the Linnean Society, 2014, 111, 863-868.

Seasonal Modulation of Testosterone during Breeding of the Rufous-Collared Sparrow (Zonotrichia capensis australis) in Southern Patagonia

The roles of testosterone (T) during reproduction are often complex and vary among and within vertebrate taxa and locations, making general hypotheses relating T to breeding behavior and success difficult to integrate. In birds, T is thought to influence degrees of territoriality and associated aggression in males to maximize breeding success. Importantly, most work supporting these ideas has been conducted in the Northern Hemisphere. However, accumulating work on tropical species has shown divergent patterns of T in association with breeding behavior. The compilation of work from northern temperate and tropical species suggests that the function of T in relation to breeding behavior varies across latitude and environmental conditions. We investigate the patterns of T in relation to breeding behavior in a subspecies of the rufous-collared sparrow Zonotrichia capensis australis breeding at high latitude in the Southern Hemisphere (55°S). We then compare the T profiles and breeding behaviors of male Z. c. australis to conspecifics breeding in the tropics and congeners in North America to test the hypothesis that environments with breeding seasons of similar lengths will drive similar patterns of T in relation to breeding behavior. We found that Z. c. australis have high levels of T during the early-breeding periods when territories are being established and low levels of T during the parental phase of breeding, similar to temperate and Arctic birds in the Northern Hemisphere but unlike tropical Zonotrichia capensis costaricensis. In contrast, we found that Z. c. australis also exhibit similar aggressive behaviors in early breeding and midbreeding, unlike many birds in the Northern Hemisphere.

High rates of extra‐pair paternity in two equatorial populations of rufous‐collared sparrow, Zonotrichia capensis

The latitudinal increase in extra‐pair paternity (EPP) rates in birds suggests broad selective benefits to low EPP rates in the tropics. However, we have few EPP data from tropical birds, particularly from species with close relatives at high latitudes. Here, we report EPP rates in two resident equatorial populations of rufous‐collared sparrow Zonotrichia capensis, a genus well‐represented at high latitudes. We found 64% and 60% of broods contained extra‐pair offspring, and 42% and 52% of all young were extra‐pair. EPP rates were similar in these populations, despite clear differences in elevation, temperature, rainfall, and breeding season length. These findings provide evidence that EPP rates in tropical birds can be as high as those observed in temperate birds, and suggest that the selective pressures acting on EPP rates vary markedly across tropical birds.

Can nest predation explain the lack of defenses against cowbird brood parasitism in the Rufous-collared Sparrow (Zonotrichia capensis)?

ABSTRACT. Although interspecific avian brood parasitism usually lowers host productivity, some species lack any defense against parasites. We analyzed the effect of parasite egg removal or nest desertion following a parasitism event on the breeding productivity of the Rufous-collared Sparrow (Zonotrichia capensis), a common host of the Shiny Cowbird (Molothrus bonariensis). The Rufouscollared Sparrow is an effective cowbird host that does not eject parasite eggs. We removed cowbird eggs from nests of Rufouscollared Sparrows to test for differences in hatching, fledging, and nesting success among naturally unparasitized, parasitized, and experimentally unparasitized nests from which we removed the cowbird eggs. We also used simulations to test whether parasite egg removal or nest desertion provide viable strategies to counter the effects of parasitism in this species. Naturally unparasitized nests produced more nestlings and fledglings than parasitized and experimentally unparasitized nests, but there were...

Breaking down seasonality: Androgen modulation and stress response in a highly stable environment

Previous studies show that most birds inhabiting temperate regions have well defined life history stages, and they modulate the production of testosterone (T) and corticosterone (CORT) in response to changes in seasonality. In this study we aimed to examine baseline and stress-induced levels of CORT and circulating T in relation with life history stages in the rufous-collared sparrow, Zonotrichia capensis. We carried out this study for a year in a population inhabiting riparian habitats in the Atacama Desert in Chile, one of the most climatically stable and driest places in the world. This environment shows minimal yearly change in average temperature and precipitation is virtually zero. We found individuals breeding, molting and overlapping breeding and molt year round, although most individuals were molting during March and in breeding condition during October. T levels were not related to individual breeding condition, and at population level they were not significantly different across sampling months. Baseline levels of CORT did not vary across the year. Stress-induced levels of CORT were suppressed during March when most of the birds were molting. This phenomenon was also observed in birds not molting during this period suggesting a mechanism other than molt in determining the stress-response suppression. Our results strongly suggest that in this study site, long-term extremely stable conditions could have relaxed the selective pressures over the timing of life history stages which was evidenced by the breeding and molt schedules, its overlap and endocrine profiles.

Spatial Patterns of Avian Malaria Prevalence inZonotrichia capensison the Western Slope of the Peruvian Andes

Environmental heterogeneity largely dictates the spatial distributions of parasites and therefore the susceptibility to infection of host populations. We surveyed avian malaria infections in Rufous-collared sparrows (Zonotrichia capensis) across replicated altitudinal and latitudinal transects along the western slope of the Peruvian Andes to assess geographic patterns of prevalence. We found malaria infection prevalence peaked at midelevations along all 3 altitudinal transects (x ≈ 2,733 m), with highest overall prevalence observed in the northern transect. We observed low levels of malarial parasite diversity, with 94% of infected birds carrying a single Haemoproteus (subgenus Parahaemoproteus) strain. The remaining infected birds harbored either a single alternate Haemoproteus or 1 of 2 Plasmodium strains. Our data suggest that temperature and precipitation are the primary drivers of the spatial patterns in avian malaria prevalence along the western slope of the Andes.

Effects of food supplementation on a tropical bird

Tropical birds typically exhibit a 'slow pace of life' relative to higher latitude species. This is often manifested as slow development, low fecundity, and high survival. Following from this, it is predicted that tropical birds may be more likely to trade current reproductive effort to favor self-maintenance, thus supporting survival and future reproduction. To test this idea, we conducted two food supplementation experiments on tropical rufous-collared sparrows (Zonotrichia capensis) in the eastern Andes of Ecuador. In the first experiment, we food-supplemented pairs during the non-breeding life-history stage, and in the second experiment, we food-supplemented pairs that were provisioning fledglings. In both experiments, a larger proportion of food-supplemented birds exhibited pre-basic molt (replacement of feathers) than in a control group. To our knowledge, this is the first study to experimentally demonstrate that a food-supplemented bird invests extra resources into molt, a form of self-maintenance, and contrasts with the majority of food supplementation studies in high latitude birds that show they typically advance the initiation of, or extend the period of, reproduction. Our results are consistent with the syndrome of the slow pace of life in the tropics and support the concept of fundamental differences between temperate-zone and tropical birds.

Continental phylogeography of an ecologically and morphologically diverse Neotropical songbird, Zonotrichia capensis.

BackgroundThe Neotropics are exceptionally diverse, containing roughly one third of all extant bird species on Earth. This remarkable species richness is thought to be a consequence of processes associated with both Andean orogenesis throughout the Tertiary, and climatic fluctuations during the Quaternary. Phylogeographic studies allow insights into how such events might have influenced evolutionary trajectories of species and ultimately contribute to a better understanding of speciation. Studies on continentally distributed species are of particular interest because different populations of such taxa may show genetic signatures of events that impacted the continent-wide biota. Here we evaluate the genealogical history of one of the world’s most broadly-distributed and polytypic passerines, the rufous-collared sparrow (Zonotrichia capensis).ResultsWe obtained control region DNA sequences from 92 Zonotrichia capensis individuals sampled across the species’ range (Central and South America). Six additional molecular markers, both nuclear and mitochondrial, were sequenced for a subset of individuals with divergent control region haplotypes. Median-joining network analysis, and Bayesian and maximum parsimony phylogenetic analyses all recovered three lineages: one spanning Middle America, the Dominican Republic, and north-western South America; one encompassing the Dominican Republic, Roraima (Venezuela) and La Paz (Bolivia) south to Tierra del Fuego, Argentina; and a third, including eastern Argentina and Brazil. Phylogenetic analyses suggest that the Middle American/north-western South American clade is sister to the remaining two. Bayesian and maximum likelihood coalescent simulations used to study lineage demographic history, diversification times, migration rates and population expansion together suggested that diversification of the three lineages occurred rapidly during the Pleistocene, with negligible gene flow, leaving genetic signatures of population expansions.ConclusionsThe Pleistocene history of the rufous-collared sparrow involved extensive range expansion from a probable Central American origin. Its remarkable morphological and behavioral diversity probably represents recent responses to local conditions overlying deeper patterns of lineage diversity, which are themselves produced by isolation and the history of colonization of South America.

Osmoregulatory and metabolic costs of salt excretion in the Rufous-collared sparrow Zonotrichia capensis

Recent experiments on shorebirds have demonstrated that maintaining an active osmoregulatory machinery is energetically expensive. This may, in part, explain diet and habitat selection in birds with salt glands. However little is known about the osmoregulatory costs in birds lacking functional salt glands. In these birds, osmotic work is done almost exclusively by the kidneys. We investigated the osmoregulatory cost in a bird species lacking functional salt glands, the passerine Zonotrichia capensis. After 20days of acclimation to fresh water (FW) and salt water (200mM NaCl, SW), SW birds tended to be heavier than FW birds. However, this difference was not statistically significant. Total basal metabolic rate was higher in SW birds as compared with FW birds. Renal and heart masses were also higher in the SW group. We also found greater medullary development and an increase in urine osmolality in the SW group. In spite of Z. capensis' ability to tolerate a moderate salt load in the laboratory, we hypothesize that increased cost of maintenance produced by salt consumption may significantly affect energy budget, dietary, and habitat choices in the field.

Geographic Variation in the Association between Exploratory Behavior and Physiology in Rufous-Collared Sparrows

Increasing research has attempted to clarify the links between animal personality and physiology. However, the mechanisms driving this association remain largely unknown, and knowledge of how ecological factors may affect its direction and strength is scant. In this study, we quantified variation in the association between exploratory behavior, basal metabolic rate (BMR), and total evaporative water loss (TEWL) in rufous-collared sparrows (Zonotrichia capensis) inhabiting desert, Mediterranean, and cold-temperate climates. We found that the exploratory behavior score was highest in birds from the cold-temperate site, which was characterized by a moderate level of ecological variability (seasonality). Moreover, the association between exploratory behavior and physiological variables differed among localities. Only birds from the Mediterranean site showed a positive correlation between exploratory behavior and BMR. We found no association between exploration and TEWL at any study site. Our findings suggest that differences in the ecological conditions experienced by each sparrow population result in a particular combination of behavioral and physiological traits. An understanding of this intraspecific variation along ecological gradients provides unique insights into how specific ecological conditions affect the coupling of behavioral and physiological traits and the mechanisms underlying that relationship.

Tolerance to Dietary Phenolics and Diet Breadth in Three Seed‐eating Birds: Implications for Graminivory

The ability to cope with plant secondary compounds (PSCs) has profound implications for an animal's behavior. In the present study, we assessed the tolerance to dietary phenolics in three seed-eating birds: Zonotrichia capensis, Saltatricula multicolor, and Diuca diuca, which differ in their diet breadth. Seeds in their habitat have distinct chemical composition: grass seeds have less PSCs, specifically, less total phenolics than forb seeds. Based on the detoxification limitation hypothesis and using published data of the natural history of these birds in the central Monte desert, we postulate that predominantly and exclusively graminivorous birds such as D. diuca and S. multicolor, respectively, are less tolerant due, in part, to a lower detoxification capacity than those with greater diet breadth, Z. capensis. To achieve this goal, we measured the food intake of diets varying in their concentration of tannic acid (TA). Indicators of tolerance were body mass change, food, TA and water intake, and glucuronic acid output throughout the experiment. Zonotrichia capensis performed better along the tolerance experiment: it maintained body mass from 0 to 4% TA diet, food and TA intake were higher than the other two species at the end of the experiment, and glucuronic acid output by Z. capensis was greater than D. diuca and S. multicolor from 2% TA diet until the end of the experiment. Our results suggest that Z. capensis is the most tolerant species and this physiological trait may explain their greater diet breadth.

Variation in the gonadotrophin-releasing hormone-1 and the song control system in the tropical breeding rufous-collared sparrow (Zonotrichia capensis) is dependent on sex and reproductive state

Seasonal breeding in temperate zone vertebrates is characterised by pronounced variation in both central and peripheral reproductive physiology as well as behaviour. In contrast, many tropical species have a comparatively longer and less of a seasonal pattern of breeding than their temperate zone counterparts. These extended, more “flexible” reproductive periods may be associate with a lesser degree of annual variation in reproductive physiology. Here we investigated variation in the neuroendocrine control of reproduction in relation to the changes in the neural song control system in a tropical breeding songbird the rufous-collared sparrows (Zonotrichia capensis). Using in situ hybridization, we show that the optical density of GnRH1 mRNA expression is relatively constant across pre-breeding and breeding states. However, males were found to have significantly greater expression compared to females regardless of breeding state. Both males and females showed marked variation in measures of peripheral reproductive physiology with greater gonadal volumes and concentrations of sex steroids in the blood (i.e. testosterone in males; estrogen in females) during the breeding season as compared to the pre-breeding season. These findings suggest that the environmental cues regulating breeding in a tropical breeding bird ultimately exert their effects on physiology at the level of the median eminence and regulate the release of GnRH1. In addition, histological analysis of the song control system HVC, RA and Area X revealed that breeding males had significantly larger volumes of these brain nuclei as compared to non-breeding males, breeding females, and non-breeding females. Females did not exhibit a significant difference in the size of song control regions across breeding states. Together, these data show a marked sex difference in the extent to which there is breeding-associated variation in reproductive physiology and brain plasticity that is dependent on the reproductive state in a tropical breeding songbird.

Phenotypic flexibility in basal metabolic rate is associated with rainfall variability among populations of rufous-collared sparrow

Phenotypic flexibility in metabolic rates allows organisms to reversibly adjust their energy flow to meet challenges imposed by a variable environment. In turn, the food habits hypothesis (FHH) predicts that species or populations adjust their basal metabolic rate (BMR) according to the diet attributes such as food abundance or predictability. Desert ecosystems represent a temporally heterogeneous environment because of low rain pulse predictability, which is also associated with temporal variation in food resources. In the present study, we investigated the relationship between the magnitude of BMR flexibility in response to dietary acclimation and the inter-annual rainfall variability in three populations of rufous-collared sparrows. Specifically we addressed the question of whether birds from a desert environment are more flexible in BMR than those from non-desert habitats. We found a positive trend between BMR flexibility and the inter-annual rainfall variability. In fact, dietary treatments had a significant effect only in desert birds, a result that also supported the FHH. Our study confirms the existence of phenotypic variation in response to environmental conditions among populations, and also highlights the importance of considering the circumstances in which phenotypic flexibility evolves and the specific environmental cues that induce their expression.

Effects of Nutritional and Anti-Nutritional Properties of Seeds on the Feeding Ecology of Seed-Eating Birds of the Monte Desert, Argentina

Abstract. Food quality and physiological capacities and limitations in recognizing and processing food are among the factors influencing the choices of desert birds in feeding. Seed-eating birds of the central Monte desert generally select grass seeds rather than forb seeds. We studied some of the mechanisms underlying seed-selection patterns in seed-eating birds of the Monte desert, analyzing nutrients and secondary compounds of the 15 most abundant seeds in the soil and their relationship with the diet of six species of seed-eating birds. Grass seeds contained more starch and less total phenols than did forb seeds and were free from alkaloids. The diet of the most graminivorous birds was correlated with seeds' starch concentration, while generalists' diet was correlated with seed abundance. To assess the plausible mechanisms underlying birds' selection of seed, we experimented with three species differing in the breadth of their diet: a generalist, the Rufous-collared Sparrow (Zonotrichia capensis), and...

Responses in the abundance of generalist birds to environmental gradients: The rufous-collared sparrow (Zonotrichia capensis) in the southern Neotropics

Bird species are adapted to certain ranges of physical conditions and will respond to environmental heterogeneity depending on their ability to exploit a broad range of resources and their tolerance to new environmental conditions. We used the rufous-collared sparrow as a model to explore how the abundance of generalist passerine birds may vary along environmental gradients in the southern Neotropics. We analyzed variations in the abundance along 6 gradients expressing variations in climate, productivity, the proportion of native forest in the landscape, the proportion of vegetation types through the transition between regions, and the intensity of human activities (agriculture, urbanization). The rufous-collared sparrow was most abundant in seasonal and semi-arid climates. Thermal amplitude was the best climatic predictor of the species abundance at the large scale. Within regions where climatic conditions are relatively homogeneous, land use better predicted abundance patterns. The species responded positively to increasing primary productivity, agricultural intensity, and native forest degradation and conversion and negatively to increasing urbanization. The rufous-collared sparrow adapts successfully to new environments created by human activities such as agriculture, ranching, forestry, and urbanization. Some native species may be tolerant to certain types and intensity of human activities, and knowledge of how they respond to both natural and human-created environments may help efforts to anticipate the impact of human activities on native birds in a changing world.

Within-Species Digestive Tract Flexibility in Rufous-Collared Sparrows and the Climatic Variability Hypothesis

The climatic variability hypothesis (CVH) states that species are geographically more widespread at higher latitudes because individuals have a broader range of physiological tolerance or phenotypic flexibility as latitude and climatic variability increase. However, it remains unclear to what extent climatic variability or latitude, acting on the phenotype, account for any observed geographical gradient in mean range size. In this study, we analyzed the physiological flexibility within the CVH framework by using an intraspecific population experimental approach. We tested for a positive relationship between digestive-tract flexibility (i.e., morphology and enzyme activities) and latitude and climatic and natural diet variability in populations of rufous-collared sparrows (Zonotrichia capensis) captured in desert (27°S), Mediterranean (33°S), and cold-temperate (41°S) sites in Chile. In accordance with the CVH, we observed a positive relationship between the magnitude of digestive-tract flexibility and environmental variability but not latitude. The greatest digestive flexibility was observed in birds at middle latitudes, which experience the most environmental variability (a Mediterranean climate), whereas individuals from the most stable climates (desert and cold-temperate) exhibited little or no digestive-tract flexibility in response to experimental diets. Our findings support the idea that latitudinal gradients in geographical ranges may be strongly affected by the action of regional features, which makes it difficult to find general patterns in the distribution of species.

Female, but Not Male, Tropical Sparrows Respond More Strongly to the Local Song Dialect: Implications for Population Divergence

In addition to the observed high diversity of species in the tropics, divergence among populations of the same species exists over short geographic distances in both phenotypic traits and neutral genetic markers. Divergence among populations suggests great potential for the evolution of reproductive isolation and eventual speciation. In birds, song can evolve quickly through cultural transmission and result in regional dialects, which can be a critical component of reproductive isolation through variation in female preference. We examined female and male behavioral responses to local and nonlocal dialects in two allopatric populations of rufous-collared sparrows (Zonotrichia capensis) in the Andes Mountains of Ecuador. Here we show that female sparrows prefer their natal song dialect to the dialect of an allopatric population that is just 25 km away and separated by an unsuitable higher-elevation habitat (pass of 4,200 m), thus providing evidence of prezygotic reproductive isolation among populations. Males showed similar territorial responses to all conspecific dialects with no consistent difference with respect to distance, making male territoriality uninformative for estimating reproductive isolation. This study provides novel evidence for culturally based prezygotic isolation over very short distances in a tropical bird.

Homogenisation of birdsong along a natural–urban gradient in Argentina

One of the clearest examples of animal adaptation to urban habitats is the variation in their vocalisations driven by anthropogenic noise. A behavioural process that has received less attention in urban habitats is how anthropogenic barriers influence animal spacing behaviour and communication. I addressed the effects of the distribution of bird territories in urban habitats and the pattern of song differentiation over distance among neighbouring birds. I studied natural and suburban populations of the white-crowned sparrow Zonotrichia capensis in Argentina. This songbird dwells in gardens, but is here less abundant and more regularly spaced than in open woodlands. Neighbouring birds in garden suburbs sang songs that were more similar to each other than neighbouring birds in natural habitats. Song differentiation was greater within natural habitats than within suburbs at distance < 400 m. At larger distances differences among habitats were no longer significant. Neighbour songs were more homogeneous where territories were more regularly spaced, considering all habitats or urban ones only. Selection for individual recognition of neighbours through songs and greater male–male interactions could have enhanced acoustic differentiation in natural habitats. Conversely, weak territorial interactions in suburbs with abundant landmarks may have constrained song output, resulting in song homogenisation. This study points to differences in variability that may be more difficult to detect than differences in acoustic properties per se, and suggests that the effects of urbanisation on animal communication can be underestimated if inter-individual variation is not properly addressed.