Pine Siskins Research Articles

A North American climate-masting-irruption teleconnection and its change under future climate

Atmospheric variability can impact biological populations by triggering facultative migrations, but the stability of these atmosphere-biosphere connections may be vulnerable to climate change. As an example, we consider the leading mode of continental-scale facultative migration of Pine Siskins, where the associated ecological mechanism is changes in resource availability, with a mechanistic pathway of climate conditions affecting mast seeding patterns in trees which in turn drive bird migration. The three summers prior to pine siskin irruption feature an alternating west-east mast-seeding dipole in conifer trees with opposite anomalies over western and eastern North America. The climate driver of this west-east mast-seeding dipole, referred to as the North American Dipole, occurs during summer in the historical record, but shifts to spring in response to future climate warming during this century in a majority of global climate models. Identification of future changes in the timing of the climate driver of boreal forest mast seeding have broadly important implications for the dynamics of forest ecosystems.

Response to food restriction, but not social information use, varies seasonally in captive cardueline finches.

For songbirds, temperate winters can impose severe conditions on songbirds that threaten survival, including shorter days and often lower temperature and food availability. One well-studied mechanism by which songbirds cope with such conditions is seasonal acclimatization of thermal metabolic traits, with strong evidence for both preparative and responsive changes in thermogenic capacity (i.e., the ability to generate heat) to low winter temperature. However, a bird's ability to cope with seasonal extremes or unpredictable events is likely dependent on a combination of behavioral and physiological traits that function to maintain allostatic balance. The ability to cope with reduced food availability may be an important component of organismal response to temperate winters in songbirds. Here we compare responses to experimentally reduced food availability at different times of year in captive red crossbills (Loxia curvirostra) and pine siskins (Spinus pinus) - two species that cope with variable food resources and live in cold places - to investigate seasonal changes in the organismal response to food availability. Further, red crossbills are known to use social information to improve response to reduced food availability, so we also examine whether use of social information in this context varies seasonally in this species. We find that pine siskins and red crossbills lose less body mass during time-restricted feedings in late winter compared to summer, and that red crossbills further benefit from social information gathered from observing other food restricted red crossbills in both seasons. Observed changes in body mass were only partially explained by seasonal differences in food intake. Our results demonstrate seasonal acclimation to food stress and social information use across seasons in a controlled captive environment and highlight the importance of considering diverse physiological systems (e.g., thermogenic, metabolic, digestive, etc) to understand organismal responses to environmental challenges.

Food and social cues modulate reproductive development but not migratory behavior in a nomadic songbird, the Pine Siskin

Abstract Many animals rely on photoperiodic and non-photoperiodic environmental cues to gather information and appropriately time life-history stages across the annual cycle, such as reproduction, molt, and migration. Here, we experimentally demonstrate that the reproductive physiology, but not migratory behavior, of captive Pine Siskins (Spinus pinus) responds to both food and social cues during the spring migratory-breeding period. Pine Siskins are a nomadic finch with a highly flexible breeding schedule and, in the spring, free-living Pine Siskins can wander large geographic areas and opportunistically breed. To understand the importance of non-photoperiodic cues to the migratory-breeding transition, we maintained individually housed birds on either a standard or enriched diet in the presence of group-housed heterospecifics or conspecifics experiencing either the standard or enriched diet type. We measured body condition and reproductive development of all Pine Siskins and, among individually housed Pine Siskins, quantified nocturnal migratory restlessness. In group-housed birds, the enriched diet caused increases in body condition and, among females, promoted reproductive development. Among individually housed birds, female reproductive development differed between treatment groups, whereas male reproductive development did not. Specifically, individually housed females showed greater reproductive development when presented with conspecifics compared to heterospecifics. The highest rate of female reproductive development, however, was observed among individually housed females provided the enriched diet and maintained with group-housed conspecifics on an enriched diet. Changes in nocturnal migratory restlessness did not vary by treatment group or sex. By manipulating both the physical and social environment, this study demonstrates how multiple environmental cues can affect the timing of transitions between life-history stages with differential responses between sexes and between migratory and reproductive systems.

Research on Factors Influence the Migration of Pine Siskins

Avian migration is a remarkable phenomenon, and the migration behavior of Pine Siskins serves as a fascinating example, offering insights into the complex interplay of external and internal factors that shape migratory patterns. The migration behavior of Pine Siskins is influenced by various factors, including food availability, climatic influences, intrinsic regulation, and social interactions. This facultative migratory species exhibits year-to-year variations in migration patterns, with movements driven by the availability of food resources. Seasonal expressions, such as quiescence and early morning activity, contribute to their migratory behavior. Food availability plays a critical role in their decision to migrate, while climatic dipoles shape their movement patterns. Social interactions also influence their settlement decisions. Physiological regulation involves endocrine signaling components, although differences between pre-migratory and migratory periods were not observed. Biological rhythms, such as photoperiod, contribute to the transition to a migratory state. Understanding these multifaceted aspects of Pine Siskins' migration enhances the knowledge of avian migration and provides insights into the adaptive strategies employed by migratory birds. Further research is needed to fully comprehend the complexities of migratory behavior and develop effective conservation measures for these remarkable avian travelers.

Body mass and triglycerides predict departure of free‐living nomadic pine siskins

Abstract Facultative migrations are observed across vertebrate taxa, include irruptive and nomadic movements, and occur in response to ephemeral and unpredictably variable resources. While the physiology underlying seasonal, obligate migrations is thoroughly studied, much less is known about the physiological mechanisms of facultative movements. We test two hypotheses in a free‐living, nomadic bird, the pine siskin (Spinus pinus). The Prepare Hypothesis predicts that, like obligate migrants, siskins increase fuel stores to prepare for migratory movements and elevations of baseline corticosterone (CORT) support departure. The Escape Hypothesis predicts that siskins do not prepare for departure, body condition declines as food availability declines, and stress‐related levels of CORT induce escape from resource‐poor areas. Under the controlled lab conditions of previous studies, food restriction induces declines in body condition and increases in CORT and locomotor activity, supporting the Escape Hypothesis. This study evaluates the ecological relevance of these captive findings by testing the Prepare and Escape Hypotheses in the field for the first time. During two fall field seasons, we radio‐tagged siskins and tracked their local movements using handheld and automated telemetry. We assess how body condition and CORT relate to feeding behaviour [estimated via plasma triglycerides (TRIG)], space use and departure. We do not find support for either the Prepare or Escape Hypothesis, but rather observe an intermediate pattern. Birds with higher TRIG, and therefore greater food intake, are more likely to depart. Birds in poor condition stay longer near the field site; however, above a threshold body mass, body condition does not predict departure. These findings suggest moderate energy stores are necessary for departure, but movement decisions depend on other factors among birds with sufficient fuel. Siskin movements are physiologically distinct from both obligate and fugitive movements, and we discuss how food availability and body condition interact to drive different types of movement. Read the free Plain Language Summary for this article on the Journal blog.

Telomere length predicts timing and intensity of migratory behaviour in a nomadic songbird.

Our understanding of state-dependent behaviour is reliant on identifying physiological indicators of condition. Telomeres are of growing interest for understanding behaviour as they capture differences in biological state and residual lifespan. To understand the significance of variable telomere lengths for behaviour and test two hypotheses describing the relationship between telomeres and behaviour (i.e. the causation and the selective adoption hypotheses), we assessed if telomere lengths are longitudinally repeatable traits related to spring migratory behaviour in captive pine siskins (Spinus pinus). Pine siskins are nomadic songbirds that exhibit highly flexible, facultative migrations, including a period of spring nomadism. Captive individuals exhibit extensive variation in spring migratory restlessness and are an excellent system for mechanistic studies of migratory behaviour. Telomere lengths were found to be significantly repeatable (R = 0.51) over four months, and shorter pre-migratory telomeres were associated with earlier and more intense expression of spring nocturnal migratory restlessness. Telomere dynamics did not vary with migratory behaviour. Our results describe the relationship between telomere length and migratory behaviour and provide support for the selective adoption hypothesis. More broadly, we provide a novel perspective on the significance of variable telomere lengths for animal behaviour and the timing of annual cycle events.

Social environment influences termination of nomadic migration.

The final stage of migration, when animals terminate migratory movements and transition to a more sedentary state, remains the least understood phase of migration. Whereas migrants that return to the same locations each year may use mechanisms associated with locating a specific destination, migrants with low site fidelity, such as nomadic migrants, may rely on local environmental cues to determine when to cease migratory movements. Using an experiment with captive birds, we tested whether the presence of a conspecific influences the termination of migration, indicated by changes in behaviour and physiology, in a nomadic migrant (the pine siskin, Spinus pinus). We paired migratory birds with a non-migratory individual or left migratory and non-migratory individuals unpaired. Migratory paired birds had a significant decline in nocturnal activity immediately after pairing and activity levels remained lower two weeks later, with significant declines in energetic reserves and flight muscle size also observed. By contrast, migratory unpaired birds maintained high levels of activity and energetic reserves. These results provide evidence for a role of the social environment in migratory termination decisions. Social cues may be particularly useful in nomadic migrants, such as pine siskins, to facilitate settling in high quality, but unfamiliar, habitats.

Seasonal Patterns of Fat Deposits in Relation to Migratory Strategy in Facultative Migrants

Physiological preparations for migration generally reflect migratory strategy. Migrant birds fuel long-distance flight primarily with lipids, but carrying excess fuel is costly; thus, the amount of fat deposited prior to departure often reflects the anticipated flight duration or distance between refueling bouts. Seasonal pre-migratory deposition of fat is well documented in regular seasonal migrants, but is less described for more facultative species. We analyze fat deposits of free-living birds across several taxa of facultative migrants in the songbird subfamily Carduelinae, including house finches (Haemorhous mexicanus), American goldfinches (Spinus tristis), pine siskins (Spinus pinus) and four different North American ecotypes of red crossbills (Loxia curvirostra), to evaluate seasonal fat deposition during facultative migratory periods. Our data suggest that the extent of seasonal fat deposits corresponds with migratory tendency in these facultative taxa. Specifically, nomadic red crossbills with a seasonally predictable annual movement demonstrated relatively large seasonal fat deposits coincident with the migratory periods. In contrast, pine siskins, thought to be more variable in timing and initiation of nomadic movements, had smaller peaks in fat deposits during the migratory season, and the partial migrant American goldfinch and the resident house finch showed no peaks coincident with migratory periods. Within the red crossbills, those ecotypes that are closely associated with pine habitats showed larger peaks in fat deposits coincident with autumn migratory periods and had higher wing loading, whereas those ecotypes associated with spruces, Douglas-fir and hemlocks showed larger peaks coincident with spring migratory periods and lower wing loading. We conclude that population averages of fat deposits do reflect facultative migration strategies in these species, as well as the winter thermogenic challenges at the study locations. A difference in seasonal fattening and wing loading among red crossbill ecotypes is consistent with the possibility that they differ in their migratory biology, and we discuss these differences in light of crossbill reproductive schedules and phenologies of different conifer species.

Migratory state and patterns of steroid hormone regulation in the pectoralis muscle of a nomadic migrant, the pine siskin (Spinus pinus)

The endocrine system is known to mediate responses to environmental change and transitions between different life stages (e.g., a non-breeding to a breeding life stage). Previous works from the field of environmental endocrinology have primarily focused on changes in circulating hormones, but a comprehensive understanding of endocrine signaling pathways requires studying changes in additional endocrine components (e.g., receptor densities) in a diversity of contexts and life stages. Migratory birds, for instance, can exhibit dramatic changes in their physiology and behavior, and both sex steroids as well as glucocorticoids are proposed mediators of the transition into a migratory state. However, the role of changes in endocrine signaling components within integral target tissues, such as flight muscles, in modulating the transition into a migratory state remains poorly understood. Here, we examined changes in gene expression levels of and correlational patterns (i.e., integration) between 8 endocrine signaling components associated with either glucocorticoids or sex steroid signaling in the pectoralis muscles of a nomadic migratory bird, the pine siskin (Spinus pinus). The pectoralis muscle is essential to migratory flight and undergoes conspicuous changes in preparation for migration, including hypertrophy. We focus on endocrine receptors and enzymes (e.g., 5α-reductase) that modulate the signaling capacity of circulating hormones within target tissues and may influence either catabolic or anabolic functioning within the pectoralis. Endocrine signaling components were compared between captive birds sampled prior to the expression of vernal migratory preparation and during the expression of a vernal migratory state. While birds exhibited differences in the size and color of the flight muscle and behavioral shifts indicative of a migratory state (i.e., zugunruhe), none of the measured endocrine components differed before and after the transition into the migratory state. Patterns of integration amongst all genes did, however, differ between the two life stages, suggesting the contrasting demands of different life stages may shape entire endocrine signaling networks within target tissues rather than individual components. Our work aligns with previous endocrine studies on pine siskins and, viewed together, suggest additional studies are needed to understand the endocrine system’s role in mediating the development and progression of the vernal migratory state in this species. Further, the patterns observed in pine siskins, a nomadic migrant, differ from previous studies on obligate migrants and suggest that different mechanisms or interactions between endocrine signaling components may mediate the migratory transition in nomadic migrants.

Physiology and behavior under food limitation support an escape, not preparative, response in the nomadic pine siskin (Spinus pinus).

Migration allows animals to use resources that are variable in time and/or space, with different migratory strategies depending on the predictability of resource variation. When food varies seasonally, obligate migrants anticipate and prepare for migration. In contrast, facultative migrants, whose movements are unpredictable in timing and destination, may prepare for either migration or escape when resources are depleted. We propose and test two alternative hypotheses regarding the behavioral and physiological responses of facultative migrants to declining food availability. (1) The prepare hypothesis predicts that facultative migrants prepare for departure by increasing fuel stores in response to declining food availability, and elevations of baseline corticosterone (CORT) facilitate increased activity. (2) The escape hypothesis predicts that facultative migrants do not prepare for departure, body condition declines as food availability declines, and stress-induced levels of CORT induce escape behavior when both energetic condition and food resources are low. We conducted a 16-day experiment, measuring body composition (using quantitative magnetic resonance), activity (using force perches) and baseline CORT in pine siskins (Spinus pinus) given ad libitum food or a slow decline, fast decline or randomly changing amount of food. Our results support the escape hypothesis: body condition declined as food declined, decreases in body and fat mass were associated with increases in baseline CORT, and activity increased only when food availability was low. This work suggests that facultative migration in autumn allows birds to escape low-resource areas and that the underlying physiological mechanisms differ from those driving both seasonal, obligate migrations and spring nomadic movements.

Variation in chronotype is associated with migratory timing in a songbird.

Like many organisms, birds exhibit daily (circadian) and seasonal biological rhythms, and within populations both daily and seasonal timing often vary among individuals. Because photoperiod interacts with the circadian rhythms of many organisms to induce seasonal changes in behaviour and physiology, it is hypothesized that differences in daily timing, called chronotypes, underpin differences among individuals in the timing of seasonal events. For seasonal events stimulated by increasing daylength, this hypothesis predicts a positive relationship between the timing of daily and seasonal activities of individuals, with advanced chronotypes expressing events earlier in the year. The few previous tests of this hypothesis have focused on seasonal reproductive timing in birds. However, the hypothesis predicts that this relationship should extend to other photoinduced seasonal events. Therefore, we tested whether variation in chronotype was associated with variation in spring migratory timing in a captive songbird model, the pine siskin (Spinus pinus). We found that pine siskins expressing migratory restlessness exhibited repeatable chronotypes in their timing of nocturnal activity. Further, chronotype was significantly associated with the onset date of migratory behaviour, consistent with the hypothesized relationship between chronotype and seasonal timing.

Declining food availability, corticosterone, and migratory response in a nomadic, irruptive migrant

While obligate migrants time their movements to respond to predictable changes in the environment, facultative migration is characterized by more variable movements that are driven by unpredictable changes in resource availability. The proximate cues that trigger facultative migrations and the endocrine mechanisms involved in these responses remain poorly understood, though corticosterone may be a key mediator of facultative migration due to its effects on activity and metabolic processes. We conducted experiments in the fall and spring to examine the response of pine siskins (Spinus pinus), a facultative migrant, to a two-stage food restriction. Our goals were to examine whether declining food availability stimulated behavioral and/or physiological changes consistent with a migratory response, whether anticipatory behavioral and physiological adjustments occurred when birds were initially presented with changing food availability, and if observed changes corresponded to changes in circulating corticosterone levels. We found no evidence of preparatory physiological changes for migration, but food-restricted birds in the spring had increased daytime activity indicative of a migratory response. Corticosterone increased at each stage of the restriction and the change in corticosterone corresponded to the magnitude of decline in body condition. Increased corticosterone was also correlated with larger increases in activity during the initial stage of food restriction, but only during the spring, when birds also displayed higher levels of initial body condition. These results suggest that initial energetic state and corticosterone response may interact to determine an individual's behavioral and physiological response to declining food availability and ultimately the facultative migratory response.

Migratory state is not associated with differences in neural glucocorticoid or mineralocorticoid receptor expression in pine siskins

Abstract Although the endocrine system likely plays an important role in orchestrating the transition to a migratory state, the specific mechanisms by which this occurs remain poorly understood. Changes in glucocorticoid signaling are one proposed mechanism that may be important in migratory transitions. Although previous work has focused on the role of changes in circulating glucocorticoids, another potential mechanism is changes in the expression of its cognate receptors. Here, we test this hypothesis by comparing mRNA expression of the genes for the mineralocorticoid receptor (MR) and glucocorticoid receptor (GR) in two brain regions implicated in the regulation of migratory behavior (the hippocampus and hypothalamus) in pine siskins (Spinus pinus) sampled before or after the transition to a spring nomadic migratory state. Compared to pre-migratory birds, migratory birds had body conditions more indicative of physiological preparations for migration (e.g., larger body mass), and greater levels of nocturnal migratory restlessness. However, we found no differences between pre-migratory and migratory birds in the expression of GR or MR mRNA in either the hippocampus or hypothalamus. Thus, differences in expression of receptors for glucocorticoids do not appear to underly the observed differences in physiology and behavior across a migratory transition. Taken together with previous results showing no change in circulating corticosterone levels during this transition, our findings provide no evidence for a role of glucocorticoid signaling in the spring migratory transition of this species.

Increasing photoperiod stimulates the initiation of spring migratory behaviour and physiology in a facultative migrant, the pine siskin.

The transition to a migratory state involves coordinated changes in physiology and behaviour. In species with regular, predictable (obligate) migrations, increasing day length triggers the expression of a spring migratory state and androgens play an important role in stimulating its development. By contrast, we know little about the environmental cues and endocrine mechanisms that regulate migration in species with less predictable (facultative) migrations. Here, we tested whether photoperiod stimulates a migratory state in a facultative nomadic migrant, the pine siskin (Spinus pinus). We exposed wintering birds to either a naturally increasing or short-day photoperiod and measured physiological and behavioural changes indicative of a migratory state. We also examined changes in circulating hormones that may play a role in the migratory transition. Natural-day, but not short-day, birds displayed physiological preparations for migration, including increases in fat deposition, and showed increased levels of migratory restlessness. We found no evidence for a role of corticosterone in the migratory transition, but testosterone may be important. This study is the first experimental test of the role of photoperiod in regulating facultative migration and demonstrates that the predictive cue used by many obligate migrants to time spring migration is also important in a facultative migrant.

Incursion of Post-Breeding Pine Siskins in the Dune- Ridge Forest, Delta Marsh, Manitoba, 1985

Incursion of Post-Breeding Pine Siskins in the Dune- Ridge Forest, Delta Marsh, Manitoba, 1985

Major Histocompatibility Complex Allele Persistence in Eurasia and America in the Genus Carduelis (Spinus) During Million Years

Introduction:GenusCarduelis(Fringillidaefamily) includes goldfinches, siskins, redpolls, greenfinches and crossbills. Many of the species classified within this genus and other related genera have been grouped by using molecular systematics and the mitochondrial cytochrome b (mt cyt b) gene. According to this, the Eurasian siskin (C. spinus)is the only one extant direct ancestor of several North American finches; North American / South American radiations may have been originated by Eurasian siskin (or extinct relative). In the present work, we aim to perform a study of transpecies and transcontinental analyses of MHC (Major Histocompatibility Complex) Class I alleles in several genusCarduelis/Spinusspecies in order to draw evolutionary conclusions in several wild bird species belonging to the genusCarduelis / Spinus.Materials and Methods:Blood was taken from worldwide wild bird species. Passerine phylogeny was done after analysing mtDNA with Maximun Likelihood and Bayesian dendrograms. Major histocompatibility complex alleles were obtained by standard DNA cloning and sequencing.Results:We found two matches between MHC-I DNA alleles from different South American siskins at DNA level. Also, it was observed that the Eurasian siskin shares a protein with pine siskin and another with three South American siskins. Eight South American siskins species also share the same MHC protein. In addition, studied songbirds MHC class I intron 2 is longer than that ofGallus gallus.Conclusion:We have drawn the following conclusions: 1) We present the first direct evidence that “Minimal Essential MHC” does not exist for birds; one of its main definition characters,i.e.: small intron size does not hold for songbirds. 2) We also report that MHC genes transpecies evolution exist in birds by showing also for the first time that worldwide bird species keep the same MHC protein and DNA alleles. 3) New evidences on MHC alleles conservation from EurasianCarduelis spinus(most ancient) to South American siskins (most recent) during million years support that Eurasian siskin is the parental species for American GenusCarduelis (Spinus)species. It is uncertain whether Eurasian siskin (or extant relative) had initially an Holoartic distribution, including America.

Passage Dates, Energetic Condition, and Age Distribution of Irruptive Pine Siskins during Autumn Stopovers at a Reclaimed Landfill in the New Jersey Meadowlands

Abstract Little is known about the stopover biology of Spinus pinus (Pine Siskin) and other Fringillid birds during their irruptive movements into the US from boreal Canada. Here, we report on the passage timing, energetic condition, and age distribution of 402 Pine Siskins that we captured during autumn stopovers in New Jersey in the irruption year of 2012. Pine Siskins passed through our study site for ∼3 weeks and peaked in abundance between 9 and 12 October. More birds were juveniles than adults (54% v. 46%), although the difference was not significant. Juveniles were heavier than adults, but fat scores did not differ by age. Neither age group appeared to gain significant mass during the stopover. We encourage migration banding stations like ours that experience irruptions to report the information they collect and help improve our understanding of the migration biology and behavior of irruptive species.

Estrogen levels influence medullary bone quantity and density in female house finches and pine siskins

Medullary bone, a non-structural osseous tissue, serves as a temporary storage site for calcium that is needed for eggshell production in a number of avian species. Previous research focusing primarily on domesticated species belonging to the Anseriformes, Galliformes, and Columbiformes has indicated that rising estrogen levels are a key signal stimulating medullary bone formation; Passeriformes (which constitute over half of extant bird species and are generally small) have received little attention. In the current study, we examined the influence of estrogen on medullary bone and cortical bone in two species of Passeriformes: the Pine Siskin (Spinus pinus) and the House Finch (Haemorhous mexicanus). Females of these species received either an estradiol implant or were untreated as a control. After 4.5–5months, reproductive condition was assessed and leg (femora) and wing (humeri) bones were collected for analysis using high-resolution (10μm) micro-computed tomography scanning. We found that in both species estradiol-treated females had significantly greater medullary bone quantity in comparison to untreated females, but we found no differences in cortical bone quantity or microarchitecture. We were also able to examine medullary bone density in the pine siskins and found that estradiol treatment significantly increased medullary bone density. Furthermore, beyond the effect of the estradiol treatment, we observed a relationship between medullary bone quantity and ovarian condition that suggests that the timing of medullary bone formation may be related to the onset of yolk deposition in these species. Further research is needed to better understand the precise timing and endocrine regulation of medullary bone formation in Passerines and to determine the extent to which female Passerines rely on medullary bone calcium during the formation of calcified eggshells.

Seasonal expression of migratory behavior in a facultative migrant, the pine siskin

The vast majority of what is known about migration comes from studies of obligate migration—regular and predictable “to-and-fro” seasonal movements. Much less is known about facultative migration, which occurs less predictably in space and time. Here, we use pine siskins (Spinus pinus), nomadic and irruptive finches, as a captive model to advance our understanding of facultative migration. Building on approaches used to study obligate migration, we investigate whether pine siskins exhibit migratory restlessness in captivity, similar to obligate migrants, and whether pine siskins express a seasonal pattern of migratory behavior. Pine siskins exhibited nocturnal migratory restlessness that was remarkably similar to that in obligate migrants; it was characterized by jumping and wing whirring behavior that was preceded by a period of quiescence. This behavior was expressed seasonally, peaking in the spring, when free-living pine siskins frequently migrate. As in many obligate migrants, migratory restlessness in pine siskins coincided with elevated body condition and showed minimal overlap with body molt. Although pine siskins are both nocturnal and diurnal migrants, patterns of diurnal activity did not mirror patterns of nocturnal activity and were more difficult to interpret. Free-living pine siskins frequently migrate in the fall, but in captivity showed no clear pattern of fall migratory restlessness. We suggest that this could reflect a relatively predictable and programmed spring migration and a more facultative fall migration. Migrations are undertaken by a diverse array of animals and are important to the functioning of many ecosystems. The goal of this study was to shed light on an important, but understudied form of migration: facultative migration. Facultative migrations occur less predictably than the better-studied “to-and-fro” seasonal migrations of obligate migrants. Using a songbird, the pine siskin, as a model, we sought to determine if pine siskins express a migratory state in captivity and whether pine siskins exhibit a seasonal pattern of migratory behavior. We found that pine siskins showed behavioral and physiological changes associated with a migratory state, which were remarkably similar to those in obligate migrants. Pine siskins showed seasonal changes in migratory behavior in the study that suggest that this species has a predictable and programmed spring migratory period and a much more flexible fall migration.

Structure and variation of the Fringillidae (Aves: Passeriformes) mitochondrial DNA control region and their phylogenetic relationship

The control region is the major noncoding segment of animal mitochondrial DNA. To infer the structure and variation of Fringillidae mitochondrial DNA control region, the entire control region sequences of 25 species were analyzed. The length of the control region sequences was very conserved (1230 ± 18) and can be separated into three domains. The frequency of both substitutions and gaps was highest in the third domain and lowest in the central region. The control region has the same flanking gene order from tRNAGlu to tRNAPhe. Genetic distances between species ranged from 1.80% (between Carduelis pinus and Carduelis spinus) to 25.34% (between Carduelis sinica and Coccothraustes vespertinus). The average genetic distances among the species within the genera varied from 5.11% (Leucosticte) to 14.31% (Carpodacus). The average genetic distances showed insignificantly negative correlation with ts/tv. Domain III is the most variable of the three domains among all the genera. The control region of Fringillidae contains a putative TAS element and the highly conserved CSB-1, and F, E, D, C boxes. However, neither CSB-2 nor CSB-3 could be unambiguously identified in the Fringillidae. The maximum likelihood method was used to construct a phylogenetic tree. Control region analysis demonstrated that some currently recognized genera may be polyphyletic, including Carpodacus, Carduelis and Serinus.