Overwintering Range Research Articles

Foxtrot migration and dynamic over-wintering range of an Arctic raptor

Advances in tracking technologies have revealed the diverse migration patterns of birds, which are critical for range mapping and population estimation. Population trends are usually estimated in breeding ranges where birds remain stationary, but for species that breed in remote areas like the Arctic, these trends are often assessed in over-wintering ranges. Assessing population trends during the wintering season is challenging due to the extensive movements of birds in these ranges, which requires a deep understanding of the movement dynamics. However, these movements remain understudied, particularly in the mid-latitudes, where many Arctic breeders overwinter, increasing uncertainty in their ranges and numbers. Here, we show that the Arctic breeding raptor Rough-legged buzzard, which overwinters in the mid-latitudes, has a specific wintering strategy. After migrating ca. 1500 km from the Arctic to mid-latitudes, the birds continue to move throughout the entire over-wintering period, traveling another 1000 km southwest and then back northeast as the snowline advances. This continuous movement makes their wintering range dynamic throughout the season. In essence, this movement represents an extension of the quick migration process, albeit at a slower pace, and we have termed this migration pattern ‘foxtrot migration’, drawing an analogy to the alternating fast and slow movements of the foxtrot dance. These results highlight the potential errors in range mapping from single mid-winter surveys and emphasize the importance of this migration pattern in assessing the conservation status of bird species. Understanding this migration pattern could help to correctly estimate bird populations in over-wintering ranges, which is especially important for species that nest in hard-to-reach regions such as the Arctic.

Foxtrot migration and dynamic over-wintering range of an Arctic raptor.

Advances in tracking technologies have revealed the diverse migration patterns of birds, which are critical for range mapping and population estimation. Population trends are usually estimated in breeding ranges where birds remain stationary, but for species that breed in remote areas like the Arctic, these trends are often assessed in over-wintering ranges. Assessing population trends during the wintering season is challenging due to the extensive movements of birds in these ranges, which requires a deep understanding of the movement dynamics. However, these movements remain understudied, particularly in the mid-latitudes, where many Arctic breeders overwinter, increasing uncertainty in their ranges and numbers. Here, we show that the Arctic breeding raptor Rough-legged buzzard, which overwinters in the mid-latitudes, has a specific wintering strategy. After migrating ca. 1500 km from the Arctic to mid-latitudes, the birds continue to move throughout the entire over-wintering period, traveling another 1000 km southwest and then back northeast as the snowline advances. This continuous movement makes their wintering range dynamic throughout the season. In essence, this movement represents an extension of the quick migration process, albeit at a slower pace, and we have termed this migration pattern 'foxtrot migration', drawing an analogy to the alternating fast and slow movements of the foxtrot dance. These results highlight the potential errors in range mapping from single mid-winter surveys and emphasize the importance of this migration pattern in assessing the conservation status of bird species. Understanding this migration pattern could help to correctly estimate bird populations in over-wintering ranges, which is especially important for species that nest in hard-to-reach regions such as the Arctic.

Environmental factors influencing red knot (Calidris canutus islandica) departure times of relocation flights within the non-breeding period.

Deciding when to depart on long-distance, sometimes global, movements can be especially important for flying species. Adverse weather conditions can affect energetic flight costs and navigational ability. While departure timings and conditions have been well-studied for migratory flights to and from the breeding range, few studies have focussed on flights within the non-breeding season. Yet in some cases, overwintering ranges can be large enough that ecological barriers, and a lack of resting sites en route, may resist movement, especially in unfavorable environmental conditions. Understanding the conditions that will enable or prohibit flights within an overwintering range is particularly relevant in light of climate change, whereby increases in extreme weather events may reduce the connectivity of sites. We tracked 495 (n = 251 in 2019; n = 244 in 2020) overwintering red knots (Calidris canutus islandica) in the Dutch Wadden Sea and investigated how many departed towards the UK (on westward relocation flights), which requires flying over the North Sea. For those that departed, we used a resource selection model to determine the effect of environmental conditions on the timing of relocation flights. Specifically, we investigated the effects of wind, rain, atmospheric pressure, cloud cover, and migratory timing relative to sunset and tidal cycle, which have all been shown to be crucial to migratory departure conditions. Approximately 37% (2019) and 36% (2020) of tagged red knots departed on westward relocation flights, indicating differences between individuals' space use within the overwintering range. Red knots selected for departures between 1 and 2.5 h after sunset, approximately 4 h before high tide, with tailwinds and little cloud cover. However, rainfall and changes in atmospheric pressure appear unimportant. Our study reveals environmental conditions that are important for relocation flights across an ecological barrier, indicating potential consequences of climate change on connectivity.

Population Source of Third-Generation Oriental Armyworm in Jilin, China, Determined by Entomology Radar, Trajectory Analysis, and Mitochondrial COI Sequences.

The armyworm, Mythimna separata (Walker) (Lepidoptera: Noctuidae), is an important polyphagous pest with a strong migratory ability. Recently, third-generation larvae have become an increasingly serious pest threat in Jilin Province of northeast China. To investigate the population source of this species, scanning entomological radar observations and insect mitochondrial cytochrome oxidase I (COI) genes were used in this study. Five main results were found: (1) The peak period in captured second-generation moths was from mid to late July. The temperature and wind speeds were optimum for the moths to have migrated. Strong southwesterly winds occurred during the peak migration period. (2) Radar observations indicated that most of the moths’ migration took place at a height of 600 m, often in a dense layer which formed at heights of 350–800 m. (3) Analyses of adult ovarian development and larval haplotypes showed third-generation larvae were progeny of both locally produced progeny and immigrant moths. (4) Based on our back-tracking and haplotype analyses, immigration led to an outbreak originated in the same source area to the southwest. (5) Emigration of second-generation moths was confirmed by both radar observation and mtDNA analysis. Forward trajectories indicated that the moths were capable of immigrating far from their overwintering range. These results are useful for improving the forecasting systems of this insect pest species.

Climate warming promotes pesticide resistance through expanding overwintering range of a global pest

Climate change has the potential to change the distribution of pests globally and their resistance to pesticides, thereby threatening global food security in the 21st century. However, predicting where these changes occur and how they will influence current pest control efforts is a challenge. Using experimentally parameterised and field-tested models, we show that climate change over the past 50 years increased the overwintering range of a global agricultural insect pest, the diamondback moth (Plutella xylostella), by ~2.4 million km2 worldwide. Our analysis of global data sets revealed that pesticide resistance levels are linked to the species’ overwintering range: mean pesticide resistance was 158 times higher in overwintering sites compared to sites with only seasonal occurrence. By facilitating local persistence all year round, climate change can promote and expand pesticide resistance of this destructive species globally. These ecological and evolutionary changes would severely impede effectiveness of current pest control efforts and potentially cause large economic losses.

Genotyping-by-sequencing reveals genomic homogeneity among overwintering Pacific Dunlin (Calidris alpina pacifica) aggregations along the Pacific coast of North America

AbstractInformation on how migratory populations are genetically structured during the overwintering season of the annual cycle can improve our understanding of the strength of migratory connectivity and help identify populations as units for management. Here, we use a genotype-by-sequencing approach to investigate whether population genetic structure exists among overwintering aggregations of the Pacific Dunlin subspecies (Calidris alpina pacifica) sampled at 2 spatial scales (within and among overwintering sites) in the eastern Pacific Flyway. Genome-wide analyses of 874 single nucleotide polymorphisms across 80 sampled individuals revealed no evidence for genetic differentiation among aggregations overwintering at 3 locations within the Fraser River Estuary (FRE) of British Columbia. Similarly, comparisons of aggregations in the FRE and those overwintering in southern sites in California and Mexico indicated no genetic segregation between northern and southern overwintering areas. These results suggest that Pacific Dunlin within the FRE, Sacramento Valley (California), and Guerrero Negro (Mexico) are genetically homogeneous, with no evident genetic structure between sampled sites or regions across the overwintering range. Despite no evidence for differentiation among aggregations, we identified a significant effect of geographical distance between sites on the distribution of individual genotypes in a redundancy analysis. A small proportion of the total genotypic variance (R2 =0.036, P = 0.011) was explained by the combined effect of latitude and longitude, suggesting weak genomic patterns of isolation-by-distance that are consistent with chain-like migratory connectivity between breeding and overwintering areas. Our study represents the first genome-scale investigation of population structure for a Dunlin subspecies and provides essential baseline estimates of genomic diversity and differentiation within the Pacific Dunlin.

An Evaluation of Studies on the Potential Threats Contributing to the Decline of Eastern Migratory North American Monarch Butterflies (Danaus plexippus)

The migratory monarch butterflies (Danaus plexippus) of eastern North America have undergone large-scale declines, which may be attributable to a variety of underlying causes. The uncertainty about the primary cause of declines and whether individual threats are likely to increase in the future presents challenges for developing effective conservation management and policy initiatives that aim to improve population viability. This paper identifies five potential threats and classifies these threats according to the types of studies (observational, experimental, simulation/models) and their current impact and anticipated risk. Broadly, the threats can be classified into five categories: (1) change in suitable abiotic environmental conditions; (2) deforestation in the overwintering range; (3) exposure to contaminants including the bacteria Bacillus thuringiensis, herbicides, and insecticides; (4) loss of breeding habitat; and (5) predation, parasitism, and species-specific pathogens. The vast distribution of the monarch butterfly makes it likely that population declines are attributed to a suite of interacting factors that vary spatially and temporally in their contribution. Nonetheless, the published papers we reviewed suggest the decline in suitable environmental conditions in addition to overwintering (i.e., deforestation) and breeding habitat loss are the most likely threats to continue to affect the population viability of monarch butterflies.

Delimiting Strategic Zones for the Development of Fall Armyworm (Lepidoptera: Noctuidae) on Corn in the State of Florida.

The fall armyworm, Spodoptera frugiperda (J.E. Smith) (Lepidoptera: Noctuidae), cannot survive prolonged periods of freezing temperatures, thereby limiting where it can overwinter in North America. Climate change is anticipated to reduce the frequency of freeze days in Florida over the decades, with the potential consequence of a significant expansion of the overwintering range, whose northern limit in North America was assessed between 27 and 28°N in the last century. To assess this possibility, the development of the fall armyworm on corn leaves, one of the main host plants in the United States, was determined at five constant temperatures ranging from 14 to 30°C. Based on the development time, the thermal constant and the lower threshold temperature were used to estimate the number of generations of fall armyworm at 42 locations in the state of Florida, from 2006 to 2016. Maps were constructed to provide a visual description of the interpolated data, using GIS (Geographic Information System). The highest number of generations was observed in the counties farther south, an area that showed the highest temperatures during the years and plays a strategic role in maintaining fall armyworm populations in corn fields. Additionally, we conclude that in the absence of freeze periods, the northern limit for fall armyworm overwintering should be between 28 and 29°N.

Response of antioxidant enzymes in Mythimna separata (Lepidoptera: Noctuidae) exposed to thermal stress.

The oriental army worm Mythimna separata (Lepidoptera: Noctuidae) is a migratory pest in Eastern Asia and China. Seasonal high temperatures in Southern China and low temperatures in Northern China are pressures favouring the annual migration of this species, while cold tolerance determines the northern limit of its overwintering range. A number of physiological stress responses occur in insects as a result of variations in temperature. One reaction to thermal stress is the generation of reactive oxygen species (ROS), which can be harmful by causing oxidative damage. The time-related effects (durations of 1, 4 and 7 h) of thermal stress treatments of M. separata at comparatively low (5, 10, 15 and 20°C) and high (30, 35, 40 and 45°C) temperatures on the activities of antioxidant enzymes, including superoxide dismutase (SOD), catalase (CAT), peroxidase (POX) and glutathione S-transferases (GSTs), and total antioxidant capacity (T-AOC) were determined. Thermal stress resulted in significant elevation of the activities of SOD, CAT and GSTs, indicating that these enzymes contribute to defence mechanisms counteracting oxidative damage caused by an increase in ROS. However, at high-temperatures, POX and T-AOC were also found to contribute to scavenging ROS. Our results also indicate that extreme temperatures lead to elevated ROS production in M. separata. The present study confirms that thermal stress can be responsible for oxidative damage. To overcome such stress, antioxidant enzymes play key roles in diminishing oxidative damage in M. separata.

Climate Change and Phenology: Empoasca fabae (Hemiptera: Cicadellidae) Migration and Severity of Impact.

Climate change can benefit individual species, but when pest species are enhanced by warmer temperatures agricultural productivity may be placed at greater risk. We analyzed the effects of temperature anomaly on arrival date and infestation severity of potato leafhopper, Empoasca fabae Harris, a classic new world long distance migrant, and a significant pest in several agricultural crops. We compiled E. fabae arrival dates and infestation severity data at different states in USA from existing literature reviews and agricultural extension records from 1951–2012, and examined the influence of temperature anomalies at each target state or overwintering range on the date of arrival and severity of infestation. Average E. fabae arrival date at different states reveal a clear trend along the south-north axis, with earliest arrival closest to the overwintering range. E. fabae arrival has advanced by 10 days over the last 62 years. E. fabae arrived earlier in warmer years in relation to each target state level temperature anomaly (3.0 days / °C increase in temperature anomaly). Increased temperature had a significant and positive effect on the severity of infestation, and arrival date had a marginal negative effect on severity. These relationships suggest that continued warming could advance the time of E. fabae colonization and increase their impact on affected crops.

Tracking overwintering areas of fish-eating birds to identify mercury exposure.

Migration patterns are believed to greatly influence concentrations of contaminants in birds due to accumulation in spatially and temporally distinct ecosystems. Two species of fish-eating birds, the Double-crested Cormorant (Phalacrocorax auritus) and the Caspian Tern (Hydroprogne caspia) breeding in Lake Ontario were chosen to measure the impact of overwintering location on mercury concentrations ([Hg]). We characterized (1) overwintering areas using stable isotopes of hydrogen (δ(2)H) and band recoveries, and (2) overwintering habitats by combining information from stable isotopes of sulfur (δ(34)S), carbon (δ(13)C), nitrogen (δ(15)N), and δ(2)H in feathers grown during the winter. Overall, overwintering location had a significant effect on [Hg]. Both species showed high [Hg] in (13)C-rich habitats. In situ production of Hg (e.g., through sulfate reducing bacteria in sediments) and allochthonous import could explain high [Hg] in birds visiting (13)C-rich habitats. Higher [Hg] were found in birds with high δ(2)H, suggesting that Hg is more bioavailable in southern overwintering locations. Hotspot maps informed that higher [Hg] in birds were found at the limit of their southeastern overwintering range. Mercury concentrations in winter feathers were positively related to predicted spatial pattern of [Hg] in fish using the National Descriptive Model of Mercury in Fish (NDMMF) based on bird spatial assignment (using δ(2)H). This study indicates that the overwintering location greatly influences [Hg].

Movements of boreal caribou in the James Bay lowlands

Little is known about the movements and home range of boreal woodland caribou (Rangifer tarandus) in the James Bay lowlands, northern Ontario. Our preliminary study involves the use of GPS collars with Argos satellite system uplink to monitor movements of caribou and 10 animals were collared in December 2004. Animals appeared to have reduced rates of daily movement starting approximately in mid to late December and stretching until late February. Similarly, most animals appeared to have very reduced rates of movement from the beginning of May to the end of June indicating that this is their calving period (includes both parturition as well as the period immediately after parturition). Thus the over-wintering range was assumed to be where the animals were from mid-December to late February and the calving range was defined as the area they were in from the beginning of May to the end of June. Individual home-ranges were typically large, the mean 90% kernel home range for 2004 - 2007 was 41 579 km2. Over wintering areas and calving areas were small when compared to annual home-range size and reflect the reduced rates of movement during these time periods. Female caribou show site fidelity to calving grounds, using the same core areas year after year. However, the same level of site fidelity was not observed in over-wintering areas. The caribou in the James Bay lowlands display behaviours that are characteristic of both the forest-tundra and forest-forest ecotypes which may warrant the reconsideration of the validity of proposed ecotypes with respect to protection under species-at-risk legislation.

Migratory connectivity and conservation of the Amur Falcon Falco amurensis: a stable isotope perspective

SummaryStable isotopes (δD, δ13C, δ15N) were measured in adult and juvenile Amur Falcon Falco amurensis feathers to understand the migratory connectivity of this species. Using the OIPC (Online Isotopes in Precipitation Calculator) and a calibration curve for American Kestrels Falco sparverius we predicted the breeding range of South African Amur Falcons in the Palaearctic. δD values for juvenile feathers (mean ± SE = -58.1 ± 2.5‰, range -83.9 to -25.7‰) and predicted Palaearctic annual precipitation values indicated that juvenile Amur Falcons in South Africa originated from across their entire Palaearctic range. This rejects the leapfrog migration hypothesis and suggests the widespread movement of birds south, with a funnelling effect into the subregion where they become concentrated over a narrower distribution range. Adult δDf values were more depleted (-37.4 ± 1.8‰, range = -71.3 to -9.3‰) than predicted annual precipitation values for sites where feathers moulted in South Africa (-20.2 ± 0.9‰) but there was no correlation between δDp and δDf. This, together with significant variation of δ13C among sites and annual fluctuations in roost sizes, suggests that roost site fidelity is low in the overwintering range. Populations not confined to breeding sites in South Africa are able to move widely across the subregion, feeding on a broad range of arthropods that become seasonally abundant during the austral summer. Total population estimates for South Africa are significantly lower than global estimates so a significant proportion of the population may not overwinter in South Africa or global populations are significantly lower than estimated. In South Africa, roosts that number hundreds to thousands of individuals are often in large exotic trees, e.g. Eucalyptus sp. They are often tolerant of disturbance at roosts which may be of little concern for their conservation. Habitat changes in the overwintering range in South Africa though, particularly due to agriculture, afforestation and strip mining, may affect global population numbers. There is also concern over the persecution of birds along their migratory route.

Wind Selection and Drift Compensation Optimize Migratory Pathways in a High-Flying Moth

Numerous insect species undertake regular seasonal migrations in order to exploit temporary breeding habitats [1]. These migrations are often achieved by high-altitude windborne movement at night [2-6], facilitating rapid long-distance transport, but seemingly at the cost of frequent displacement in highly disadvantageous directions (the so-called "pied piper" phenomenon [7]). This has lead to uncertainty about the mechanisms migrant insects use to control their migratory directions [8, 9]. Here we show that, far from being at the mercy of the wind, nocturnal moths have unexpectedly complex behavioral mechanisms that guide their migratory flight paths in seasonally-favorable directions. Using entomological radar, we demonstrate that free-flying individuals of the migratory noctuid moth Autographa gamma actively select fast, high-altitude airstreams moving in a direction that is highly beneficial for their autumn migration. They also exhibit common orientation close to the downwind direction, thus maximizing the rectilinear distance traveled. Most unexpectedly, we find that when winds are not closely aligned with the moth's preferred heading (toward the SSW), they compensate for cross-wind drift, thus increasing the probability of reaching their overwintering range. We conclude that nocturnally migrating moths use a compass and an inherited preferred direction to optimize their migratory track.

Estimating the Potato Leafhopper <I>Empoasca fabae</I> (Homoptera: Cicadellidae) Overwintering Range and Spring Premigrant Development by Using Geographic Information System

The potato leafhopper, Empoasca fabae (Harris), is a circular migratory pest of many crops in the United States that overwinters in the southern states. Northward migrant population arrival to the northern states occurs earlier in the north central states compared with northeastern states. Migrant leafhopper arrival to the north varies from year to year depending on factors influencing the development of spring migrants in the overwintering areas and on timing of weather systems capable of transporting the migrants northward. An estimate of the potato leafhopper minimum temperature survival, the geographic limits of the potato leafhopper overwintering range, leafhopper spring development in the overwintering areas, and the identification of the spring migration initiation northwards can help to predict the leafhopper arrival time in the northern states. In the current study, geographic information system (GIS) was used to estimate the potato leafhopper minimum temperature survival and premigrant development. The minimum winter temperature was estimated by overlaying minimum temperature isolines with potato leafhopper collection data taken during the winter, The geographic limits of the overwintering range were estimated using the minimum temperature survival to create a condition-based model by using ArcMap-GIS 8.2. The estimated overwintering range was larger and covered areas further north than previously estimated and included Missouri, Kansas, Kentucky, Virginia, and Maryland. The use of degree-day accumulation to estimate days of first adult emergence in the overwintering areas resulted in earliest adult emergence in the south central region. First adult emergence in south central and southeastern areas occurred before the detection of potato leafhoppers in the north central United States. These data suggested that the difference in population arrival between the north central states and the northeastern states was more dependent on factors affecting the migration and weather conditions encountered along the migration pathway.

Can greenhouses eliminate the development of cold resistance of the leafminers?

Latitudinal patterns for quantitative traits in insect are commonly used to investigate climatic adaptation. We compare the cold resistance of the leafminer (Liriomyza sativae) pupa among populations distributed from tropical to temperate regions, incorporating the thermal overwintering limit of the insect's range. The patterns of cold resistance for northern and southern populations differ. The southern populations significantly increased their cold resistance with latitude, showing a latitudinal pattern independent of seasons, acclimation regimes, and assay methods. In contrast, the northern populations showed no stable patterns; they were always intermediate in cold hardiness between the low-latitude and high-latitude populations within the overwintering limit. Integration of these data with those of the biologically similar congeneric leafminer, L. huidobrensis, suggests that a pattern shift in stress tolerance associated with the overwintering range limit is probably a general adaptive strategy adopted by freeze-intolerant species that have a high-latitude boundary of distribution, but can only overwinter and develop in protected greenhouses in harsh seasons. Considering the widespread availability of greenhouses for overwintering insects in northern China, we speculated that the large-scale existence of thermally-buffered microhabitats in greenhouses might eliminate the development of cold resistance of the leafminer populations. However, results suggest a strong selection for increased cold resistance for natural populations of Liriomyza species at higher latitudes that can overwinter in the field, but not for populations at latitudes above the thermal limit. Thus, habitat modification associated with greenhouses can limit gene flow and reduce cold tolerances even at latitudes above where the leafminers can overwinter in the field.

Implication of pupal cold tolerance for the northern over-wintering range limit of the leafminer Liriomyza sativae (Diptera: Agromyzidae) in China

This study investigated the cold tolerance of a laboratory-reared population and latitude-separated populations of the leafminer Liriomyza sativae in China in terms of low-temperature survival rate and supercooling capacity. Pupae of the laboratory-reared population are susceptible to freezing temperatures. The supercooling point of the pupae varied at a maximum of up to 10°C among the geographic populations. Both acclimation of the pupae at 5 and 10°C significantly increased the cold survival rate of laboratory-reared and field-collected populations. But the field population had a more active response to the same acclimation regime. Combining the experimental data with previous field investigations, the −2°C isotherm of the minimum mean temperature of January was proposed as the leafminer's over-wintering range limit. Meanwhile, with the widespread availability of greenhouses as a source of re-infestation in northern China, the leafminer's natural border of distribution should be determined by the minimum temperature in warm seasons. Our results suggest that biologically similar Liriomyza species may cope with the intensifying cold stress along the latitude by adopting a mixed cold-tolerance strategy, which is closely associated with the greenhouse microhabitats. The physiologically based modeling of the over-wintering limit provides a tool for guiding the management for greenhouse pests and predicting the source of pest infestation.

Inability of Ceratitis capitata (Diptera: Tephritidae) to overwinter in the Judean hills.

The overwintering potential of the Mediterranean fruit fly, Ceratitis capitata (Wiedemann) (Diptera: Tephritidae), in cold winter areas within its northern distribution is a key element in understanding its ecology. Recent studies have suggested that although originating in tropical Africa, the fly has become adapted to the cold weather that prevails within its northernmost areas of distribution. We address the question of whether the Mediterranean fruit fly has expanded its overwintering range to include the mountains of central Israel. Doing so would imply that the fly has developed either a behavioral or a physiological mechanism to cope with low temperature and/or damp conditions in combination with cold. We monitored adult populations year round, sampling fruit, calculating expected emergence days for overwintering flies, and studying adults captured within dense and sparse apple orchards. We also performed several manipulative experiments to study preimago ability to survive the winter under natural or seminatural conditions. The study was conducted in the central mountains of Israel at 700-m altitude from 1994 to 2003. Comparison experiments also were conducted at 400 m and at sea level. Our results show 1) no adults captured during the winter and spring, 2) an absence of new infestations during the winter and spring, and 3) inability of preimago stages to overwinter in the central mountains of Israel. Thus, we conclude that the fly does not overwinter in the central mountains of Israel. We discuss the ecological and applied significance of our findings.