The response of Mitchell grasses (Astrebla spp.) and Button grass (Dactyloctenium radulans (R. Br.)) to rainfall and their importance to the survival of the Australian plague locust,Chortoicetes terminifera (Walker), in the arid zone

The environmental conditions that allow the growth of Barley Mitchell grass, Astrebla pectinata, and the development of the Australian plague locust, Chortoicetes terminifera, are similar. A single fall of more than 20 mm rain and a mean monthly maximum temperature above 23°C ensures sustained growth of Barley Mitchell grass and hatching of eggs of the Australian plague locust. Curly Mitchell grass, Astrebla lappacea, is found in heavier clay soils and requires about 40 mm to respond. Both Mitchell grasses normally remain green for about 2 months after rain which is slightly longer than the duration of locust development to the adult stage. As a result, locusts in Mitchell grass areas can complete their development to migration and laying on a single substantial fall of rain: an adaptation of critical importance in the arid zone where follow-up rain is unlikely. Under similar circumstances, ephemerals like Button grass, Dactyloctenium radulans, are dry in approximately 6 weeks. The locusts are usually at the late nymphal stage by this time and even though Mitchell grass is still quite green, nymphal growth is retarded, resulting in adults which are smaller than normal. These adults are able to accumulate the fat needed for migration by feeding on the dry green Mitchell grasses. In the absence of dry green or green Mitchell grass, the locusts persist locally and die without laying unless there is further rain.     

河北省黄骅市三个重点蝗区两个时段土壤湿度的遥感提取

蝗灾是人类历史上重要的自然灾害之一,影响蝗灾爆发的因素很多,其中土壤湿度是重要的影响因素之一。以位于河北省黄骅市境内的3个重点蝗区——黄灶、杨官庄和藤南大洼为监测样区,利用高时间分辨率MODIS遥感影像数据,分别提取2002年(东亚飞蝗大发生年)秋蝗产卵期到夏蝗孵化期间的土壤湿度信息以及2004年(东亚飞蝗轻为害年)相同时段的土壤湿度信息,发现蝗灾大爆发年份的土壤湿度明显低于轻发生年份,在秋蝗的产卵期(9—10月)和夏蝗的孵化期(3—5月)差异尤为明显。     

Field evaluation of imidacloprid as a systemic approach to flea control in black‐tailed prairie dogs,Cynomys ludovicianus

Epizootic outbreaks of sylvatic plague have dramatically influenced prairie dog (Cynomys sp.) populations across North America. While a great deal of debate surrounds the cause and persistence of plague, flea control can stop the spread of plague epizootic outbreaks and even increase prairie dog survival under non‐epizootic conditions. We investigated a newly‐developed imidacloprid‐treated grain bait that could potentially reduce flea infestations and mitigate the effects of plague on black‐tailed prairie dogs (C. ludovicianus). We used a study design involving randomly assigned experimental and control study plots to assess the effectiveness of the systemic flea control product. We observed a significant difference in flea prevalence and abundance between experimental and control sites on three of the four sites treated with a single application of imidacloprid‐treated grain bait for up to 90 days post‐treatment. We observed an even greater reduction in flea infestations following the double application of treatment bait on two of three additional experimental sites. While we were unable to reduce flea infestations to the extent reported for more commonly used topical insecticides containing deltamethrin, imidacloprid might still be effective at reducing the risk of plague and halting epizootics. In addition, this systemic product can be more rapidly applied than topical insecticides, providing managers with a tool to quickly reduce flea infestations. Future research is needed to evaluate the effectiveness of different application timing and rates, the utility of the product in limiting plague, and the potential effects on non‐target species that might also consume the treated bait.     

Sylvatic plague management and prairie dogs – a meta‐analysis

Yersinia pestis, a bacterial pathogen that causes sylvatic plague, is present in the prairie dogs (Cynomys spp.) of North America. Epizootics of sylvatic plague through transmission in vectors (fleas) commonly completely extirpate colonies of prairie dogs. Wildlife managers employ a wide variety of insecticidal treatments to suppress plague and conserve prairie dog colonies. I compiled and statistically compared the available literature describing methods of plague control and their relative effectiveness in managing plague outbreaks by using meta‐analyses. Natural log response ratios were used to calculate insecticide‐induced vector mortality and vaccine‐conferred survival increases in prairie dogs in 37 publications. Further, subgroupings were used to explore the most effective of the available vector suppression insecticides and plague suppression vaccines. After accounting for the type of treatment used and the method by which it was applied, I observed plague reduction through use of both insecticides and vaccines. Insecticides resulted in a significant reduction of the abundance of vectors by 91.34% compared to non‐treated hosts (p<0.0001). Vaccines improved survival of prairie dog hosts by 4.00% (p<0.0001) compared to control populations. The use of insecticides such as deltamethrin and carbaryl is recommended to stop actively spreading epizootics, and dual antigen oral vaccines to initially suppress outbreaks.     

Hybrid songbirds employ intermediate routes in a migratory divide

Migratory divides are contact zones between populations that use different routes to navigate around unsuitable areas on seasonal migration. Hybrids in divides have been predicted to employ intermediate and potentially inferior routes. We provide the first direct test of this hypothesis, using light‐level geolocators to track birds breeding in a hybrid zone between Swainson's thrushes in western Canada. Compared to parental forms, hybrids exhibited increased variability in their migratory routes, with some using intermediate routes that crossed arid and mountainous regions, and some using the same routes as one parental group on fall migration and the other on spring migration. Hybrids also tended to use geographically intermediate wintering sites. Analysis of genetic variation across the hybrid zone suggests moderately strong selection against hybrids. These results indicate that seasonal migratory behaviour might be a source of selection against hybrids, supporting a possible role for migration in speciation.     

Isolation from a marching band increases haemocyte density in wild locusts (Chortoicetes terminifera)

1. The density‐dependent prophylaxis hypothesis predicts that individuals in high‐density populations will invest more resources in immune defence than individuals at lower densities. 2. However, recent work suggests that this prediction may not apply to all situations; solitarious species may paradoxically have higher scores than crowded counterparts in certain immune assays. 3. To investigate the relationship between a key immune parameter and field population densities, the total haemocyte counts (THCs) of Australian plague locusts (Chortoicetes terminifera) from three population densities in Western Australia were compared. 4. THCs were negatively correlated with field population densities, and locusts removed from a marching band and kept in isolation had increased THCs relative to group‐housed controls. 5. These results demonstrate that immune investment can inversely relate to population density in field conditions. 6. We suggest that isolated locusts increase their haemocyte densities relative to crowded conspecifics in response to potentially greater exposure to parasitoids and nematodes.     

Perceived wintering latitude determines timing of song output in a migratory bird

Migratory bird populations frequently consist of individuals that overwinter variable distances from the breeding site. Seasonal changes in photoperiod, which varies with latitude, underlie seasonal changes in singing frequency in birds. Therefore, migratory populations that consist of individuals that overwinter at different latitudes with large overwintering ranges could experience within‐population variation in seasonal production of song. To test the influence of overwintering latitude on intrapopulation variance in song production in the spring, we subjected two groups of Eastern Song Sparrows (Melospiza melodia melodia) from the same partially migratory breeding population to different photoperiodic schedules associated with a 1,300‐km difference in overwintering location. One group remained on the natural photoperiodic schedule of the breeding site (resident group) while the other group experienced a nonbreeding photoperiod that mimicked a southern migration in the fall followed by a northern migration back to the breeding site in the spring (migratory group). We compared song output between the two groups in three different stages (nonbreeding, prebreeding, and breeding). Little singing occurred during nonbreeding stage sample dates (20 November, 6 December) for the resident group, and no singing occurred for the migrant group. During the prebreeding stage (27 January, 7 February), significantly more singing occurred in the resident group than in the migrant group. During the breeding stage (21 March, 4 April), after a simulated migration for the migrants, song output was similar in both groups. These results suggest that within‐population variation in wintering latitude may contribute to variation in seasonal changes in singing behavior, which may covary with readiness to breed. Studies utilizing confirmed migrants and residents, rather than merely simulated migrants and residents, are also needed to better understand these processes.     

Differential plague susceptibility in species and populations of prairie dogs

Laboratory trials conducted over the past decade at U.S. Geological Survey National Wildlife Health Center indicate that wild populations of prairie dogs (Cynomys spp.) display different degrees of susceptibility to experimental challenge with fully virulent Yersinia pestis, the causative agent of plague. We evaluated patterns in prairie dog susceptibility to plague to determine whether the historical occurrence of plague at location of capture was related to survival times of prairie dogs challenged with Y. pestis. We found that black‐tailed prairie dogs (Cynomys ludovicianus) from South Dakota (captured prior to the detection of plague in the state), Gunnison's prairie dogs (Cynomys gunnisoni) from Colorado, and Utah prairie dogs (Cynomys parvidens) from Utah were most susceptible to plague. Though the susceptibility of black‐tailed prairie dogs in South Dakota compared with western locations supports our hypothesis regarding historical exposure, both Colorado and Utah prairie dogs have a long history of exposure to plague. It is possible that for these populations, genetic isolation/bottle necks have made them more susceptible to plague outbreaks.     

Local factors associated with on‐host flea distributions on prairie dog colonies

Outbreaks of plague, a flea‐vectored bacterial disease, occur periodically in prairie dog populations in the western United States. In order to understand the conditions that are conducive to plague outbreaks and potentially predict spatial and temporal variations in risk, it is important to understand the factors associated with flea abundance and distribution that may lead to plague outbreaks. We collected and identified 20,041 fleas from 6,542 individual prairie dogs of four different species over a 4‐year period along a latitudinal gradient from Texas to Montana. We assessed local climate and other factors associated with flea prevalence and abundance, as well as the incidence of plague outbreaks. Oropsylla hirsuta, a prairie dog specialist flea, and Pulex simulans, a generalist flea species, were the most common fleas found on our pairs. High elevation pairs in Wyoming and Utah had distinct flea communities compared with the rest of the study pairs. The incidence of prairie dogs with Yersinia pestis detections in fleas was low (n = 64 prairie dogs with positive fleas out of 5,024 samples from 4,218 individual prairie dogs). The results of our regression models indicate that many factors are associated with the presence of fleas. In general, flea abundance (number of fleas on hosts) is higher during plague outbreaks, lower when prairie dogs are more abundant, and reaches peak levels when climate and weather variables are at intermediate levels. Changing climate conditions will likely affect aspects of both flea and host communities, including population densities and species composition, which may lead to changes in plague dynamics. Our results support the hypothesis that local conditions, including host, vector, and environmental factors, influence the likelihood of plague outbreaks, and that predicting changes to plague dynamics under climate change scenarios will have to consider both host and vector responses to local factors.     

Isotopic (δ2Hf) evidence of “loop migration” and use of the Gulf of Maine Flyway by both western and eastern breeding populations of Blackpoll Warblers

Declining numbers of Blackpoll Warblers (Setophaga striata) have been documented at long‐term migration monitoring sites as well as in breeding areas. However, the “loop migration” of Blackpoll Warblers makes it difficult to ascribe population change at migration monitoring sites to specific breeding populations. Individuals from all populations across the breeding range of Blackpoll Warblers concentrate in fall along the Atlantic coastline of eastern North America prior to initiating a transoceanic flight to wintering areas. In spring, Blackpoll Warblers return along a different route, moving north into the southeastern United States where birds from eastern and western breeding populations then diverge during migration to reach their respective breeding areas. To monitor breeding populations outside of breeding areas and identify factors potentially affecting those populations, we must be able to identify where birds captured during migration breed and map seasonal variation in population‐specific flyways. To “map” population‐specific migration movements of Blackpoll Warblers, we used feather deuterium (δ²H_f) values and a spatially explicit model to assign molt origins of 289 Blackpoll Warblers moving through sites in the Gulf of Maine (GOM) region and at three locations further west and south (northern Great Lakes area, Pennsylvania, and Florida). The assignment method was validated with feather samples from 35 birds captured during the breeding season at Churchill, Manitoba, Canada. As predicted, the spatial pattern of movement within and between seasons reflected “loop migration.” Blackpoll Warblers captured during fall migration in the GOM region included birds from across their breeding range, whereas birds captured during the spring were exclusively from northeastern populations. During fall migration, Blackpoll Warblers captured at two sites west of the GOM were from breeding areas further northwest than those from western Canada that were captured in the GOM. Blackpoll Warblers captured in eastern Florida during spring migration were assigned exclusively to breeding areas in the northeast, suggesting that eastern and western populations diverge soon after entering the United States. Finally, most Blackpoll Warblers sampled at Manomet Bird Observatory originated from breeding populations in Alaska and western Canada that have shown a similar (70–90%) decline over the same period. Our results, therefore, not only document the “loop migration” pattern of Blackpoll Warblers, but, by mapping patterns of connectivity between breeding and non‐breeding areas, may help target conservation efforts for breeding populations of Blackpoll Warblers where most needed.     

Validation of Inverse Seasonal Peak Mortality in Medieval Plagues,Including the Black Death,in Comparison to Modern Yersinia pestis-Variant Diseases

Background

Recent studies have noted myriad qualitative and quantitative inconsistencies between the medieval Black Death (and subsequent “plagues”) and modern empirical Y. pestis plague data, most of which is derived from the Indian and Chinese plague outbreaks of A.D. 1900±15 years. Previous works have noted apparent differences in seasonal mortality peaks during Black Death outbreaks versus peaks of bubonic and pneumonic plagues attributed to Y. pestis infection, but have not provided spatiotemporal statistical support. Our objective here was to validate individual observations of this seasonal discrepancy in peak mortality between historical epidemics and modern empirical data.

Methodology/Principal Findings

We compiled and aggregated multiple daily, weekly and monthly datasets of both Y. pestis plague epidemics and suspected Black Death epidemics to compare seasonal differences in mortality peaks at a monthly resolution. Statistical and time series analyses of the epidemic data indicate that a seasonal inversion in peak mortality does exist between known Y. pestis plague and suspected Black Death epidemics. We provide possible explanations for this seasonal inversion.

Conclusions/Significance

These results add further evidence of inconsistency between historical plagues, including the Black Death, and our current understanding of Y. pestis-variant disease. We expect that the line of inquiry into the disputed cause of the greatest recorded epidemic will continue to intensify. Given the rapid pace of environmental change in the modern world, it is crucial that we understand past lethal outbreaks as fully as possible in order to prepare for future deadly pandemics.     

The relationship between outbreaks, rainfall and low density populations of the African armyworm, Spodoptera exempta, in Kenya

This study has demonstrated that low numbers of African armyworms (Spodoptera exempta) (Walker) (Lepidoptera: Noctuidae) can be found throughout the year in parts of the coastal and highland regions of Kenya where there is frequent rainfall. During the study period the numbers of moths caught in pheromone traps in these regions built up during the short and long rains, and decreased dramatically during the intervening dry seasons. There was a lag of one to two months between the peaks for rainfall and moth numbers. This contrasts with the situation at sites of the seasonal outbreaks of armyworm larvae, where a sudden preceding influx of moths coincides with the rainfall. A positive correlation was found, for the long rains seasons only, between the number of armyworm outbreaks throughout the country and the peak numbers of moths in coastal and highland regions.It is suggested therefore that the peak numbers of moths trapped in the eastern highlands and coastal regions during the rainy seasons arise principally from outbreaks in the extensive seasonal grasslands. These grassland areas diminish considerably during the prolonged dry periods between the two rainy seasons, and it is only the eastern highland and coastal regions that are likely to provide suitable habitats for breeding during the dry seasons. The significance of these populations for initiating the first outbreaks of the following season is discussed.     

GPS‐tracking reveals non‐breeding locations and apparent molt migration of a Black‐headed Grosbeak

Black‐headed Grosbeaks (Pheucticus melanocephalus) have been observed to undergo prebasic molt during fall in the North American Monsoon region of the southwestern United States and northwestern Mexico, but it is unknown whether molt migration is pervasive across populations of the species. During the 2014 breeding season, we GPS‐tagged (where GPS is global positioning system) nine adult Black‐headed Grosbeaks in Yosemite National Park with archival GPS tags to determine specific locations where grosbeaks breeding in Yosemite spent portions of the non‐breeding season, and to assess whether those locations were consistent with molt migration. On 2 June 2015, one of these birds, a male GPS‐tagged on 19 June 2014, was recaptured with its GPS unit still attached. Data downloaded from the unit revealed that, by 20 August 2014, the bird had moved 1300 km from Yosemite National Park to Sonora, Mexico, where it remained until at least 15 October 2014. By 24 November 2014, the grosbeak had moved >1300 m from Sonora to the Michoacán‐Jalisco border region, where it remained until the last GPS‐determined location was obtained on 24 March 2015. The seasonal timing of these movements and the length of stay in Sonora are consistent with the expected behavior of a molt‐migrating bird. Remote‐sensed enhanced vegetation index (EVI) data indicated that the grosbeak arrived in the monsoon region near the area's annual peak in EVI, and then, as the index was declining sharply, departed for the Michoacán‐Jalisco region, where the index also declined during the same period, but substantially less so than in Sonora. Climate change in the coming decades is expected to delay the annual onset of the monsoon while also accelerating the initiation of arid, summer‐like conditions throughout much of western North America, possibly yielding a temporal mismatch between fall migration and the monsoon‐driven conditions that may be critical for molt‐migrating birds.     

Increased temperature induces leaffolder outbreak in rice field

The rice leaffolder, Cnaphalocrosis medinalis Güenée (Lepidoptera: Crambidae), has emerged as a serious pest with significant outbreaks over the last decade in several rice‐growing countries, including China and Bangladesh and resulting in heavy rice yield losses. Climate changes (particularly high temperatures in late winter in Bangladesh) coincide with upsurge in outbreaks of this pest. We generated a statistical model using more than two decades of data to show that increased temperatures associate with this upsurge. Over the 22‐year model period, leaffolder populations in November increased significantly, corresponding to significant monthly temperature trends (but not rainfall) over the same period. Utilizing a linear model, we find that increasing temperature interacts with the amount of rainfall. With the variable month as a proxy for all seasonal effects affecting leaffolder abundance, the model reveals a significant correspondence with climate variations compared to average conditions; specifically, the model predicts that increasing maximum temperatures will lead to more leaffolder, while more rain will decrease their abundance. This study suggests that warmer environment contributed to recent outbreaks of leaffolder in rice‐growing countries; thus, climate change increases rice yield losses by increasing pest population in the field.     

中国鼠疫宿主鼠类丰富度格局及疫区环境因子分析

鼠疫等媒介疾病地理分布规律的认识对于疫病防控具有重要意义,分析影响疾病地理分布的因素进而预测疾病发生趋势已经成为目前研究的热点。本文在分析鼠间鼠疫及其宿主鼠类地理分布数据以及相关环境数据的基础上,探讨了鼠疫宿主丰富度与环境因子的关系以及影响鼠间鼠疫发生的主要环境因素。我国的鼠疫宿主鼠类在干旱区和季风区过渡带的县级行政单元中物种丰富度最高,物种丰富度与高度差正相关性最高。发生鼠间鼠疫的地区,宿主鼠类丰富度较高,未发生鼠间鼠疫的地区,宿主鼠类丰富度较低。影响鼠间鼠疫发生的主要环境因子包括蚤类物种数、年均气温、年降雨量、年均相对湿度和年日照时数和气候因子,其次是包括高度差、植被类型数、土壤类型数和地貌类型数的景观因子,以及反映海拔和宿主鼠类物种数的地形和宿主鼠类因子。在未来全国气候变干变暖条件下,我国鼠间鼠疫发病区可能随宿主鼠类的迁移而扩展。     

Are recession populations of the desert locust (Schistocerca gregaria) remnants of past swarms?

The desert locust (Schistocerca gregaria) undergoes crowding-induced phase transformation from solitary to gregarious, which involves changes in behaviour, colour, development, morphometry, fecundity and endocrine physiology. During recession, solitary locusts persist in the central, drier part of the species' range in small pocket populations that are prone to extinction. During the intermittent upsurges and the subsequent plagues, gregarious swarms attain huge population size and invade a vast area causing major damage to agriculture. A highly variable nuclear DNA marker, a noncoding 3' end fragment of an antennapedia-class homeobox gene, was screened in locust samples from Eritrea. Despite the homogenizing potential of plague swarms, the last of which was in 1986-89 and originated in this region, the population genetic structure of solitary phase locusts along the Red Sea coast of Eritrea revealed significant divergence. The pattern of divergence indicated that the invasion of the western and northern plains in the summer of 1995 may not, as reported then, have originated in eastern Chad or western Sudan. A number of interrelated hypotheses have been presented to explain the observed genetic heterogeneity between the sampled populations. We conclude, with caution due to the limited sample sizes, that: (i) geographical isolation between breeding sites during plagues and recession; (ii) the marked differences in the flight behaviour of plague swarms and recession populations; (iii) possible failure of gregarious locusts to solitarize and re-establish in recession areas; and (iv) the effect of repeated extinction and recolonization in the meta-population contribute to the maintenance of the genetic structure of recession populations. Potentially productive future research has been identified.     

Breeding latitude leads to different temporal but not spatial organization of the annual cycle in a long‐distance migrant

The temporal and spatial organization of the annual cycle according to local conditions is of crucial importance for individuals’ fitness. Moreover, which sites and when particular sites are used can have profound consequences especially for migratory animals, because the two factors shape interactions within and between populations, as well as between animal and the environment. Here, we compare spatial and temporal patterns of two latitudinally separated breeding populations of a trans‐Equatorial passerine migrant, the collared flycatcher Ficedula albicollis, throughout the annual cycle. We found that migration routes and non‐breeding residency areas of the two populations largely overlapped. Due to climatic constraints, however, the onset of breeding in the northern population was approximately two weeks later than that of the southern population. We demonstrate that this temporal offset between the populations carries‐over from breeding to the entire annual cycle. The northern population was consistently later in timing of all subsequent annual events – autumn migration, non‐breeding residence period, spring migration and the following breeding. Such year‐round spatiotemporal patterns suggest that annual schedules are endogenously controlled with breeding latitude as the decisive element pre‐determining the timing of annual events in our study populations.     

Integrating stable isotopes,parasite, and ring‐reencounter data to quantify migratory connectivity—A case study with Barn Swallows breeding in Switzerland,Germany, Sweden,and Finland

Ecosystems around the world are connected by seasonal migration. The migrant animals themselves are influenced by migratory connectivity through effects on the individual and the population level. Measuring migratory connectivity is notoriously difficult due to the simple requirement of data conveying information about the nonbreeding distribution of many individuals from several breeding populations. Explicit integration of data derived from different methods increases the precision and the reliability of parameter estimates. We combine ring‐reencounter, stable isotope, and blood parasite data of Barn Swallows Hirundo rustica in a single integrated model to estimate migratory connectivity for three large scale breeding populations across a latitudinal gradient from Central Europe to Scandinavia. To this end, we integrated a non‐Markovian multistate mark‐recovery model for the ring‐reencounter data with normal and binomial mixture models for the stable isotope and parasite data. The integration of different data sources within a mark‐recapture modeling framework enables the most precise quantification of migratory connectivity on the given broad spatial scale. The results show that northern‐breeding populations and Southern Africa as well as southern‐breeding populations and Western–Central Africa are more strongly connected through Barn Swallow migration than central European breeding populations with any of the African wintering areas. The nonbreeding distribution of Barn Swallows from central European breeding populations seems to be a mixture of those populations breeding further north and south, indicating a migratory divide.     

蝗虫多型现象的神经内分泌调控

蝗虫有两种型,即散居型和群居型。蝗灾通常由群居型蝗虫所引发。多年来人们试图找到控制蝗虫由散居型向群居型转变的关键因子,以期控制蝗虫危害。该文主要从神经内分泌的角度概述了蝗虫多型性的生理机制,重点介绍了保幼激素、蜕皮激素和脑神经肽［His7］-corazonin在蝗虫多型性中的主要作用和机制。     

Evidence from traditional and new technologies for northward migrations of Australian plague locusts (Chortoicetes terminifera) (Walker) (Orthoptera: Acrididae) to western Queensland

Abstract The development of recent infestations of the Australian plague locust (Chortoicetes terminifera) (Walker) (Orthoptera: Acrididae) has been traced using traditional survey data combined with information from several modern technologies including simulation of windborne transport trajectories, direct observation with entomological radar and satellite imagery. The results indicate that migration from spring generations in the southern and eastern parts of the species range, including agricultural areas, to the summer rainfall areas in arid western Queensland (Qld) has contributed to the development of infestations on several occasions. Migration from swarm populations in New South Wales to western Qld in November and December 1999 contributed to a rapid population increase that, over a sequence of generations, led to the major infestation of agricultural areas in March and April 2000. There is evidence that northward migrations also occurred in 1995, 1997 and 2000. These contributed to the early summer populations in Qld, but did not result in large migrations to the south in autumn. These observations suggest that a pattern of exchange migration across much of the geographical range of the species between regions of winter and summer rainfall characterizes the spatial dynamics of this species. This pattern appears to be adaptive and suggests migration in C. terminifera is sustained by contemporary natural selection.