Eastern Migratory Population Research Articles

Experimental Suppression of Red Imported Fire Ants (Solenopsis invicta) Has Little Impact on the Survival of Eggs to Third Instar of Spring-Generation Monarch Butterflies (Danaus plexippus) Due to Buffering Effects of Host-Plant Arthropods

The eastern migratory population of the monarch butterfly (Danaus plexippus) has shown evidence of declines in recent years. During early spring, when the population is at its smallest, red imported fire ants (RIFA) (Solenopsis invicta) have been implicated as having devastating effects on monarch egg and larval survival, but there are no conclusive experimental data to support this contention. The purpose of this study was to determine the effect of RIFA on the survival of spring monarch eggs to third instar larvae. Three treatments were analyzed: control plots, RIFA-suppressed plots, and RIFA-enhanced plots. Other host-plant arthropods were also documented. In control plots, monarch survival was unrelated to RIFA abundance on or around the plants. For both years combined, RIFA suppression had little impact on monarch survival. In one of the two years, higher survival occurred in the suppressed treatment, but confidence in this difference was low. In control plots, monarch survival increased with increasing numbers of other arthropods (not including RIFA) on the host plant. Predator pressure did not vary relative to arthropod abundance, and RIFA only occupied plants in large numbers when large numbers of other arthropods were also present. The presence of RIFA did not affect predator pressure. RIFA artificially drawn onto host plants created artificially high predator pressure, and monarch survival was low. Long-term use of bait to control RIFA may not be cost-effective provided surrounding biodiversity is high. Efforts to promote spring monarchs should focus on promoting biodiversity in addition to planting milkweed.

Multi-season climate projections forecast declines in migratory monarch butterflies.

Climate change poses a unique threat to migratory species as it has the potential to alter environmental conditions at multiple points along a species' migratory route. The eastern migratory population of monarch butterflies (Danaus plexippus) has declined markedly over the last few decades, in part due to variation in breeding-season climate. Here, we combined a retrospective, annual-cycle model for the eastern monarch population with climate projections within the spring breeding grounds in eastern Texas and across the summer breeding grounds in the midwestern U.S. and southern Ontario, Canada to evaluate how monarchs are likely to respond to climate change over the next century. Our results reveal that projected changes in breeding-season climate are likely to lead to decreases in monarch abundance, with high potential for overwintering population size to fall below the historical minimum three or more times in the next two decades. Climatic changes across the expansive summer breeding grounds will also cause shifts in the distribution of monarchs, with higher projected abundances in areas that become wetter but not appreciably hotter (e.g., northern Ohio) and declines in abundance where summer temperatures are projected to increase well above those observed in the recent past (e.g., northern Minnesota). Although climate uncertainties dominate long-term population forecasts, our analyses suggest that we can improve precision of near-term forecasts by collecting targeted data to better understand relationships between breeding-season climate variables and local monarch abundance. Overall, our results highlight the importance of accounting for the impacts of climate changes throughout the full-annual cycle of migratory species.

Natal dispersal of Whooping Cranes in the reintroduced Eastern Migratory Population.

Natal dispersal is a key demographic process for evaluating the population rate of change, especially for long‐lived, highly mobile species. This process is largely unknown for reintroduced populations of endangered avian species. We evaluated natal dispersal distances (NDD) for male and female Whooping Cranes (Grus americana) introduced into two locations in central Wisconsin (Necedah National Wildlife Refuge, or NNWR, and the Eastern Rectangle, or ER) using a series of demographic, spatial, and life history‐related covariates. Data were analyzed using gamma regression models with a log‐link function and compared using Akaike information criterion corrected for small sample sizes (AICc). Whooping Cranes released in the ER dispersed 261% further than those released into NNWR, dispersal distance increased 4% for each additional nesting pair, decreased about 24% for males as compared to females, increased by 21% for inexperienced pairs, and decreased by 3% for each additional year of age. Natal philopatry, habitat availability or suitability, and competition for breeding territories may be influencing observed patterns of NDD. Whooping Cranes released in the ER may exhibit longer NDD due to fragmented habitat or conspecific attraction to established breeding pairs at NNWR. Additionally, sex‐biased dispersal may be increasing in this population as there are more individuals from different natal sites forming breeding pairs. As the population grows and continues to disperse, the drivers of NDD patterns may change based on individual or population behavior.

High Survivorship of First-Generation Monarch Butterfly Eggs to Third Instar Associated with a Diverse Arthropod Community.

Simple SummaryThe eastern migratory population of the monarch butterfly has been the focus of extensive conservation efforts in recent years. However, there are gaps in our knowledge about the survival of first, or spring generation, monarchs in their core areas of Texas, Oklahoma, and Louisiana. This is important because the spring generation represents the first stage of annual recovery from overwinter mortality. It is, therefore, an important stage for monarch conservation efforts. This study showed that, in the context of a complex arthropod community in north Texas, first generation monarch survival was high. The study found that survival was not directly related to predators on the host plant, but was higher on host plants that harbored a greater number and variety of other, non-predatory arthropods. This is possibly because the presence of alternate, preferable prey enabled monarch eggs and larvae to be overlooked by predators. The implication is that, at least in the southern U.S., monarch conservation should consider strategies that promote diverse functional arthropod communities.Based on surveys of winter roost sites, the eastern migratory population of the monarch butterfly (Danaus plexippus) in North America appears to have declined in the last 20 years and this has prompted the implementation of numerous conservation strategies. However, there is little information on the survivorship of first-generation monarchs in the core area of occupancy in Texas, Oklahoma, and Louisiana where overwinter population recovery begins. The purpose of this study was to determine the survivorship of first-generation eggs to third instars at a site in north Texas and to evaluate host plant arthropods for their effect on survivorship. Survivorship to third instar averaged 13.4% and varied from 11.7% to 15.6% over three years. The host plants harbored 77 arthropod taxa, including 27 predatory taxa. Despite their abundance, neither predator abundance nor predator richness predicted monarch survival. However, host plants upon which monarchs survived often harbored higher numbers of non-predatory arthropod taxa and more individuals of non-predatory taxa. These results suggest that ecological processes may have buffered the effects of predators and improved monarch survival in our study. The creation of diverse functional arthropod communities should be considered for effective monarch conservation, particularly in southern latitudes.

First Description of Nesting Behavior of a Same-Sex Pair of Whooping Cranes (Grus americana) in the Reintroduced Eastern Migratory Population

Accounts of same-sex nesting behaviors have been observed in several wild and captive bird species, but the evolutionary adaptation of this behavior remains unclear. There have been reports of male-male and female-female territorial pairs of Whooping Cranes (Grus americana) in two reintroduced populations. However, this is the first documentation of nesting behavior of a same-sex pair of Whooping Cranes. The incubation rhythm and nesting behaviors of a female-female Whooping Crane pair nesting at McMillan Marsh Wildlife Area in Marathon County, Wisconsin, USA, were documented using a nest camera. The female-female pair displayed comparable nest constancy (99.7%) and incubation behaviors to male-female Whooping Crane pairs in the same population. Both females of the pair incubated the nest; however, the older female incubated for longer periods of time than the younger female (235.8 ± 20.7 min and 168.3 ± 17.2 min, respectively). The pair exchanged incubation duties 5.7 ± 0.2 and 1.1 ± 0.2 times daily during daylight hours and nighttime hours, respectively. The two females incubated the nest for at least 28 days, but fate of the eggs was not determined. Ultimately, the cause of this pairing remains unknown. This behavior could have implications for the breeding success of this small, reintroduced population of Whooping Cranes.

Invasive paper wasp turns urban pollinator gardens into ecological traps for monarch butterfly larvae

Invasive species can be particularly disruptive when they intersect with organisms of conservation concern. Stabilizing the declining eastern migratory population of monarch butterflies (Danaus plexippus) is projected to require extensive habitat restoration across multiple land use sectors including metropolitan areas. Numerous conservation programs encourage urban citizens to plant gardens with milkweeds, the obligate larval host plants of the monarch. Here, we show that predation by Polistes dominula, an invasive paper wasp that is particularly abundant in urban settings, can turn such sites into ecological traps for monarch larvae. Polistes dominula was the predominant paper wasp seen foraging in central Kentucky pollinator gardens. In 120 observed encounters with monarch larvae on milkweeds in gardens, most second to fourth instars were killed, whereas most fifth instars escaped by thrashing or dropping. The wasps bit and carried off second instars whole, whereas third and fourth instar kills were first gutted, then processed and carried away piecemeal. Predation on sentinel larvae was much higher in urban gardens than in rural settings. The wasps exploited ornamental butterfly “hibernation boxes” in pollinator gardens as nesting habitat. Polistes dominula is an under-recognized predator that may diminish the urban sector’s contributions to monarch habitat restoration.

Evidence for a Growing Population of Eastern Migratory Monarch Butterflies Is Currently Insufficient

The eastern migratory population of monarch butterflies has experienced a multi-decadal decline, but a recent increase in abundance (to 6.05 ha in winter 2018) has led some observers to question whether the population has reversed its long-standing decline and embarked on a trajectory of increasing abundance. We examined this possibility through changepoint analyses, assessing whether a change in trajectory existed. We found evidence of a change in trajectory in 2014, but insufficient statistical support for a significantly increasing population since that time (β = 0.285, 95% CI = -0.127, 0.697). If the population estimate for winter 2019 is ≥4.0 ha, we will then be able to credibly assert the population has been increasing since 2014. However, given estimated levels of time series variability, presumed habitat capacity and no recent change in status or trend, there was a 13.5% probability of observing a population estimate as large or larger than was reported for winter 2018. Our analyses highlight the incredible difficulty in drawing robust conclusions from annual changes in abundance over a short time series, especially for an insect that commonly exhibits considerable year-to-year variation. Thus, we urge caution when drawing conclusions regarding species status and trends for any species for which limited data are available.

Monarch butterfly population decline in North America: identifying the threatening processes.

The monarch butterfly (Danaus plexippus) population in North America has sharply declined over the last two decades. Despite rising concern over the monarch butterfly's status, no comprehensive study of the factors driving this decline has been conducted. Using partial least-squares regressions and time-series analysis, we investigated climatic and habitat-related factors influencing monarch population size from 1993 to 2014. Potential threats included climatic factors, habitat loss (milkweed and overwinter forest), disease and agricultural insecticide use (neonicotinoids). While climatic factors, principally breeding season temperature, were important determinants of annual variation in abundance, our results indicated strong negative relationships between population size and habitat loss variables, principally glyphosate use, but also weaker negative effects from the loss of overwinter forest and breeding season use of neonicotinoids. Further declines in population size because of glyphosate application are not expected. Thus, if remaining threats to habitat are mitigated we expect climate-induced stochastic variation of the eastern migratory population of monarch butterfly around a relatively stationary population size.

Restoring monarch butterfly habitat in the Midwestern US: ‘all hands on deck’

The eastern migratory population of monarch butterflies (Danaus plexippus plexippus) has declined by >80% within the last two decades. One possible cause of this decline is the loss of ≥1.3 billion stems of milkweed (Asclepias spp.), which monarchs require for reproduction. In an effort to restore monarchs to a population goal established by the US Fish and Wildlife Service and adopted by Mexico, Canada, and the US, we developed scenarios for amending the Midwestern US landscape with milkweed. Scenarios for milkweed restoration were developed for protected area grasslands, Conservation Reserve Program land, powerline, rail and roadside rights of way, urban/suburban lands, and land in agricultural production. Agricultural land was further divided into productive and marginal cropland. We elicited expert opinion as to the biological potential (in stems per acre) for lands in these individual sectors to support milkweed restoration and the likely adoption (probability) of management practices necessary for affecting restoration. Sixteen of 218 scenarios we developed for restoring milkweed to the Midwestern US were at levels (>1.3 billion new stems) necessary to reach the monarch population goal. One of these scenarios would convert all marginal agriculture to conserved status. The other 15 scenarios converted half of marginal agriculture (730 million stems), with remaining stems contributed by other societal sectors. Scenarios without substantive agricultural participation were insufficient for attaining the population goal. Agricultural lands are essential to reaching restoration targets because they occupy 77% of all potential monarch habitat. Barring fundamental changes to policy, innovative application of economic tools such as habitat exchanges may provide sufficient resources to tip the balance of the agro-ecological landscape toward a setting conducive to both robust agricultural production and reduced imperilment of the migratory monarch butterfly.

The Disconnect Between Summer and Winter Monarch Trends for the Eastern Migratory Population: Possible Links to Differing Drivers

The decline of the eastern population of the migratory monarch has become a topic of great concern, but has been based entirely on patterns observed in overwinter colony sizes. Less attention has been paid to population trends during other phases of the migratory cycle. Here, we present an analysis of trends using three monitoring programs, one focused on overwinter colony size and two focused on summer breeding grounds. We discovered an alarming steepening in the decline of winter colony size since 2008. However, population indices from two independent summer monitoring programs were characterized by high year-to-year variability and no statistically detectable trends over time. Despite the mismatch in summer and winter patterns, there is still an association between the yearly fluctuations between these key periods, suggesting a link in population dynamics throughout the year. Further, a suggestion of a downturn near the end of the summer time-series should be carefully tracked into the future. We discuss two possible reasons for this disconnect: 1) higher levels of variance or possibly biased sampling could weaken any statistical signal, and 2) losses during fall migration could potentially contribute to overwinter declines.

A hierarchical nest survival model integrating incomplete temporally varying covariates

Nest success is a critical determinant of the dynamics of avian populations, and nest survival modeling has played a key role in advancing avian ecology and management. Beginning with the development of daily nest survival models, and proceeding through subsequent extensions, the capacity for modeling the effects of hypothesized factors on nest survival has expanded greatly. We extend nest survival models further by introducing an approach to deal with incompletely observed, temporally varying covariates using a hierarchical model. Hierarchical modeling offers a way to separate process and observational components of demographic models to obtain estimates of the parameters of primary interest, and to evaluate structural effects of ecological and management interest. We built a hierarchical model for daily nest survival to analyze nest data from reintroduced whooping cranes (Grus americana) in the Eastern Migratory Population. This reintroduction effort has been beset by poor reproduction, apparently due primarily to nest abandonment by breeding birds. We used the model to assess support for the hypothesis that nest abandonment is caused by harassment from biting insects. We obtained indices of blood-feeding insect populations based on the spatially interpolated counts of insects captured in carbon dioxide traps. However, insect trapping was not conducted daily, and so we had incomplete information on a temporally variable covariate of interest. We therefore supplemented our nest survival model with a parallel model for estimating the values of the missing insect covariates. We used Bayesian model selection to identify the best predictors of daily nest survival. Our results suggest that the black fly Simulium annulus may be negatively affecting nest survival of reintroduced whooping cranes, with decreasing nest survival as abundance of S. annulus increases. The modeling framework we have developed will be applied in the future to a larger data set to evaluate the biting-insect hypothesis and other hypotheses for nesting failure in this reintroduced population; resulting inferences will support ongoing efforts to manage this population via an adaptive management approach. Wider application of our approach offers promise for modeling the effects of other temporally varying, but imperfectly observed covariates on nest survival, including the possibility of modeling temporally varying covariates collected from incubating adults.

Which uncertainty? Using expert elicitation and expected value of information to design an adaptive program

Natural resource management is plagued with uncertainty of many kinds, but not all uncertainties are equally important to resolve. The promise of adaptive management is that learning in the short-term will improve management in the long-term; that promise is best kept if the focus of learning is on those uncertainties that most impede achievement of management objectives. In this context, an existing tool of decision analysis, the expected value of perfect information (EVPI), is particularly valuable in identifying the most important uncertainties. Expert elicitation can be used to develop preliminary predictions of management response under a series of hypotheses, as well as prior weights for those hypotheses, and the EVPI can be used to determine how much management could improve if uncertainty was resolved. These methods were applied to management of whooping cranes ( Grus americana), an endangered migratory bird that is being reintroduced in several places in North America. The Eastern Migratory Population of whooping cranes had exhibited almost no successful reproduction through 2009. Several dozen hypotheses can be advanced to explain this failure, and many of them lead to very different management responses. An expert panel articulated the hypotheses, provided prior weights for them, developed potential management strategies, and made predictions about the response of the population to each strategy under each hypothesis. Multi-criteria decision analysis identified a preferred strategy in the face of uncertainty, and analysis of the expected value of information identified how informative each strategy could be. These results provide the foundation for design of an adaptive management program.

Associations between host migration and the prevalence of a protozoan parasite in natural populations of adult monarch butterflies

Summary1. Monarch butterfliesDanaus plexippus(L.) (Lepidoptera: Nymphalidae) are susceptible to infection by the obligate protozoan parasiteOphryocystis elektroscirrha(McLaughlin and Myers) (Apicomplexa: Neogregarinida). Because monarchs form resident and migratory populations in different parts of the world, this host–parasite system provides the opportunity to examine how variation in parasite prevalence relates to host movement patterns.2. Parasite prevalence was evaluated using 14 790 adult monarchs captured between 1968 and 1997. Comparison of three populations in North America indicated that parasite prevalence is associated negatively with host dispersal distances. A continuously breeding, nonmigratory population in southern Florida showed high prevalence (over 70% heavily infected). The western population migrates moderate distances to overwintering sites on the Pacific Coast and has intermediate prevalence (30% heavily infected). The eastern migratory population, which travels the longest distance to Mexican overwintering sites, has exhibited less than 8% infection throughout the past 30 years.3. Variation in parasite loads within North American migratory populations was investigated to determine whether the prevalence of heavy infection and average parasite loads declined during migration or overwintering. Average parasite loads of summer‐breeding adults in western North America decreased with increasing distance from overwintering sites. This suggests that heavily infected monarchs are less likely to remigrate long distances in spring. No differences in the frequency of heavily infected adults were found among eastern or western North American monarchs throughout the overwintering period, however, suggesting that this parasite does not affect overwintering mortality.4. Changes in the prevalence of monarchs with low parasite loads demonstrate that spore transfer occurs during migration and overwintering, possibly when adult butterflies contact each other as a result of their clustering behaviour.5. This study of geographical and temporal variation inO. elektroscirrhaamong populations ofD. plexippusdemonstrates the potential role of seasonal migration in mediating interactions between hosts and parasites, and suggests several mechanisms through which migratory behaviour may influence parasite prevalence.