RECONNECTING LANDSCAPES IN A CHANGING WORLD

• Sponsoring Institution: National Institute of Food and Agriculture
• Project Status: COMPLETE
• Funding Source: HATCH
• Reporting Frequency: Annual
• Accession No.: 1018091
• Project Start Date: Dec 1, 2018
• Project End Date: Nov 30, 2023
• Program Code: [(N/A)]- (N/A)

Recipient Organization
MICHIGAN STATE UNIV
(N/A)
EAST LANSING,MI 48824

Performing Department
Kellogg Biological Station

Non Technical Summary
Habitat loss and fragmentation are the most important threats to biodiversity (Wilcove et al. 1998), and these threats are exacerbated by land use and climate change. Landscape modification is so widespread that impacts on diversity and ecosystem function are likely to be devastating. As one indicator, we conducted a global analysis of forest loss and fragmentation and found that 70% of the world's forest is now within a kilometer of the edge of the forest. Through a review of all long-term habitat fragmentation experiments, we found that habitat fragmentation has degrading effects that propagate through entire ecological systems. The Haddad Lab seeks to understand threats of land use and climate change, and, more importantly, to test conservation strategies that reduce them. Our central focus is restoration of connectivity in degraded and fragmented landscapes, especially in agricultural and managed forest landscapes. We will accomplish our objectives in our unique studies that are at the interface of basic and applied ecology, are large experiments (hectare scale), and are of decades duration. We will pursue our objectives in three experiments. First, we will continue research on US Forest Service land to test the effects of landscape corridors on biodiversity. Second, we will continue research on Department of Defense, US Fish and Wildlife Service, and National Park Service lands to test the effects of restoration of connectivity on rare species. Third, we will work in the Long Term Ecological Research Site at Kellogg Biological Station to test the effects of restored connectivity on agriculture and the environment.

Animal Health Component

50%

Research Effort Categories

Basic

50%

Applied

50%

Developmental

0%

Classification

Knowledge Area (KA)	Subject of Investigation (SOI)	Field of Science (FOS)	Percent
136	0860	1070	33%
131	0510	1070	67%

Knowledge Area
136 - Conservation of Biological Diversity; 131 - Alternative Uses of Land;

Subject Of Investigation
0510 - Wilderness; 0860 - Endangered species;

Field Of Science
1070 - Ecology;

Keywords

corridor

connectivity

habitat loss

habitat fragmentation

endangered species

conservation

prairie strips

Goals / Objectives
I. Landscape CorridorsMy lab collaborates with a large team and we are currently in the middle of a 10 year study funded by NSF-LTREB (Long-term Research in Environmental Biology). This study is built on a 25 year long experiment. We will answer three overarching questions:Question 1: By what mechanisms are connectivity effects on diversity generated and maintained? We have identified between-patch connectivity as a key factor affecting diversity, with connectivity increasing species richness by 20% within five years of initiating our experiment. This is the strongest empirical evidence for a positive landscape connectivity-species diversity relationship, yet the mechanisms that lead to this pattern remain unknown. Although we have good evidence that colonization dynamics caused the initial connectivity effects on species richness, we will now determine how connectivity increased and is currently maintaining elevated species richness.Question 2: Does the importance of connectivity for species diversity change over time? Our data to date illustrate an initial pattern of rapid increase in species richness in connected relative to unconnected patches, followed by an apparent plateau. We will examine the role of connectivity for species richness over the long-term.Question 3: Does connectivity increase the temporal stability of species richness? We have observed a persistent effect of connectivity on plant species richness, but our current data also suggest substantial inter-annual variation in this effect, varying by a factor of two since the plateau was reached. We predict that connectivity stabilizes diversity and community composition following disturbance - increasing resistance and resilience to disturbance - because of connectivity's role in increasing colonization following disturbance. We will test how connectivity affects temporal stability of species richness in response to disturbance caused by (prescribed) fires and variation in precipitation - broadly relevant perturbations in context of ecosystem management and climate change.II. Landscape RestorationA complete understanding of restoration in a metapopulation framework is the most urgent need for recovery of St. Francis' Satyr, and of many species that occur in disturbance-dependent landscapes. St. Francis' Satyr is federally endangered and is only found on Ft. Bragg army installation. Perhaps the only reason it has not already gone entirely extinct at Ft. Bragg is because of artillery impact zones, which retain reliable habitat patches in a spatially complex landscape because of direct disturbance (artillery ignites near-annual fires) and because beavers are not managed there as pests. Our objectives are to test metapopulation management strategies for St. Francis' Satyr on Ft. Bragg outside of artillery impact zones and use what we learn to advance the establishment of new, viable metapopulations off of Ft. Bragg.Although critical to the long-term persistence of these disturbance-dependent butterflies, the metapopulation framework has never been investigated. This deficit has left the species in peril and has prevented reintroduction. Our proposed research would permit us to make the most important advance needed in St. Francis' Satyr recovery: scaling up from our recent focus on the restoration of small, isolated populations of St. Francis' Satyr to a new understanding of metapopulations at larger spatial scales.Without natural disturbances or management interventions, we relegate existing populations to extirpation because of loss of host plants. Further, because butterflies cannot swim or survive fire, disturbance (natural and managed) also causes widespread death of individuals in discrete populations. However, because this process regenerates habitat, if there is a robust metapopulation St. Francis' Satyr can disperse from elsewhere to repopulate restored or regenerated habitat. Recovering St. Francis' Satyr will require the creation and maintenance of habitat patches at different stages of plant succession. Simultaneous succession or restoration of large areas at once is undesirable as it may cause extirpation of entire metapopulations and result in synchronous decline of habitat. Instead, ideal St. Francis' Satyr habitat dynamics are strongly asynchronous at the local scale with patches that are: 1) recently restored (i.e., disturbed); 2) ready for recolonization; 3) at peak St. Francis' Satyr population, producing potential colonists; and 4) beginning to undergo successional decline, all intermixed at a spatial scale such that individual butterflies can move between them. In the next five years, we will answer two questions:Question 1: What is the optimal frequency and patch size/spatial arrangement of habitat restoration or management (e.g., controlled burns) to minimize metapopulation extinction risk, taking into account the desired asynchrony between populations that are close enough to be linked by dispersal?Question 2: Given the spatial scale of population synchrony, will dispersal reduce the risk that entire metapopulations will go extinct? How should corridors best be used to enhance movement, both between populations on the army base and between on-base and off-base populations?3. KBS-LTER and Conservation Strips.Our overarching question is: When does diversity, and management that promotes it, increase resilience to land use change? We will test whether diversity-resilience relationships commonly observed in restoration are also prevalent in annual row crops or harvested perennial crops. We will also test how diversity-resilience relationships vary across ecosystem functions.We will create diverse, perennial Conservation Strips within agricultural treatments. The existing treatments contain winter cover crops that are tilled into the soil in early spring prior to planting the main crop. This is a major annual disturbance creating temporally unfavorable conditions for many taxa in the main crop area. Given that dispersal is a key driver of biodiversity, especially in heavily disturbed annual crops, we test how Conservation Strips influence arthropod diversity and the resilience of ecosystem services provided by arthropods.We predict that following disturbance, more transient taxa such as mobile predators and parasitoids will preferentially move to Conservation Strips that can harbor aphid prey and provide pollen (Woltz et al. 2012, Woltz and Landis 2013), a protein resource used by some aphid natural enemies. These species may not permanently colonize conservation strips but rather use them as temporal or spatial stepping stones as they move through the landscape (e.g., aphid pests of field crops and their natural enemies, principally lady beetles and Hymenopteran parasitoids). We also predict that other taxa, including carabids, ants, and in future work microbes will establish persistent populations, from which individuals can move into adjacent crops.To investigate effects of the wider arthropod community on the resilience of ecosystem functions, we will sample ground-dwelling arthropods with pitfall traps in all treatments (Bestelmeyer et al. 2000). Carabid communities have been periodically sampled at the MSCE in the past, and they show clear responses to management intensity (Clark et al. 1997), which cascades to influence post-dispersal weed seed predation (Menalled et al. 2007). Ant abundance and diversity at KBS vary with crop identity and perenniality (Wills & Landis 2017). We will test the effects of carabids and ants on pest suppression by use of sentinel weed seed and pest insect egg card exposures (Werling et al. 2011, Meehan et al. 2012).

Project Methods
I. Savannah River Site Corridor ExperimentIn 2000, we created eight large-scale experimental landscapes (hereafter "blocks") in USDA Forest Service land at the Savannah River Site, South Carolina that explicitly test for connectivity effects while controlling for patch area and shape (see Figure 1). Each block is comprised of five patches (~1 ha each) and one corridor that connects two of the patches. Within each block, one center patch (100 x 100m, 1ha) is surrounded by four peripheral patches, each the same distance (150m) from the center patch. The center patch is connected to one peripheral patch ("connected patch") by a 150m x 25m corridor. The other three peripheral patches are unconnected ("unconnected patches"). The orientation of the corridor relative to north was chosen at random, so that corridor orientation is unbiased with respect to wind and animal movement. All patches were created by cutting mature plantation loblolly pine forest in 2000 and burning. The blocks are maintained with prescribed fires every 2-3 years, mimicking historical disturbance regimes. All patches have been planted with longleaf pine seedlings and with understory species characteristic of longleaf pine savanna, with the ultimate goal of restoring the entire landscape to longleaf pine savanna. A forested buffer is maintained within a circle whose radius extends 150m beyond the outside boundary of peripheral patches.We focus on plant species richness, colonization, and extinction. We will first quantify rates of extinction and colonization with our data on patch-level occupancy. Each year, we will tally the total numbers of extinction or colonization events, quantified as instances of a species being newly absent or newly present, relative to the previous year, in each patch as response variables. This will allow us to test whether colonization or extinction rates differ among patch types and whether these rates change over time. To distinguish between mechanisms of lower extinction rates (rescue effects and/or population sizes). We will focus on colonization rates and extinction rates of individual species. For all species occurring within a connected and unconnected patch across at least four experimental landscapes, we will estimate colonization and extinction rates for each experimental patch type (connected andunconnected) (Morris and Doak 2002). For these same species, we will evaluate whether our metrics of population size vary between connected and unconnected patches.II. Restoring Population Sources for Rare ButterfliesWe will employ two methods of de novo habitat restoration for St. Francis' satyr butterfly: hardwood removal and wetland creation (Figure 2). A key advantage of implementing restoration as part of this project is that we can design restoration as a factorial experiment in a completely blocked design to control for landscape effects. We will restore St. Francis' Satyr habitat first by removing hardwood trees to retard wetland succession. Hardwoods block light and transpire large quantities of water, reducing the abundance of sedges on which St. Francis' Satyr larvae feed. We will also restore St. Francis' satyr habitat by damming streams to create wetlands, then reduce or remove dams to permit sedge development. Aquadams (http://www.aquadam.com/) provide a flexible means to create temporary dams.We chose sites identified as having the potential for suitable habitat within the dispersal range of extant St. Francis' Satyr populations, based on habitat and movement models from our previous research. We will initiate populations at all restored sites by introducing captive reared individuals.We will measure adult survival and population size in each restored site using mark-recapture methods, which we have optimized for St. Francis' satyr (Haddad et al. 2008). Briefly, adult butterflies will be monitored daily along fixed transects within each restored site. Butterflies will be captured with a hand net and marked with a unique number on the underside of both wings. If possible, marks will be resighted using close-focus binoculars to avoid disturbing butterflies. We will estimate butterfly daily survival rate, detectability, and total population sizes through each adult flight period.Larval St. Francis' Satyr are extremely cryptic, and can be difficult to find even when restricted to one plant. Therefore we will use the ratio of emerging adults to initial, newly hatched larvae to estimate hatching-to-adult survival. Specifically, we will surround three 1x1m2 plots in each restored wetland with aluminum flashing on which caterpillars cannot climb. Within each plot, we will add 25 newly hatched caterpillars, and will leave plots open to the environment for the duration of the larval period.We will test effects of restoration on St. Francis' Satyr demography, behavior, and population growth . Specifically, we will determine when we are creating population sources and population sinks via restoration. We will also construct spatially explicit individual-based models that integrate demography in remnant, managed, and restored patches with movement across the landscape. We will then link the population processes of demography and dispersal with dynamic GIS maps of the temporal dynamics of habitat quality in managed and restored habitat.III. KBS-LTER Conservation StripsAll objectives will be accomplished within KBS-LTERs Main Cropping System Experiment (MCSE). It is the core focus of KBS LTER, and treatments within this experiment will be used to address all proposed hypotheses (Figure 3). The MCSE (established in 1988) includes treatments spanning a gradient of agricultural intensities that include four annual cropping systems (corn-soybean-wheat rotations under Conventional, No-till, Reduced Input, and Biologically Based/organic managements), two perennial cropping systems (Switchgrass and Hybrid Poplar), unmanaged Early Successional grasslands, and mid- and late-successional forests. This experiment thus represents a gradient of ecological complexity (along with gradients of disturbance frequency, resource subsidization, and perenniality), and we have monitored numerous ecosystem functions continuously since its establishment.In the spring of 2019, we will plant Conservation Strips, into our Reduced Input and Biologically Based treatments at the MCSE. Conservation Strips are widely used to enhance ecosystem services in Europe (Batáry et al. 2015) and increasingly proposed for the US (Schulte et al. 2017). They will allow us to investigate whether perennial habitat within annual crops influences dispersal, community assembly, diversity, and, therefore, resilience of ecosystem functions in crops exposed to frequent disturbance. Specifically, we will install 4.6 m wide strips consisting of perennial grasses and forbs in the center of 1 ha plots, replacing approximately 5% of the existing crop area. They will be designed to enhance diversity and ecological function. Plant species chosen for inclusion will be those commonly included in restorations (Grman et al. 2013, Grman et al. 2015), most attractive to pollinators and natural enemies (Fiedler and Landis 2007) and beneficial microorganisms, and flowering at a range of times throughout the growing season.Within Conservation Strips, we will use sticky cards and sweepnet sampling to determine presence of transient taxa. In addition, we will use pitfall traps and other techniques to determine the population size and area covered by resident species. Outside of Conservation Strips, we will create four sampling transects perpendicular to the strips to assess selected insect communities within and at increasing distances from the strips, and our long-term data will allow for pre-strip vs. poststrip comparisons of a variety of ecosystem functions.

Progress 10/01/19 to 09/30/20

Outputs
Target Audience:My target audience includes the scientific community, land managers at USDA and DOE, and the general public in regard to rare butterflies (especially with book release) and Kellogg Biological Station Longterm Ecological Research study Changes/Problems: Nothing Reported What opportunities for training and professional development has the project provided?My lab employs sixteen people at various stages of professional development, from undergraduate to postdoctoral. All of them are trained in field biology, including in agricultural, forested, and conservation lands. I employed two undergraduates and three postbac students that assisted with research on prairie strips; three postbac students who did work on rare butterflies; two postdocs who did work on prairie strips; and a research assistant each that did work on corridors, rare butterflies, prairie strips, and website development (ConservationCorridor.org). In nearly all of these positions, the people are in training for the next level, typically a permanent job or graduate school. Thus, in addition to training, I provide professional development in other ways that promote their career goals. How have the results been disseminated to communities of interest?In addition to journal articles, I appeared in or was featured in eight stories on rare butterflies, nineteen studies on landscape corridors, thirty five stories on landscape diversity, four stories on conversion from conventional agriculture to no-till, and sixteen stories on landscape fragmentation. These stories appeared in the NY Times, Washington Post, LA Timess, AP (100s of newspapers), NPR, among others. My lab also runs ConservationCorridor.org that reaches 40,000 users annually. What do you plan to do during the next reporting period to accomplish the goals?I. Corridors In the coming year we will study how corridors affect species interactions, specifically moth pollinator diversity and their effects as pollinators. We will study microbial ecology -- the diversity and composition of microbial communities, as well as the ecosystem services they generate in connected patches. Finally, we will make further progress in understanding what affects the stability of plant diversity, including whether the number of species maintains constant over time, or whether weather or fire influence year to year fluctuations II Restoration for butterflies We are near finished collecting data needed to parameterize population models for the endangered St. Francis' Satyr. This year, we will make strong advances in the modeling effort, understanding how restoration and re-restoration interact with landscape connectivity to promote butterfly recovery. We will initiate studies of a butterfly in Michigan, Poweshiek Skipperling. Specifically, we will begin a captive rearing study, to generate individuals to be used in restoration. III. KBS LTER and Prairie Strips We will continue with a third year of data collection on effects effects of strips on pollination, decomposition, ant diversity and function, butterfly abundance and diversity, and other responses. We will begin new studies of effects of growing season drought and increased temperature on demography of butterflies and moths. We will expand studies of long-term dynamics of populations by synthesizing studies across the LTER network.

Impacts
What was accomplished under these goals? I. Landscape Corridor In 2019-2020, we made major progress in showing the impacts of reconnecting landscapes where habitats had been lost or fragmented on biodiversity and on mechanisms underlying connectivity effects. Working in the largest, best replicated experiment in the world to test the role of landscape corridors, we provided the best demonstration of the long-term impacts of corridors on biodiversity, results that were published in Science in October 2020. Specifically, in an 18 year study of plant species in 40, 1ha fragments, some of which were connected by corridors. we found that corridors increased diversity by 14%, and that the effects of corridors are still growing. These results corresponded to mechanisms of effects that we are discovering. First, corridors are increasing colonization by bees. This would promote species interactions (pollination) that would serve to boost biodiversity. Second, we accumulated plant traits that will be used to disentangle corridor effects. Third, we tracked plants populations that are part of a restoration experiment, testing for fragmentation effects on dispersal. Our work is conducted in partnership with the US Forest Service, Savannah River. They assist us in creation and maintanence of our experimental landscapes, and we return our results to aid in their conservation management to optimize biodiversity in working landscapes. This year, we made particular progress in management our experiment through controlled burns that are implemented by the Forest Service. II Landscape Restoration. For the 19th year, we continued population monitoring of the endangered St. Francis' Satyr butterfly. This follows active restoration. In 2019-2020 we restored new sites at Ft. Bragg Army Installation. We also increased efforts for restoration on lands owned by The Nature Conservancy. We made progress in new studies of "re-restoration" -- the natural dynamics needed to impeded vegetative succession and promote butterfly population growth. We advanced modeling studies to understand the effects of precipitation dynamics on populations of the rare Miami Blue butterfly. We made new discoveries about how connectivity must be maintained for vertebrates to promote their migration to cooler climates, all while avoiding the effects of urbanization. III KBS-LTER and conservation strips Much of this year was devoted to studying the effects of Prairie Strips and landscape diversity in reduced input and organic agricultural treatments. We conducted second year studies of pollination, decomposition, and ant and beetle dynamics. We discovered how landscape diversity promotes biodiversity of ants and the ecosystem services ants provide -- primaily through pest predation and weed seed consumption. We analyzed the effects of no-till agriculture on productivity and income over thirty years (the length of our experiment. We found that yield in no-till increases every year for thirty years, soil moisture remains elevated every year, and that analysis of shorter time periods likely leads to spurious results. When longterm dynamics are accounted for, conversion to no-till becomes profitable after a decade, and benefits accrue for every year after that.

Publications

• Type: Journal Articles Status: Published Year Published: 2019 Citation: Damschen, E.I., Brudvig, L.A., Burt, M.A., Fletcher, R.J., Haddad, N.M., Levey, D.J., Orrock, J.L., Resasco, J. and Tewksbury, J.J., 2019. Ongoing accumulation of plant diversity through habitat connectivity in an 18-year experiment. Science, 365(6460), pp.1478-1480.
• Type: Journal Articles Status: Published Year Published: 2019 Citation: Chaplin-Kramer, R., Sharp, R.P., Weil, C., Bennett, E.M., Pascual, U., Arkema, K.K., Brauman, K.A., Bryant, B.P., Guerry, A.D., Haddad, N.M. and Hamann, M., 2019. Global modeling of natures contributions to people. Science, 366(6462), pp.255-258.
• Type: Journal Articles Status: Published Year Published: 2020 Citation: Henry, E.H., Burford Reiskind, M.O., Land, A.D. and Haddad, N.M., 2020. Maintaining historic disturbance regimes increases species' resilience to catastrophic hurricanes. Global Change Biology, 26(2), pp.798-806.
• Type: Journal Articles Status: Published Year Published: 2020 Citation: Harvey, Jeffrey A., Robin Heinen, Inge Armbrecht, Yves Basset, James H. Baxter-Gilbert, T. Martijn Bezemer, Monika B�hm et al. "International scientists formulate a roadmap for insect conservation and recovery." Nature Ecology & Evolution 4, no. 2 (2020): 174-176.
• Type: Journal Articles Status: Published Year Published: 2020 Citation: Helms IV, J.A., Ijelu, S.E., Wills, B.D., Landis, D.A. and Haddad, N.M., 2020. Ant biodiversity and ecosystem services in bioenergy landscapes. Agriculture, Ecosystems & Environment, 290, p.106780.
• Type: Journal Articles Status: Published Year Published: 2020 Citation: Cusser, S., Bahlai, C., Swinton, S.M., Robertson, G.P. and Haddad, N.M., 2020. Long?term research avoids spurious and misleading trends in sustainability attributes of no?till. Global Change Biology.
• Type: Journal Articles Status: Published Year Published: 2020 Citation: Cusser, S., Pechal, J.L. & Haddad, N.M. Carrion increases pollination service across an urban gradient. Urban Ecosyst (2020). https://doi.org/10.1007/s11252-020-01032-z
• Type: Journal Articles Status: Published Year Published: 2020 Citation: Costanza, Jennifer K., James Watling, Ron Sutherland, Curtis Belyea, Bistra Dilkina, Heather Cayton, David Bucklin, Stephanie S. Roma�ach, and Nick M. Haddad. "Preserving connectivity under climate and land-use change: No one-size-fits-all approach for focal species in similar habitats." Biological Conservation 248 (2020): 108678.

Progress 12/01/18 to 09/30/19

Outputs
Target Audience:My target audience includes other scientists and conservation land managers Changes/Problems: Nothing Reported What opportunities for training and professional development has the project provided?My lab employs sixteen people at various stages of professional development, from undergraduate to postdoctoral. All of them are trained in field biology, including in agricultural, forested, and conservation lands. I employed two undergraduates and four postbac students that assisted with research on prairie strips; three postbac students who did work on rare butterflies; two postdocs who did work on prairie strips; and a research assistant each that did work on corridors, rare butterflies, and website development (ConservationCorridor.org). In nearly all of these positions, the people are in training for the next level, typically a permanent job or graduate school. Thus, in addition to training, I provide professional development in other ways that promote their career goals. How have the results been disseminated to communities of interest?Besides through scientific papers, I maintain a website ConservationCorridor.org that reseaches 50,000 people per year. Associated with my book release, I have given talks at eight science museums. I have been small tours to teachers and stakeholders at my research station. What do you plan to do during the next reporting period to accomplish the goals?I. Corridors We understand the effects of corridors on dispersal and diversity. What we do not know is how corridors affect population processes. In the coming year we will study bee population dynamics and how they are affected by corridors. We will introduce bees that otherwise do not occur in our sites. We will introduce them to one patch, and then test if we can recover them in connected or unconnected habitats. As we now know that corridors increase plant diversity, we will test what affects the stability of plant diversity, including whether the number of species maintains constant over time, or whether weather or fire influence year to year fluctuations II Restoration for butterflies We are collecting detailed data on butterfly demography to create population models and use them to promote recovery of endangered species. To do this, we are observing butterflies at all life stages -- egg, larval, adult -- in treated and control habitats. Further, we will measure dispersal through high and low quality habitat. Then we will create population models to predict when and where restoration will promote butterfly populations. III. Prairie Strips This will be a big year with the experimental strips projects. We will continue with a second year of data collection on effects effects of strips on pollination, decomposition, ant diversity and function, butterfly abundance and diversity, and other responses. Although these things are of interest in their own right, the holy grail is to test for their effects on agricultural yield near strips.

Impacts
What was accomplished under these goals? I. Landscape Corridor In 2019, we made major progress in showing the impacts of reconnecting landscapes where habitats had been lost or fragmented. Working in the largest, best replicated experiment in the world to test the role of landscape corridors, we provided the best demonstration of the long-term impacts of corridors on biodiversity. Specifically, in an 18 year study of plant species in 40, 1ha fragments, some of which were connected by corridors. we found that corridors increased diversity by 14%, and that the effects of corridors are still growing. Further, we discovered the mechanisms by which this occurs. Corridors increase the rate of plant colonization of connected patchess by 5% per year, and they decrease the rate of plant extinction within connected patches by 2% per year. These annual rates accrue over time, causing the higher plant diversity that we observed. Our work is conducted in partnership with the US Forest Service, Savannah River. They assist us in creation and maintanence of our experimental landscapes, and we return our results to aid in their conservation management to optimize biodiversity in working landscapes. II. This was a big year in my work with conservation of rare butterflies, as I published a book on the rarest butterflies in the world. In a synthesis across the world's six rarest species, I discovered how important natural history is to butterfly conservation. I also discovered a common thread that occurs across all the rarest butterflies: we need to kill some butterflies to preserve species. Most rare butterflies live in habitats maintained by disturbance, for example fire. Fire can kill rare butterflies. However, without fire, natural plant succession reduces the presence of important food resources, causing butterfly populations to decline. As natural fires are now prevented, controlled fires are important to conservation. I also discovered the key theme that multiple butterfly populations must be connected. This connects back to habitat: the natural disturbances can cause loss of some populations while others thrive. Thriving populations can then repopulate disturbed areas causing populations to grow again. We also discovered an ironic effect of hurricances. Bartram's scrub hairstreak lives in distrubance-maintained environments in South Florida (maintained by fire). My student created an experiment to test effects of fire. Then, Hurricane Irma struck right over her experiments. It turns out that the plots that were disturbed previously did well when also disturbed by Irma. This was particularly true when the habitat was on the outer ring of the hurricance (high, but not severely high winds). III KBS-LTER and conservation strips This was a big year for our new experiments at KBS-Longterm Ecological Research sites. Our experiment has been running for 30 years. Our funding from the National Science Foundation was renewed, and I assumed directorship. We instituted new treatments within 30 year old experimental plots. One new treatment was Prairie Strips. Much of this year was devoted to establishing Prairie Strips in reduced input and organic treatments. Prairie strips are composed of 22 native species, the types of which were determined by their ability to support pollinators and beneficial insects helpful in controlling pests. We seeds strips and then mowed them regularly to reduce weed abundance. We conducted plant surveys to determine the abundancne and diversity. Then we conducted many first year studies, of pollination, of decomposition by beetles, of ants diversity and abundance, and more. Results are still coming in.

Publications

• Type: Journal Articles Status: Published Year Published: 2019 Citation: Helms, J.A., Ijelu, S.E. and Haddad, N.M., 2019. Range expansion in an introduced social parasite-host species pair. Biological Invasions, 21:2751-2759.
• Type: Books Status: Published Year Published: 2019 Citation: Haddad, N.M. The Last Butterflies: A Scientists Quest to Save a Rare and Vanishing Creature. Princeton University Press.
• Type: Journal Articles Status: Published Year Published: 2019 Citation: Wepprich T, Adrion JR, Ries L, Wiedmann J, Haddad NM (2019) Butterfly abundance declines over 20 years of systematic monitoring in Ohio, USA. PLoS ONE 14(7): e0216270
• Type: Journal Articles Status: Published Year Published: 2019 Citation: Schultz, C. B., N. M. Haddad, E. H. Henry, and E. E. Crone. 2019. Movement and Demography of At-Risk Butterflies: Building Blocks for Conservation. Annual Review of Entomology 64:167-184 doi:10.1146/annurev-ento-011118-112204
• Type: Journal Articles Status: Published Year Published: 2019 Citation: Caughlin, T. T., E. I. Damschen, N. M. Haddad, D. J. Levey, C. Warneke, and L. A. Brudvig. 2019. Landscape heterogeneity is key to forecasting outcomes of plant reintroduction. Ecological Applications 29(2):e01850. doi:10.1002/eap.1850
• Type: Journal Articles Status: Published Year Published: 2019 Citation: Henry, E., E. Brammer-Robbins, E. Aschehoug, and N.M. Haddad. 2019. Do substitute species help or hinder endangered species management? Biological Conservation 232:127-130