Disproportionate declines of formerly abundant species underlie insect loss.

Studies have reported widespread declines in terrestrial insect abundances in recent years1-4, but trends in other biodiversity metrics are less clear-cut5-7. Here we examined long-term trends in 923 terrestrial insect assemblages monitored in 106 studies, and found concomitant declines in abundance and species richness. For studies that were resolved to species level (551 sites in 57 studies), we observed a decline in the number of initially abundant species through time, but not in the number of very rare species. At the population level, we found that species that were most abundant at the start of the time series showed the strongest average declines (corrected for regression-to-the-mean effects). Rarer species were, on average, also declining, but these were offset by increases of other species. Our results suggest that the observed decreases in total insect abundance2 can mostly be explained by widespread declines of formerly abundant species. This counters the common narrative that biodiversity loss is mostly characterized by declines of rare species8,9. Although our results suggest that fundamental changes are occurring in insect assemblages, it is important to recognize that they represent only trends from those locations for which sufficient long-term data are available. Nevertheless, given the importance of abundant species in ecosystems10, their general declines are likely to have broad repercussions for food webs and ecosystem functioning.

InsectChange: a global database of temporal changes in insect and arachnid assemblages.

Insects are the most ubiquitous and diverse group of eukaryotic organisms on Earth, forming a crucial link in terrestrial and freshwater food webs. They have recently become the subject of headlines because of observations of dramatic declines in some places. Although there are hundreds of long-term insect monitoring programs, a global database for long-term data on insect assemblages has so far remained unavailable. In order to facilitate synthetic analyses of insect abundance changes, we compiled a database of long-term (≥10 yr) studies of assemblages of insects (many also including arachnids) in the terrestrial and freshwater realms. We searched the scientific literature and public repositories for data on insect and arachnid monitoring using standardized protocols over a time span of 10 yr or longer, with at least two sampling events. We focused on studies that presented or allowed calculation of total community abundance or biomass. We extracted data from tables, figures, and appendices, and, for data sets that provided raw data, we standardized trapping effort over space and time when necessary. For each site, we extracted provenance details (such as country, state, and continent) as well as information on protection status, land use, and climatic details from publicly available GIS sources. In all, the database contains 1,668 plot-level time series sourced from 165 studies with samples collected between 1925 and 2018. Sixteen data sets provided here were previously unpublished. Studies were separated into those collected in the terrestrial realm (103 studies with a total of 1,053 plots) and those collected in the freshwater realm (62 studies with 615 plots). Most studies were from Europe (48%) and North America (29%), with 34% of the plots located in protected areas. The median monitoring time span was 19 yr, with 12 sampling years. The number of individuals was reported in 129 studies, the total biomass was reported in 13 studies, and both abundance and biomass were reported in 23 studies. This data set is published under a CC-BY license, requiring attribution of the data source. Please cite this paper if the data are used in publications, and respect the licenses of the original sources when using (part of) their data as detailed in Metadata S1: Table 1.

Complex Messages in Long-Term Monitoring of Regal Fritillary (Speyeria idalia) (Lepidoptera: Nymphalidae) in the State of Wisconsin, USA, 1988-2015.

The regal fritillary ("regal") (Speyeria idalia) is endangered in Wisconsin, USA, and declining and at risk range-wide. During 1988-2015, we surveyed 24 known regal sites and >100 areas of potential habitat in Wisconsin. We recorded 9037 individuals in 742.7 km on the peak survey per year at occupied sites. At six sites surveyed over 5-25 years, we found regal fritillaries in only one year, mostly in the latter half of the study. The three populations in the state with more favorable trends than the median had a never-burned refugium and/or infrequent fire management. They also all had substantial amounts of grazing, haying, and/or mowing managements. Sites with trends below the regional median trend had frequent or moderate fire management, and either a diminishing never-burned refugium or none at all. Regal populations at sites with ≤15 ha of grassland have become undetectable. Nonetheless, Hogback, a slightly larger than 15 ha site, had the most favorable trend, a significant increase. Nearly all Wisconsin Regal populations known before 1990 declined to consistent non-findability, even though these were conserved sites. More favorable trends at more recently discovered populations may be attributable to species-specific habitat management protocols implemented in the 1990s. Two sites with better than median long-term trends represent the longest consistent land ownership of known Regal populations in the state. This wide range of population outcomes illustrates both the need for long-term monitoring and the challenges of explaining the outcomes. Despite evidence of increasing Regal dispersal, this species remains very localized, indicating the unsuitability of the wider landscape as regal habitat. The number of significantly declining or no longer detectable populations in Wisconsin indicates an ever more adverse landscape for this species. Sites will need to have habitat characteristics that are ever more optimal in a wide range of climatic conditions for Regal populations to persist.

Status and Trend of Regal Fritillary (Speyeria idalia) (Lepidoptera: Nymphalidae) in the 4th of July Butterfly Count Program in 1977-2014.

Regal Fritillary (Speyeria idalia) primarily inhabits prairie, a native grassland of central North America, and occurs rarely in nonprairie grasslands further east. This butterfly has experienced widespread decline and marked range contraction. We analyze Regal Fritillary incidence and abundance during 1977-2014 in 4th of July Butterfly Counts, an annual census of butterflies in North America. Volunteers count within the same 24 km diameter circle each year. Only 6% of counts in range reported a Regal, while 18% of counts in core range in the Midwest and Great Plains did. 99.9% of Regal individuals occurred in core range. Only four circles east of core range reported this species, and only during the first half of the study period. All individuals reported west of its main range occurred in two circles in Colorado in the second half of the study. The number of counts per year and survey effort per count increased during the study. During 1991-2014, >31 counts occurred per year in core Regal range, compared to 0-23 during 1975-1990. During 1991-2014, all measures of Regal presence and abundance declined, most significantly. These results agree with other sources that Regal Fritillary has contracted its range and declined in abundance.

Twenty Years of Elfin Enumeration: Abundance Patterns of Five Species of Callophrys (Lycaenidae) in Central Wisconsin, USA.

We recorded five species of elfins (Callophrys) during annual spring surveys targeting frosted elfin C. irus (state-listed as threatened) in 19 pine-oak barrens in central Wisconsin USA during 1994-2013. At the northwest end of its range here, C. irus co-varied with spring temperature, but declined significantly over time (eight sites verified extant of originally 17). Two other specialists increased significantly. The northern specialist, hoary elfin C. polios (nine sites), correlated positively with the previous year's growing season precipitation. The southern specialist, Henry's elfin C. henrici (11 sites), co-varied with winter precipitation and spring temperature and dryness. The two resident generalists had stable trends. For all species, the first observed date per year became earlier over time and varied more than the last observed date. Thus, flight period span increased with earlier first observed dates. Elfin abundance increased significantly with earlier first observed dates in the current and/or prior year. Three species (C. irus, C. henrici, a generalist) had more positive population trends in reserves than non-reserves. This suggests that C. irus declines correspond to habitat conditions. Thus, monitoring programs and habitat management specifically for C. irus appear necessary to obtain a long-term stable trend for this species in Wisconsin.

Decline of Hesperia ottoe (Lepidoptera: Hesperiidae) in Northern Tallgrass Prairie Preserves.

We counted butterflies on transect surveys during Hesperia ottoe flight period in 1988-2011 at tallgrass prairie preserves in four states (Illinois, Iowa, Minnesota, Wisconsin), divided into units cross-referenced to vegetation type and management history. H. ottoe occurred only in dry and sand prairie types, and was significantly more abundant in undegraded than semi-degraded prairie, and in discontinuous sod (with numerous unvegetated areas due to bare sand and/or rock outcrops) than in continuous sod. This skipper was significantly more abundant in small sites compared to medium and large sites, even when the analysis was limited to undegraded prairie analyzed separately by sod type. H. ottoe was significantly under-represented in year-burn 0 (the first growing season after fire) compared to an expected distribution proportional to survey effort. However, H. ottoe was also over-represented in fire-managed units compared to non-fire-managed units. However, by far most units and sites were in fire management and most populations declined to subdetection during this study. Peak abundance post-fire occurred in a later year-burn in discontinuous sod and was much higher than in continuous sod. We also analyze H. ottoe status and trend in midwestern prairie preserves by compiling a dataset of our and others' butterfly surveys from 1974 to 2011. Only 1/9 sites with continuous sod had detectable H. ottoe in recent year(s). In discontinuous sod, 2/6 did, with two sites lacking data for the last few years. The number of years H. ottoe was still detectable after preservation and the number of years to consistent non-detection were both significantly higher in discontinuous than continuous sod. Both measures of population persistence averaged over twice as long in discontinuous than continuous sod, and correlated negatively with prairie size. The year when consistent non-detection began varied over several decades among sites. Despite the currently urgent need to identify how to manage preserves successfully for H. ottoe, such research now needs to be very cautious, because of the extreme fragility of the few remaining populations and the ruggedness of the preserves where H. ottoe is still known to occur.