Earthworms in the diet of Herring Gulls Larus argentatus breeding on an off-shore island

ABSTRACT

Pellets of indigestible material regurgitated by Herring Gulls Larus argentatus breeding on Lady Isle, Firth of Clyde, Scotland, were collected in 2018 and 2019 and examined for earthworm chaetae. Nearly two-thirds (65.6%) of the 314 pellets came from gulls that had consumed one or more earthworms. Significantly fewer pellets (57.6%) contained chaetae in 2018, a relatively dry May to July, than in 2019 (72.0%) when rainfall was close to the long-term average for May to July. There were significant associations between the presence of large quantities of terrestrial vegetation in the pellet and the detection of large numbers of earthworm chaetae and/or fragments of terrestrial arthropods, suggesting that recent consumption of these food items is likely when pellets contain large quantities of vegetation.     

White-tailed deer (Odocoileus virginianus) fecal pellet decomposition is accelerated by the invasive earthworm Lumbricus terrestris

Exotic European earthworms have expanded into worm-free forests of the United States. Concurrently, populations of the white-tailed deer, Odocoileus virginianus, have also increased. During winter, deer use hemlock stands for cover while browsing elsewhere, creating a net organic matter flux into these stands. Deer fecal pellets can provide annual inputs of 48.1 kg C, 1.4 kg N, and 1.3 kg Ca per hectare. We tested the hypothesis that these pellets were readily consumed by invading earthworms. The litter-feeding anecic earthworm Lumbricus terrestris redistributed fecal pellets and accelerated mass and nutrient loss rates. These losses are likely due to the combination of enhanced fragmentation and decomposition as earthworms drag pellets into their burrows for consumption. This nutrient subsidy may be an important source of high quality “litter” input to hemlock stands, which may in turn facilitate the invasion of these stands by earthworms under high deer densities.     

Determination of the diet of the stone curlew (Bwhinus oedicnemus) by faecal analysis

The value of faecal analysis for estimating the composition of the diet of stone curlews was assessed by examination of the faeces of a captive bird fed on a measured diet. Remains of soft–bodied prey were under–represented in the faeces but the results could be adjusted to allow for this. Estimation of the size of prey was possible by measurement of remnants such as earthworm chaetae and insect mandibles. In some types of prey large specimens left more remnants than small ones and a correction was devised to allow for this. Regurgitated pellets contained large, hard remains and there was no trace of some important prey types such as earthworms. A method is described for estimating the composition of the diet of the wild stone curlews from counts of prey remains in their faeces.     

Do non-native earthworms in Southeast Alaska use streams as invasional corridors in watersheds harvested for timber?

Exotic earthworms from Europe and Asia have invaded previously earthworm-free areas of North America where they consume leaf litter, mix soil horizons, and alter nutrient cycling. Primarily, earthworm introductions occur through human activities; we hypothesized that the combination of logging (i.e., road construction and soil disturbance) and stream transport (i.e., hydrochory) allows earthworms to invade new ecosystems and spread within watersheds. On Prince of Wales Island, AK, we surveyed riparian zones in 11 watersheds with varying timber harvest intensity for terrestrial oligochaetes. Additionally, common invasive earthworms were experimentally submerged in a local stream to test for tolerance to prolonged immersion: all taxa survived immersion for at least 6 days. Using principal components analysis, watershed and harvest variables describing the watersheds upstream of our sampled riparian areas were reduced to two principal components describing harvest intensity (PC1) and harvest style (PC2). Logistic models successfully predicted earthworm abundance (r ²  = 0.70) from PC1, which indicated that watersheds with older, intense upstream timber harvest contained larger earthworm populations. Earthworm species richness was best predicted by PC2 (r ²  = 0.39), which suggested that earthworm communities in watersheds containing large clear-cut stands were more species-rich. Collectively, these results suggest that (1) invasive earthworms may use streams for dispersal and (2) upstream introductions via timber harvest can initiate downstream earthworm invasions. Hydrochory would allow invasive earthworms to spread at rates (tens of km d⁻¹) that are much greater than previously reported rates of terrestrial spread (5–10 m y⁻¹). Effective control of exotic earthworms in riparian zones will require watershed-level management and surveillance.     

Conventional tillage decreases the abundance and biomass of earthworms and alters their community structure in a global meta‐analysis

The adoption of less intensive soil cultivation practices is expected to increase earthworm populations and their contributions to ecosystem functioning. However, conflicting results have been reported on the effects of tillage intensity on earthworm populations, attributed in narrative reviews to site‐dependent differences in soil properties, climatic conditions and agronomic operations (e.g. fertilization, residue management and chemical crop protection). We present a quantitative review based on a global meta‐analysis, using paired observations from 165 publications performed over 65 years (1950–2016) across 40 countries on five continents, to elucidate this long‐standing unresolved issue. Results showed that disturbing the soil less (e.g. no‐tillage and conservation agriculture [CA]) significantly increased earthworm abundance (mean increase of 137% and 127%, respectively) and biomass (196% and 101%, respectively) compared to when the soil is inverted by conventional ploughing. Earthworm population responses were more pronounced when the soil had been under reduced tillage (RT) for a long time (>10 years), in warm temperate zones with fine‐textured soils, and in soils with higher clay contents (>35%) and low pH (<5.5). Furthermore, retaining organic harvest residues amplified this positive response to RT, whereas the use of the herbicide glyphosate did not significantly affect earthworm population responses to RT. Additional meta‐analyses confirmed that epigeic and, more importantly, the bigger‐sized anecic earthworms were the most sensitive ecological groups to conventional tillage. In particular, the deep burrower Lumbricus terrestris exhibited the strongest positive response to RT, increasing in abundance by 124% more than the overall mean of all 13 species analysed individually. The restoration of these two important ecological groups of earthworms and their burrowing, feeding and casting activities under various forms of RT will ensure the provision of ecosystem functions such as soil structure maintenance and nutrient cycling by “nature's plough.”     

Exotic Ecosystem Engineers Change the Emergence of Plants from the Seed Bank of a Deciduous Forest

The anthropogenic spread of exotic ecosystem engineers profoundly impacts native ecosystems. Exotic earthworms were shown to alter plant community composition of the understory of deciduous forests previously devoid of earthworms. We investigated the effect of two exotic earthworm species (Lumbricus terrestris L. and Octolasion tyrtaeum Savigny) belonging to different ecological groups (anecic and endogeic) on the emergence of plants from the seed bank of a northern North American deciduous forest using the seedling emergence method. We hypothesized that (1) exotic earthworms change the seedling emergence from the plant seed bank, (2) L. terrestris increases the emergence of plant seedlings of the deeper soil layer but decreases that of the upper soil layer due to plant seed burial, and (3) O. tyrtaeum decreases plant seedling emergence due the damage of plant seeds. Indeed, exotic earthworms altered the emergence of plant seedlings from the seed bank and the functional composition of the established plant seedlings. Surprisingly, although L. terrestris only marginally affected seedling emergence, O. tyrtaeum changed the emergence of native plant species from the seed bank considerably. In particular, the number of emerging grass and herb seedlings were increased in the presence of O. tyrtaeum in both soil layers. Moreover, the impacts of earthworms depended on the identity of plant functional groups; herb species benefited, whereas legumes suffered from the presence of exotic earthworms. The results highlight the strong effect of invasive belowground ecosystem engineers on aboveground ecosystem characteristics and suggest fundamental changes of ecosystems by human-spread earthworm species.     

Influence of driver ant swarm raids on earthworm prey densities in the Mount Kenya forest: implications for prey population dynamics and colony migrations

African driver ants are nomadic social mesopredators feeding on a highly diverse array of prey species at different trophic levels. Colonies of certain driver ant species have a biomass which can equal that of medium-sized mammalian carnivores and the ultimate cause of their nomadic life-style is thought to be local prey depletion. The impact of driver ant swarm raids is therefore expected to be strong but the degree to which they reduce prey populations has not been quantified and it is unknown whether these spectacular predators exert significant top-down effects. We examined the combined effect of driver ant (Dorylus molestus) and swarm-attending bird (Alethe poliocephala) predation on the population dynamics of earthworms, which constitute the ants’ main prey type in the montane forest of Mount Kenya. Pre-raid earthworm biomass densities in the soil layer down to a depth of 8 cm varied by a factor of 31. The immediate effect of swarm raids was a reduction in earthworm numbers in this layer, but 8 days later earthworm numbers had recovered to pre-raid levels. When earthworm biomass densities were compared, no significant effect of swarm raids was detected. The estimated proportion of earthworm prey biomass extracted from 0 to 8 cm layer by driver ants and birds together was about 2.2%. Although colony distribution was overdispersed as expected based on knowledge of D. molestus migratory behaviour, predation events were highly localized. Predation frequency was low (once every 62 days on average) and highly variable. These results indicate that earthworm prey is highly abundant but at the same time so difficult to harvest that swarm raids exert only a marginal influence on earthworm populations. Longer-term studies would be required to determine whether earthworm populations are limited by swarm raids. The small impacts of individual raids and rapid recovery of earthworm prey populations likely underlie the low frequency of migrations and short distances travelled by migrating colonies of D. molestus.     

Are non-migrant white storks (Ciconia ciconia) able to survive a cold-induced fast?

Northwestern European populations of White Storks (Ciconia ciconia) have been restored by settling young birds which, after having been maintained captive for 3 years, do not migrate when released. Since they are still supplied with food, the question we address here is how settled White Storks would cope with a fast resulting from a cold spell if food were no longer available. We therefore measured daily body mass loss, nitrogen excretion and daily energy expenditure under natural photoperiod and climatic conditions in six captive White Storks exposed to 5 days of food deprivation during winter. Daily ambient temperature ranged between -10 and 17 degrees C. After an initial decrease in both daily body mass loss and nitrogen excretion during the first day of fasting, these two parameters stabilized at low values. Lipids were the main fuel energy source (91%) and body proteins accounted for the remainder (9%). The rate of CO(2) production, measured with the doubly labeled water method, decreased significantly (P<0.001) during fasting when compared to the ad libitum value. Our data show that the metabolic adjustments of White Storks facing starvation are similar to those found in birds well-adapted to long-term fasting. From the determination of lipid and protein stores and of lipid and protein depletion rates, we conclude that White Storks would be able to survive at least during 4 weeks, a period of time twice as long as the duration of cold spells usually occurring in the Alsace region.     

The magnitude and timing of migration by soaring raptors, pelicans and storks over Israel

The magnitude and timing of the autumn and spring migrations of 35 species of medium-and large-sized raptors, White Pelicans Pelicanus onocrotalus and White Storks Ciconia ciconia were studied in Israel. Observations were carried out from the ground by a line of observers covering most of the width of Israel across the line of migration and by radar. There was a high correlation between the counts obtained by ground observers and by radar. On average, about half a million raptors (mainly Lesser Spotted Eagles Aquila po-marina, Honey Buzzards Pernis apivorus and Levant Sparrowhawks Accipiter brevipes), 250,000 White Storks and 70,000 White Pelicans passed during autumn, and about a million raptors (mainly Honey Buzzards, Steppe Buzzards Buteo vulpinus, Steppe Eagles Aquila nipalensis and Black Kites Milvus migrans) and 450,000 White Storks passed during spring. Peak numbers were higher–over a million raptors and half a million White Storks. There was high interyear variation in the number of migrants recorded during the study, probably caused by weather and counting efforts. For some species, the whole world (Lesser Spotted Eagle and Levant Sparrowhawk) or Palaearctic (White Pelican) population passes over Israel during migration, allowing an estimate of the world populations of these species. Mean dates of arrival of most raptors are highly predictable, with confidence limits ranging between 1.5 and 5.5 days. The migration periods of White Storks and White Pelicans are longer and their mean day of appearance is less predictable (confidence limits range from 4.2 to 13.8 days). During autumn, 90% of the migrating populations of nocking species, such as Levant Sparrowhawk, Lesser Spotted Eagle, Honey Buzzard and Red-footed Falcon Falco vespertinus, pass within 13, 15, 16 and 18 days, respectively, while nonflocking species, such as Egyptian Vulture Neophron percnopterus, Marsh Harrier Circus aeruginosus and Short-toed Eagle Circaetus gallicus, generally take twice as long to pass. Similar passage periods were recorded in spring. For most species, the autumn migration period was longer than the spring migration period, probably because in autumn adults move before the young birds. Three factors affected the timing and spread of the migration wave: age at first breeding, diet and size of the breeding area.     

White Storks, <Emphasis Type="Italic">Ciconia ciconia</Emphasis>, forage on rubbish dumps in Poland—a novel behaviour in population

Information on the foraging of White Storks on rubbish dumps, a novel behaviour in Central European populations, is presented. Observations were first made in 1999 and to date; they have been recorded on sixteen locations in Poland. From one to 348 White Storks (median = 2, N = 116) were observed on rubbish dumps, and most of the records (86%) were of 1–3 birds. Birds foraging on rubbish dumps were recorded from late March to early September, but the highest numbers were seen during the summer months. Most of the birds (77%, N = 171) were recorded foraging directly on an area where rubbish had been thrown, while a minor fraction (16%) was seen on neighbouring recultivated grassy areas. Dumps were used as an additional food source probably in areas where natural foraging grounds are limited. The increased frequency of observations during the summer months probably resulted from the greater nutritional needs of large nestlings which force the adults to find alternative food sources or foraging habitat shift by non-breeding birds.     

The effect of anthropogenic arsenic contamination on the earthworm microbiome

Earthworms are globally distributed and perform essential roles for soil health and microbial structure. We have investigated the effect of an anthropogenic contamination gradient on the bacterial community of the keystone ecological species Lumbricus rubellus through utilizing 16S rRNA pyrosequencing for the first time to establish the microbiome of the host and surrounding soil. The earthworm‐associated microbiome differs from the surrounding environment which appears to be a result of both filtering and stimulation likely linked to the altered environment associated with the gut micro‐habitat (neutral pH, anoxia and increased carbon substrates). We identified a core earthworm community comprising Proteobacteria (～50%) and Actinobacteria (～30%), with lower abundances of Bacteroidetes (～6%) and Acidobacteria (～3%). In addition to the known earthworm symbiont (Verminephrobacter sp.), we identified a potential host‐associated Gammaproteobacteria species (Serratia sp.) that was absent from soil yet observed in most earthworms. Although a distinct bacterial community defines these earthworms, clear family‐ and species‐level modification were observed along an arsenic and iron contamination gradient. Several taxa observed in uncontaminated control microbiomes are suppressed by metal/metalloid field exposure, including eradication of the hereto ubiquitously associated Verminephrobacter symbiont, which raises implications to its functional role in the earthworm microbiome.     

The soil fauna as a food source for moles

Moles feed mainly on insects and earthworms and may eat 18 to 36 k (40 to 80 lb) per annum, collected in an area of about 1/25 hectare (1/10 acre). This largely restricts them to deciduous woodland and old grassland on "mull" soils since field populations of soilinsects cannot provide them with enough food but earthworm populations can. Slugs and millipedes may be relatively unpalatable.
Caches of earthworms stored by moles consist almost entirely of Lumbricus terrestris immobilized by removal or multilation of the anterior 3–5 segments. It is still uncertainhow and when the caches are made. Earthworm cocoons are prominent in the diet of moles in pastures but how they are located is not yet known.     

Tree leaf litter composition and nonnative earthworms influence plant invasion in experimental forest floor mesocosms

Dominant tree species influence community and ecosystem components through the quantity and quality of their litter. Effects of litter may be modified by activity of ecosystem engineers such as earthworms. We examined the interacting effects of forest litter type and earthworm presence on invasibility of plants into forest floor environments using a greenhouse mesocosm experiment. We crossed five litter treatments mimicking historic and predicted changes in dominant tree composition with a treatment of either the absence or presence of nonnative earthworms. We measured mass loss of each litter type and growth of a model nonnative plant species (Festuca arundinacea, fescue) sown into each mesocosm. Mass loss was greater for litter of tree species characterized by lower C:N ratios. Earthworms enhanced litter mass loss, but only for species with lower C:N, leading to a significant litter × earthworm interaction. Fescue biomass was significantly greater in treatments with litter of low C:N and greater mass loss, suggesting that rapid decomposition of forest litter may be more favorable to understory plant invasions. Earthworms were expected to enhance invasion by increasing mass loss and removing the physical barrier of litter. However, earthworms typically reduced invasion success but not under invasive tree litter where the presence of earthworms facilitated invasion success compared to other litter treatments where earthworms were present. We conclude that past and predicted future shifts in dominant tree species may influence forest understory invasibility. The presence of nonnative earthworms may either suppress of facilitate invasibility depending on the species of dominant overstory tree species and the litter layers they produce.     

Capturing American White Ibises in urban South Florida using two novel techniques

As many wildlife species, including wading birds, adapt to anthropogenic landscapes and, in some cases, exhibit altered behaviors, studies that involve capturing birds may require new methods better suited for use in urban areas and to accommodate altered animal behavior. We developed two novel techniques, a leg lasso and flip net, for capturing American White Ibises (Eudocimus albus) in urban environments in southern Florida, and also used a traditional technique (mist‐nets) in non‐urban wetland habitats. The flip net and leg lasso were developed to capture White Ibises habituated to the presence of humans. Ibises were captured in urban and wetland environments from October 2015 to August 2017 in Palm Beach, Broward, and Lee counties, Florida. We captured 6.0 ± 13.5 ibis/h with the flip net, 1.6 ± 0.8 ibis/h with the leg lasso, and 0.5 ± 2.6 ibis/h with mist‐nets. We captured larger (higher mass to tarsus length ratio) birds using the flip net and leg lasso than using mist‐nets, and captured more males with leg lassos than with other two techniques. The novel techniques we used are efficient, cost effective, easy to use, and also potentially useful for capturing other species of birds. Leg lassos and flip nets are also safe to use in populated areas for both birds and humans.     

Invasion by a non‐native ecosystem engineer alters distribution of a native predator

Aim

Shifts in diet composition, abundance or distribution of native predators can occur as a result of exotic prey introductions. We examined effects of non‐native earthworms and anthropogenic landscape disturbance on habitat selection by the American robin (Turdus migratorius), a generalist predator, at landscape and local levels. We also investigated whether robins could act as vectors of spread for earthworm cocoons (egg cases).

Location

Boreal forest of Alberta, Canada.

Methods

We conducted robin and earthworm surveys at campgrounds, well pads, roads, pipelines, seismic lines and forest interiors across northern Alberta. At a subset of paired locations that had similar habitats and anthropogenic disturbance levels, we sampled both robins and earthworms.

Results

Both groups were most likely to occur at campgrounds, well pads and roads. Furthermore, robins were more likely to occur at locations where earthworms were present in our paired local‐level surveys. This correlation between robin and earthworm distributions could be due to robins acting as a vector for earthworm spread, rather than robins’ use of earthworms as prey. However, in tests using captive robins, earthworm cocoons did not survive digestion.

Main conclusions

Robin and earthworm distributions were correlated, likely due to robins’ use of earthworms as prey. These results suggest exotic prey can strongly influence native predators at both landscape and local levels, with shifts in native predator distributions occurring as a result of spatial variability in exotic prey distributions. Although the impacts of ecosystem engineering by earthworms have been previously demonstrated, our study provides evidence that effects of earthworms can also cascade upwards via trophic interactions.     

两种代表性蚯蚓对设施菜地土壤微生物群落结构及理化性质的影响

不同生态型蚯蚓的取食偏好和生境有所差异,因此蚯蚓的生态型差异可能关乎其对土壤性质的不同影响;有关不同生态型蚯蚓对土壤性质尤其是微生物学性质影响的研究有助于了解蚯蚓生态功能的作用机制。在野外调控试验的第4年采集土壤,研究了牛粪混施和表施处理下内层种威廉腔环蚓(Metaphire guillelmi)和表层种赤子爱胜蚓(Eisenia foetida)对设施菜地土壤微生物群落结构和主要理化性质的影响。结果表明,土壤微生物群落结构同时受到蚯蚓种类和牛粪施用方式的影响。牛粪表施时,两种蚯蚓均显著降低了菌根真菌、真菌生物量和原生动物生物量(P0.05);牛粪混施时,不同蚯蚓的影响有所差异,威廉腔环蚓明显增加了菌根真菌、真菌生物量和放线菌生物量,而赤子爱胜蚓的作用不明显。此外,两种蚯蚓均提高了土壤孔隙度、团聚体稳定性和土壤p H、矿质氮以及微生物生物量碳氮水平,但提高幅度取决于蚯蚓种类和牛粪施用方式。冗余分析表明蚯蚓影响下土壤微生物群落结构的变化与团聚体稳定性、p H、速效磷、矿质氮呈正相关,而与土壤容重呈负相关。     

Nest mounds of red wood ants (Formicaaquilonia): hot spots for litter-dwelling earthworms

A previously undocumented association between earthworms and red wood ants (Formicaaquilonia Yarr.) was found during an investigation of the influence of wood ants on the distribution and abundance of soil animals in boreal forest soil. Ant nest mounds and the surrounding soil of the ant territories were sampled. The ant nest mound surface (the uppermost 5-cm layer) harboured a much more abundant earthworm community than the surrounding soil; the biomass of the earthworms was about 7 times higher in the nests than in the soil. Dendrodrilusrubidus dominated the earthworm community in the nests, while in soils Dendrobaenaoctaedra was more abundant. Favorable temperature, moisture and pH (Ca content), together with abundant food supply (microbes and decomposing litter) are likely to make a nest mound a preferred habitat for earthworms, provided that they are not preyed upon by the ants. We also conducted laboratory experiments to study antipredation mechanisms of earthworms against ants. The experiments showed that earthworms do not escape predation by avoiding contact with ants in their nests. The earthworm mucus repelled the ants, suggesting a chemical defence against predation. Earthworms probably prevent the nest mounds from becoming overgrown by moulds and fungi, indicating possible mutualistic relationships between the earthworms and the ants. Received: 21 November 1996 / Accepted: 3 April 1997     

Nonlinearity of effects of invasive ecosystem engineers on abiotic soil properties and soil biota

Invasions of non‐indigenous species into natural communities are currently rated as one of the most important threats to biodiversity. Particularly exotic ecosystem engineers such as earthworms potentially have profound impacts on community assembly and functioning. We investigated the impact of invasion by the lumbricid earthworms into an aspen forest of the Canadian Rocky Mountains on soil organic matter, microorganisms and microarthropod communities. Building on the results of previous studies in this forest, we expected positive effects of Lumbricus terrestris middens and negative effects of Octolasion tyrtaeum on soil biota (increase and decrease in soil nutrient concentrations, microbial parameters and soil microarthropod density and diversity, respectively). Further, we expected that earthworm effects change with time. Combined results of previous and the present study suggest a wavelike colonization pattern for Dendrobaena octaedra and O. tyrtaeum and that indeed the impact of earthworms on soil biota changed with time, likely due to changes in earthworm density. Unexpectedly, L. terrestris middens neither affected soil abiotic nor soil biotic properties. By contrast and in contrast to our hypothesis, carbon and nitrogen concentration and C‐to‐N ratio in deeper soil layers increased in presence of O. tyrtaeum, thereby likely enhancing nutrient availability for soil microorganisms and microarthropods. Even though the density of this endogeic species was rather low, presence of O. tyrtaeum resulted in increased densities of a number of microarthropod taxa and increased microarthropod diversity. The results suggest that at low density, invasive ecosystem engineers, such as O. tyrtaeum, cause disturbances of intermediate strength thereby beneficially affecting soil microorganisms and most microarthropods. This contrasts earlier effects during the wavelike invasion of O. tyrtaeum into the aspen forest when densities of O. tyrtaeum were high resulting in generally detrimental effects on soil biota. The results emphasize the nonlinearity of earthworm effects on abiotic and biotic soil properties and call for further long‐term investigations.     

Distribution and at‐sea behavior of Bermudan White‐tailed Tropicbirds (Phaethon lepturus catesbyi) during the non‐breeding season

The movements and behavior of many taxa of seabirds during the non‐breeding season remain poorly known. For example, although studies conducted in the Pacific and Indian oceans suggest that White‐tailed Tropicbirds (Phaethon lepturus) seldom fly more than a few thousand kilometers from nest colonies after breeding, little is known about the post‐breeding movements and behavior of a subspecies of White‐tailed Tropicbirds (P. l. catesbyi) that breeds on islands in the North Atlantic Ocean. Our objective, therefore, was to use light‐based geolocators to identify the ranges and pelagic activities of White‐tailed Tropicbirds from Bermuda during the non‐breeding periods in 2014–2015 (N = 25) and 2015–2016 (N = 16). Locations were estimated based on changes in light intensity across time, and pelagic activities were determined based on whether geolocators attached to leg bands were wet (i.e., birds resting on the water's surface) or dry (i.e., birds in flight). In 2014, birds spent late summer (July–September) near Bermuda and the British Virgin Islands; by mid‐September, most (N = 17; 68%) birds took a direct easterly route to the Sargasso Sea. In 2015, most post‐breeders (N = 15; 94%) flew east from Bermuda and to the Sargasso before the end of late summer. For both years combined, fall and winter (October–February) ranges extended as far west as North Carolina and as far east as the mid‐Atlantic Ridge. In both years, all birds were located between Bermuda and the British Virgin Islands during the spring (April–May). All birds then flew north to Bermuda in both years, with variations in timing, during April and May. We also found extensive overlap in the ranges of males and females during the non‐breeding season in both years. During the non‐breeding season, White‐tailed Tropicbirds spent 5% of night periods and 41% of day periods in flight in 2014; in 2015, birds spent 8% and 42% of night and day periods, respectively, in flight. Tropicbirds spent more time flying during the day because they hunt by day, detecting prey on the wing by sight. Overall, our results suggest that White‐tailed Tropicbirds that breed in Bermuda are diurnal, nomadic wanderers that range over an extensive area of the Atlantic Ocean during the non‐breeding season.     

Earthworm invasions of ecosystems devoid of earthworms: effects on soil microbes

Recent studies document North American earthworm invasions and their profound effects on the structure of the soil profile, which is the habitat for soil microorganisms (mainly fungi and bacteria). Dramatic alterations made to these layers during earthworm invasion significantly change microbial community structure and therefore microbial activities such as C transformations. Understanding the impacts of earthworm invasion on the microbes themselves will give insight into earthworm effects on microbial activities. Bacterial and actinomycete communities in earthworm guts and casts have not been studied in environments recently invaded by earthworms. Earthworm invasion tended to decrease fungal species density and fungal species diversity and richness. The presence of earthworms decreased zygomycete species abundance probably due to disruption of fungal hyphae. Physical disruption of hyphae may also explain decreased mycorrhizal colonization rates, decreased mycorrhizal abundance and altered mycorrhizal morphology in the presence of earthworms. Mixing of organic layers into mineral soil during earthworm invasion tended to decrease microbial biomass in forest floor materials while increasing it in mineral soil. In newly invaded forest soils, microbial respiration and the metabolic quotient tended to decline. In forests where either the microbial community has had time to adapt to earthworm activities, or where the destruction of the forest floor is complete, as in invasions by the Asian Amynthas hawayanus, the presence of earthworms tends to increase the metabolic quotient indicating a shift to a smaller, more active microbial community.