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Bats play crucial ecosystem services as seed dispersers, pollinators, controllers of insects, and nutrient recyclers. However, there has not been a thorough global review evaluating these roles in bats across all biogeographical regions of the world. We reviewed the literature published during the last two decades and identified 283 relevant studies: 78 dealt with the control of potential insect pests by bats, 80 related to the suppression of other arthropods, 60 on the dispersal of native or endemic seeds, 11 dealt with the dispersal of seeds of introduced plants, 29 on the pollination of native or endemic plants, 1 study on pollination of introduced plants, and 24 on the use of guano as fertilizer. Our literature search showed that queries combining the terms "seed dispersal," "insectivorous bats," "nectarivorous bats," "use of guano," and "ecosystem services" returned 577 studies, but half were experimental in nature. We found that the evaluation of ecosystem services by bats has been mostly conducted in the Neotropical and Palearctic regions. To detect differences across relevant studies, and to explain trends in the study of ecosystem services provided by bats, we performed generalized linear mixed models (GLMM) fitted with a Poisson distribution to analyze potential differences among sampling methods. We identified 409 bat species that provide ecosystem services, 752 insect species consumed by bats and 549 plant species either dispersed or pollinated by bats. Our review summarizes the importance of conserving bat populations and the ecological services they provide, which is especially important during the current pandemic.

The Inca Empire was the largest ancient civilization in South America, extending across almost 4000 km of distinct environments from the Pacific coast to the Andes, and throughout most of the arid desert between. The present study highlights the importance of guano birds (Guanay Cormorant Leucocarbo bougainvillii, Peruvian Pelican Pelecanus thagus and Peruvian Booby Sula variegata) for the expansion and prosperity of the Inca Empire. The use of guano as a fertilizer was fundamental to sustain the agricultural development of the empire and, it has been argued, was the basis for its rapid growth. The access to guano on coastal islands and its subsequent transportation to highlands provided food security for a population of more than 8 million. The importance of guano birds to the Inca Empire led to the development of management plans based on a penal code aiming to preserve these species and their natural habitats. These protective actions may represent the first conservation measures ever implemented by humans based on the importance of species protection for human activities and livelihoods.

The soils of the Maritime and sub-Antarctic experience extreme environmental conditions but nonetheless host biological communities that can survive low temperatures, limited water availability and short day lengths or even the complete absence of solar radiation during the winter. We determined the soil organic carbon (SOC) and total nitrogen (N), soil microbial biomass (SMB), labile carbon (LC) and respiration rate (RR) in soil samples from the longest latitudinal transect (approximately 2,000 km) ever sampled exclusively in the Maritime and sub-Antarctic, comprising 69 sites located between South Georgia (54°S) and south-eastern Alexander Island (72°S). With the exception of the most northerly location (South Georgia), the greatest SMB values occurred at 67–68°S, despite the harsh environmental conditions. This is consistent with the relative nutrient richness of the soils at these latitudes, as indicated by large SOC and total soil N concentrations, which are in turn probably linked to more abundant guano and excreta deposition by sea birds and seals, which have fewer and smaller areas of ice-free terrain to land or haul out on further south. South of 68°S, SOC values declined with increasing latitude, which is probably due to increasingly extreme environmental conditions. We also found that the SOC and SMB values, the proportion of SOC considered labile (LC/SOC) and the carbon mineralization rates expressed as either RR/SOC or RR/SMB were all small compared with values from less extreme temperate and tropical regions. However, the proportion of the SOC in the microbial biomass (SMB/SOC) was substantially greater than that reported for non-polar soils. We conclude that although the soils of the Maritime and sub-Antarctic have small and relatively inactive microbial communities, they are characterized by efficient conversion of organic resources into microbial biomass and large affinities for added substrates. Highlights: Soil carbon parameters are reported from sites along the longest latitudinal transect in Antarctica ever sampled. The soil microbial biomass tended to peak at 67–68°S, coinciding with relative SOC and N abundance in the soil, probably arising from guano and excreta from sea birds and mammals. The soil microbial community was small but highly efficient at converting organic resources into microbial biomass, as indicated by large microbial biomass/soil organic carbon ratios.

The ability of some ant species (including Camponotus spp.) to forage on vertebrate urine to extract urea may extend their niche in competitive and strongly nitrogen‐limited environments. We examined the preference of Camponotus terebrans, a sand‐dwelling ant widespread in southern Australia, for baits including urine, and the duration of their foraging on those baits. We baited ants with liquid stains of urine (human and kangaroo), urea in water (2.5%. 3.5%, 7.0%, 10.0%) and sucrose in water (20% and 40%) poured directly on the ground, as well as hard baits in plots drawn on sandy soil (Kangaroo Island, South Australia). We counted individuals of this mostly nocturnal species to determine their attraction to different baits for one month. We checked plant growth on the plots after nine and 13 months. Ants collected insects and meat; they foraged for at least 29 days on stains. Ants were most numerous on 10% urea, followed by 7% urea, 3.5% urea, urine (which contains ~2.5% urea) and 2.5% urea, 40% sucrose and 20% sucrose; sucrose was less attractive to them than equimolar urea bait. Ants were attracted to human, kangaroo, and unidentified urines, and they collected bird guano. Baits and ant foraging did not affect plant recruitment in plots. We observed incidentally Camponotus consobrinus foraging on urine, which may be a common resource for this genus at the site. The remarkable ability of C. terebrans to extract nitrogen from dry sand over weeks explains partly its success on sandy soils. Foraging on urine may be an important strategy to address nitrogen limitation on sandy soils and exploit commensally niches in which hosts are kangaroos, wallabies and other vertebrates. The understanding of plant–vertebrate interactions must factor in the role of ants as commensal organisms. Such ants could also reduce greenhouse gas emissions from urine.

A 285‐cm core of bat guano was recovered from Magurici Cave in north‐west Romania and analyzed for δ13C, δ15N and pollen. Guano deposition occurred from AD 881 until 1240 and from AD 1651 to 2013, allowing for the interpretation of summer variations in precipitation and temperature during the Medieval Warm Period (MWP) and the Little Ice Age (LIA). A 2‰ increase in δ13C, 1.5‰ decrease in δ15N, and the presence of Ulmus, Quercus and Carpinus betulus indicate a warm and dry MWP occurred in the region. The lack of deposition during the beginning of the LIA suggests a possible climate‐induced change in prey availability resulting in bats vacating the cave. Variation of δ13C values between −25 and −23‰ at AD 1650 (LIA) indicates similar drier conditions as at the end of MWP. However, a 2‰ decrease in δ13C values that occurred between AD 1790 and 1900 suggests climate was trending towards wetter conditions at the end of the LIA. From AD 1938 to 2013, δ13C values appear to be more influenced by temperature, indicating that this parameter had a more significant effect on carbon discrimination than water availability.

Two amoeboid organisms of the genera Sappinia Dangeard, 1896 and Rosculus Hawes, 1963 were identified in a sample containing king penguin guano. This sample, collected in the Subantarctic, enlarges the list of fecal habitats known for the presence of coprophilic amoebae. The two organisms were co-isolated and subcultured for over 6 mo, with continuous efforts being invested to separate each one from the mixed culture. In the mixed culture, Rosculus cells were fast growing, tolerated changes in culturing conditions, formed cysts, and evidently were attracted by Sappinia trophozoites. The separation of the Rosculus strain was accomplished, whereas the Sappinia strain remained intermixed with inseparable Rosculus cells. Sappinia cell populations were sensitive to changes in culturing conditions; they improved with reduction of Rosculus cells in the mixed culture. Thick-walled cysts, reportedly formed by Sappinia species, were not seen. The ultrastructure of both organisms was congruent with the currently accepted generic characteristics; however, some details were remarkable at the species level. Combined with the results of phylogenetic analyses, our findings indicate that the ultrastructure of the glycocalyx and the presence/absence of the Golgi apparatus in differential diagnoses of Sappinia species require a critical re-evaluation.

Availability of iron limits productivity in the Southern Ocean where vast regions of high-nitrogen low-chlorophyll conditions are punctuated by hotspots of production surrounding the subAntarctic islands. Here, we compare patterns in accumulation of iron, as indicated by [Fe] and Fe : C (from Wing et al. 2014), with proxies for uptake of iron within food webs, as indicated by Fe : Al and δ56Fe within the subAntarctic Auckland Island ecosystem. We compare these proxies for dynamics of iron within egested material collected from six seabird species, representing three distinct foraging guilds. Fractions of biogenic iron, estimated from Fe : Al, were lowest in the coastal foraging guild (0.83 ± 0.01), and higher in the oceanic and predator guilds (0.94 ± 0.01 and 0.95 ± 0.01), consistent with high rates of biological recycling. The isotopic ratio of iron 56Fe/54Fe, reported as δ56Fe, is depleted in the heavy isotope 56Fe by recycling in the microbial loop, and is likely further depleted in 56Fe when taken up by animals within the food web. δ56Fe was most negative for the oceanic foraging group indicating intensive recycling (−1.16 ± 0.16‰), most positive in the coastal group where direct lithogenic sources of iron are more available (−0.14 ± 0.15‰), and intermediate in the predatory group, representing a mixture and possible fractionation within food webs (−0.37 ± 0.14‰). These data reveal strong contrasts in bioaccumulation and recycling of iron between coastal and oceanic food webs, and important interactions between marine food web structure and biogeochemical processes supporting productivity in the Southern Ocean.

Chemical weathering of archaeological material is well known; however, while there have been a number of experimental studies demonstrating different types and degrees of chemical alteration on faunal remains, little attention has been paid to stone tools, and the few studies that exist relate mainly to siliceous materials and water-induced chemical alteration. Azokh Cave, located in the South Caucasus, contains a Middle Pleistocene to Holocene infill, and detailed macro- and microscopic examination of the lithic assemblages recovered there indicates potential chemical weathering of the stone artefacts. The cave is also currently home to one of the largest bat colonies in the region, and their guano forms a significant component of the infill of the inner galleries. Based on these two factors, an experimental pilot study was set up to artificially chemically alter a range of stone flakes produced specifically for the task, in order to determine the nature and likely cause of weathering. The experimental flakes, produced from different raw materials, were buried in fresh bat guano for up to two years. The results reported herein demonstrate that in a relatively short time, the highly acidic composition of bat guano strongly affects calcium-bearing rocks (e.g., limestone, basalt) altering their entire surface. Similar comparisons may be made with chemical alteration evident on archaeological lithics from Azokh Cave, suggesting that bat guano has played a significant role in diagenetic alteration.

Seabird islands house large colonies of seabirds that spend most of their time at sea but return to land to breed. Seabirds deposit large amounts of nutrients of marine origin on islands; many species also disturb soil and vegetation during the building and maintenance of nests. Physical and chemical disturbance by seabirds alters the chemical composition of soils and vegetation and modifies plant and animal communities. In many cases, the additional nutrients brought in by seabirds subsidise native animal populations, including endemic species frequently found on remote islands. Introduction of predators such as rats, cats and mustelids has resulted in population decreases or local extinction of many seabird species, and this leads to large changes in nutrient cycling and plant and animal populations. Recent efforts at eradication of introduced seabird predators have been successful, but re-establishment of seabirds and seabird island communities will likely take additional active restoration. Key Concepts: Seabirds deposit nutrients of marine origin in terrestrial environments. Seabirds impose physical disturbance on soils and vegetation in the process of building and maintaining nests. Seabird islands often house endemic and native plants and animals that are no longer found on the mainland. The presence of seabirds selects for plants that are tolerant of high-disturbance and high-nutrient conditions. Nutrient subsidies from guano (i.e. nitrogen and phosphorus) can support increased primary and secondary productivity. Introduced mammalian predators have eradicated seabirds and other species on many seabird islands. The reduction of seabird populations by introduced predators alters plant and animal communities and nutrient cycling. Eradication of seabird predators is a necessary first step towards restoration of seabird islands. Restoration of seabird islands often requires active management of seabirds, other animals and plant populations. Protection of seabird islands will require international cooperation on land and at sea. Keywords: allochthonous inputs; ecosystem engineering; invasive predators; island endemism; island restoration; nutrient subsidies; physical disturbance; predator eradication; superpredator

Bat guano cores have been used as a source of palaeoenvironmental information to aid in the reconstruction of past climates and vegetation. We collected a 104-cm-long (43 cm compacted) guano core from Fern Cave, Alabama, USA, that provided a c. 6000-year record of guano accumulation. Pollen, nutrients (C, N, P) and stable isotopes (δ13C, δ15N) were measured on the guano core with the objective of reconstructing the environmental history of the area from the mid-Holocene to present. Our data indicate that bats have utilized Fern Cave for at least 6000 years and that Woodland Indians also utilized the cave for a short period. A 3-cm charcoal layer was dated to 2720±30 cal. a BP and inferred to be Woodland Indian in origin from microscopic inspection and thickness. Pollen and geochemical data showed that bat diets changed in the late Holocene possibly linked to food supply and climate changes. These results demonstrate that guano cores are a useful tool of palaeoenvironmental reconstruction when other forms of palaeorecords do not exist and can add to local archaeological information.

Since the introduction of white‐nose syndrome (WNS) in North America, numerous species of bat have dwindled in numbers. These declines observed are often species‐specific and thus provides opportunity for a natural experiment to test for shifts in diet through relaxed resource partitioning in bat communities post‐introduction of WNS. Acoustic monitoring at locations in Southern Ontario pre‐ (2009–2011) and post‐WNS (2012–2014) introduction showed an increase in activity of big brown bats (Eptesicus fuscus) corresponding to a decline in the activity of little brown bats (Myotis lucifugus). Next‐generation sequencing of bat stomachs and guano in Southern Ontario before and after WNS allowed for the characterization of diet changes of these species. Post‐WNS, E. fuscus consumed a wider breadth of prey and many of the insect species once consumed by M. lucifugus, including several pest insects. These results suggest that the introduction of WNS has resulted in relaxed interspecific competition within these bat communities in Southern Ontario.

Summary Limited pedogenesis on the Antarctic continent results in unusual, but important, short-range order inorganic phases and organo-mineral complexes (both with broad peaks from X-ray diffraction) in the clay fraction, which can be studied by sequential chemical extractions. We aimed to develop a better sequential extraction procedure to obtain more detailed discrimination of the short-range order phases (SROP) of the clay fraction of soil from Maritime Antarctica. Selected chemical properties of the soil and clay fraction were related to different phases of these products. The total amount of SROP and organo-mineral complexes in the clay fraction ranged from 15.1 to 43.9%, with greater occurrence in ornithogenic soil. The continuous incorporation of guano from penguins into the soil profile has contributed to increasing amounts of organo-mineral complexes. Short-range order phases have an important effect on soil chemical properties, such as an increase in non-exchangeable acidity, quantity of hydroxyl groups on the surface of the colloidal fraction, the buffering capacity of Al3+ from non-exchangeable and structural forms of the element, and a decrease in soil CEC. The greater number of sequential extraction steps enabled the separation of SROP and organo-mineral complexes into two phases of stability, with a clear predominance of the more soluble forms.

Seabirds play an important role in coastal environments, serving as key indicators of marine ecosystem variability as well as biovectors that influence terrestrial productivity and carbon storage. Long-term estimates of seabird populations remain rare, but lakes that support large seabird populations in their watersheds can archive a history of seabird activity in their sediment records. Here we present a seabird guano-influenced sediment record from Genovesa Crater Lake, Galapagos Islands, home to the world's largest reported colony of red-footed boobies (Sula sula) and smaller populations of other species. Influx of seabird guano into Genovesa Crater Lake produces high sedimentary δ15N values, and temporal variability in sediment δ15N primarily reflects changes in guano influx through time. Two abrupt increases in sedimentary δ15N occurred at 1835 AD and 1965 AD, and variance increased following the 1965 AD shift. The largest of these abrupt shifts at 1835 AD coincided, within age model error, with an abrupt increase in marine productivity indicators in sediment records off the coast of Peru and Chile. In the latter part of the twentieth century, δ15N values increased during periods of higher landings of Peruvian anchoveta and sardines. We hypothesise that seabird presence and activity on Genovesa increased during periods of higher regional marine productivity. Enhanced variance in Genovesa δ15N following the 1965 AD shift may reflect a modern population more susceptible to climate and environmental variability than at any other time in the last 400 years.

Using nitrogen isotope ratio as proxy for nutrient transfer, we tracked nutrients from bat excreta to seed exocarp in Dipteryx panamensis that host bat roosts, suggesting that trees may benefit from nutrients of bat guano deposited into hollow tree trunks. Resume Usando el ratio de isotopos estables de nitrogeno como estima de la transferencia de nutrientes, hemos investigado la asimilacion de nutrientes procedentes de excreciones de murcielagos por el exocarpo de las semillas de arboles Dipteryx panamensis que sirven de refugio a murcielagos. Los resultados sugieren que los arboles podrian beneficiarse de los nutrientes del guano de los murcielagos depositado en los troncos huecos de los arboles.

Summary The serious forest pathogen Heterobasidion annosum sensu lato was found during mycological exploration in caves and underground tunnels. Seventeen isolates of Heterobasidion annosum s.l. were obtained in these habitats in the Czech Republic, Slovakia, Romania and Spain during 2008–2012. Three species, H. abietinum, H. annosum sensu stricto, and H. parviporum, were identified by analyses of DNA sequences of three gene regions: glyceraldehyde 3-phosphate dehydrogenase, translation elongation factor 1-alfa and transcription factor. Isolates of H. parviporum were obtained from moonmilk, air and underground gallery wall. H. parviporum recorded from Spain, Castaňar de Ibor Cave, was obtained far from the natural distribution of its main host, Picea abies. Heterobasidion annosum was obtained from cave air and sediment, isopod faeces and underground gallery wall. Heterobasidion abietinum was isolated from bat guano and outdoor air. The origin and ways of fungal dispersal are discussed. The forest stands surrounding caves and tunnels are hypothesized as the potential source of inoculum to inhabit these unusual spaces, although the real paths of the pathogen to the habitats remain unclear.

The decline of the Tiwanaku state saw the emergence of two new cultures-Pica-Tarapaca and Atacama-during the Late Intermediate Period in northern Chile. Archeological evidence suggests that both groups practised maize agriculture and pastoralism, but that their interaction zones differed significantly. Marine resources are common at Pica-Tarapaca sites, even those far from coast, while Atacama sites in the desert oases and precordilleran area seem to have directed their networks towards the highlands. Here we apply stable carbon, nitrogen, and oxygen isotope analysis on human bone and enamel to test dietary patterns and residential mobility at two sites, Pica 8 and Quitor 6, representing the Pica-Tarapaca and Atacama cultures, respectively. Our results show that diet at the two sites indeed differed: significant but variable consumption of marine resources and maize is indicated at Pica 8, despite being an inland site, while diet at Quitor 6 was based mainly on terrestrial resources. The use of seabird guano and llama dung as fertilizers and extreme aridity may have contributed to the high nitrogen isotope values observed in Pica 8 humans. The δ(18) O values in Pica 8 individuals are generally lower than for Quitor in spite of its greater distance from the Andes. All three isotopes suggest the presence of at least five nonlocals in the 30 measured at Pica 8. This evidence for human mobility is consistent with the high levels of trade and interaction observed in the archeological record, and begins to quantify the degree of movement of specific individuals.

Carbon and nitrogen stable isotopes were used to track major dietary variations in arthropods found in a cave located in a neotropical savanna in Central Brazil. We determined the δ 13 C and δ 15 N for cave crickets, cockroaches, spiders, guano of a hematophagous bat, and leaf litter found on the ground near the cave entrance. The δ 13 C and δ 15 N for the cricket and cockroaches showed that bat guano was not the only food item for these arthropods. They had intermediate δ 13 C and δ 15 N between bat guano and leaf litter, which means that they consumed additional food resources other than guano in their diet, independent of distribution of guano deposits in the cave.The spiders, predators with great mobility, seemed to not have a preferential diet item since their isotope signals indicated they hunt both crickets and cockroaches. The δ 13 C of the bat guano (−15.4‰) indicated that the diet of these hematophagous bats relied on animals fed with C4 plants.The conversion of native savanna vegetation of which the debris is mostly from C3 plants, to pasture, based on C4 African grass, a remarkably common land use transformation in the region, is indirectly influencing the diet of hematophagous bats, which basically relied on exotic fauna. The C and N stable isotope ratios showed that the arthropods inside the cave do not rely solely on bat guano for their diet, but interact directly with the external environment, through litter debris, indicating a significant exchange of energy and matter between the cave environment and the surrounding area.

Oceanic island ecosystems are particularly sensitive to El Niño effects due to their dependence on energy and nutrient inputs from marine systems. Seabirds play a key role in transporting resources of marine origin to insular ecosystems. We report tree-growth patterns showing how the effects of El Niño rainy events on tree species in a southern Pacific island depend on the presence of local seabird colonies. We performed manipulative experiments in order to assess the mechanisms underlying these patterns. Tree ring data showed that, in normal years, the growth of all tree species (Aextoxicon punctatum, Cryptocarya alba, and Pinus radiata) was significantly lower in seabird sites compared to adjacent patches without seabirds (control sites). In contrast, in El Niño years, trees formerly hosting seabird colonies grew more than those in control sites. Experiments showed that (1) pine plants on soil from seabird sites grew more than those on soil from control sites, (2) pine individuals with seabird feces on their leaves grew less than those sprayed with an aqueous solution, and (3) soil moisture had little effect on plant growth. The stress produced by massive cormorant nesting on trees, which impairs tree growth and physiological performance, is relieved during El Niño events because of seabird migration due to decreased prey availability and pouring rains that flood nests. Soils enriched by the seabird guano, together with the increased water availability associated with El Niño, foster the growth of trees from seabird sites. We suggest that El Niño may be a key determinant of tree performance in forest communities from island and coastal ecosystems of the Pacific Ocean.

Bats use large (69 cm mean diameter) ponderosa pine (Pinus ponderosa) snags for maternity roosts in northern Arizona. Anthropogenic causes have altered forest structure, decreasing density of large snags. To examine use of artificial roosts for bats in these forests, we selected 26 sites and 2 treatments (clustered, single) using 1 type of roost (resin or wood) per site. At each site, we installed 4 roosts on ponderosa pine trees with a cluster of 3 roosts (south-, east-, or west-facing

Diet has a profound influence on the fitness of adult tephritid flies. Mass-reared flies are provided yeast hydrolysate as a rich source of nutrition that supports rapid sexual development and mating success. In contrast, wild tephritid flies often live in environments where food may be hard to find, and these are the conditions that sexually immature mass-reared sterile males encounter when released into the field during sterile insect technique campaigns. The effect of natural food sources (bat guano, bird droppings, citrus pollen, and wheat pollen) on the sexual development of adult mass-reared fertile, mass-reared sterile, and wild male Queensland fruit flies, Bactrocera tryoni (Froggatt) (Diptera: Tephritidae), was determined by measuring ejaculatory apodeme size. Inclusion of yeast hydrolysate in the adult diet was associated with faster growth of the ejaculatory apodeme in comparison with all other diets. Effects of diet were far less pronounced in mass-reared males, which may indicate reduced nutritional requirements, whereas the ejaculatory apodeme of wild males fed on natural sources of food or sucrose alone did not increase in size over the first 20 days of adult life.

Large collections of Eccentrotheca helenia sp. nov. from the lower Cambrian Wilkawillina and Ajax limestones in the Arrowie Basin, South Australia, contain abundant low, cap-shaped and high, laterally compressed isolated sclerites in addition to partially articulated tubular specimens. The scleritome of Eccentrotheca helenia sp. nov. is fully described for the first time and shown to be formed by ontogenetic fusion of sclerites into successively stacked sclerite rings, forming a larger, tubular structure. The apical termination of the tube is highly variable, but is primarily constructed by low, cap-shaped sclerites and characterised by a central aperture of variable inclination. The adapical portion of the tube is predominantly constructed by high, laterally compressed sclerites, but individual sclerite rings can contain both cap-shaped and laterally compressed sclerites along with sclerites of intermediate morphology. The apical aperture presumably housed organic structures for attachment to a hard substrate, but the scleritome also occasionally preserves small lateral perforations between fused sclerites, which may have served to stabilise the scleritome by providing additional points of anchorage. In the Arrowie Basin, E. helenia is found in association with archaeocyath-microbial-spongiomorph-dominated bioherms and most likely inhabited pendant or cryptic habitats within these bioherms. Eccentrotheca-like sclerites form an integral part of the scleritomes of many tommotiids which may confuse taxonomic analysis. Sclerites previously assigned to 'E.' guano, consistently occur together with sclerites of Kulparina rostrata in stratigraphic intervals consistently older than strata hosting E. helenia. Rare fused specimens indicate that the sclerites of K. rostrata and 'E.' guano belong to the same scleritome.

Batina, M. C. & Reese, C. A. 2010: A Holocene pollen record recovered from a guano deposit: Round Spring Cavern, Missouri, USA. Boreas, 10.1111/j.1502-3885.2010.00186.x. ISSN 0300-9483. Bat guano deposits have been the subject of relatively few palynological studies. The studies to date have focused on the most modern samples only, and therefore the full potential of bat guano as an archive for fossil pollen analysis remains unknown. In this paper we report the results of pollen analysis on an 88-cm vertical profile of fossil bat guano from Round Spring Cavern, Missouri. Four AMS radiocarbon dates were in sequence and showed that the guano pile was active between 8155 and approximately 550 cal. yr BP. Pollen is well preserved throughout the profile, with most samples containing less than 10% indeterminable pollen. Wind-pollinated taxa are better represented than insect-pollinated taxa, and pollen concentrations range from 842 to 371 660 grains cm−3. As expected, the dominant vegetation types represented by the pollen assemblage are oak and pine, in accordance with the region's oak–hickory–pine forest. However, there are discrepancies between the guano pollen record and the results of a lake pollen study conducted in southern Missouri. We propose several possible factors that may be influencing the guano pollen record and provide direction for further investigation into the palaeoecological potential of bat guano.

Invasive species are widespread and can have devastating effects on biota, especially insular biota. Invasive species eradications are increasingly employed to promote island recovery to preinvasion states. However, it remains unclear if additional restoration actions may be required on islands that were once heavily reliant on seabird guano for ecosystem functions. Active seabird augmentation has been suggested as necessary to exact ecosystem recovery on contemporary timescales in some cases. I use two experiments on offshore islands in Cook Strait, New Zealand, to test the hypothesis that seabird restoration will restore island ecosystem functioning following invasive rodent removal. The first is a small-scale single-island fertilization experiment that simulates seabird recovery. This experiment tested the recovery potential of offshore islands and was used to infer the density of seabirds needed to elicit ecosystem recovery. The second is a large-scale natural experiment that takes advantage of eight islands with differing rodent eradication and seabird restoration histories. I compared ecosystem functioning variables (delta 15N, C:N ratios in soil, plants, and spiders, as well as arthropod abundance and diversity) on two islands that had rodents eradicated and two islands undergoing seabird augmentation with two control islands (never invaded by rodents) and two positive control islands (currently invaded by rodents). The results suggest that islands do have the potential for recovery given nutrient amendments, but that islands with rodents eradicated and islands undergoing seabird augmentation have not recovered most of their ecosystem function. Finer, intra-island analysis showed that seabird restoration projects have the potential to speed the recovery process, but that the projects on the studied seabird restoration islands were not advanced enough to produce island-wide recovery. The results suggest that high seabird densities (5-10 burrows/m2) are needed to promote recovery to never-invaded control levels. Seabird augmentation, through chick translocation and/or social facilitation with decoys, vocalization playbacks, and/or mirrors can supplement passive seabird recovery on islands where seabirds have been extirpated or extremely reduced by invasive predators. Such restoration efforts may be necessary to promote ecosystem recovery on contemporary timescales.

Phosphorite, or guano, or insular phosphorite, commonly containing 10-35% P2O5, is one of the sources of phosphatic fertilizer. Taking the modern insular phosphorite on the Xisha Islands as an example, this paper discusses the ore-forming mechanism of phosphate and the important contribution made by animals to this process. The phosphorite occurs in the middle part of carbonate sand cays of coral reef surrounded by longshore sandbanks, which assumes a dish-shape. The surface of the cays is covered by unconsolidated guano. The phosphorite, yellowish brown to dark brown, is made up of thick-bedded sand and gravel cemented by collophanite; and partly replaced by carbonate apatite. The mechanism of mineralization is as follows: The average content of phosphorus in the sea water is only 0. 07 ppm, but through the sea food. chain concentration, phosphorus content may become higher. The phytoplanktons, as producers, suck up phosphorus out of sea water. The primary consumers, zooplanktons, obtain phosphorus by eating phytoplanktons, then they are followed by the second and third consumers. Sea birds living on fish, as the third consumers, concentrate and transport the phosphorus to the carbonate sand cays. Finally layers of guano are deposited under the arbores. Under the tropical climatic conditions with high temperature and plentiful rainfall, soluble matters are leached out of the guano and infiltrated into deeper horizon. In alkaline groundwater with pH greater than 7. 0, the phosphoritization takes place, bioclastics are cemented and replaced by collophanite or carbonate apatite. Finally, the phosphorus is fixed and phosphate formed.

SUMMARY At a breeding colony on Ascension, many Black Noddies remained throughout the year. The main part of the paper describes their behaviour, with special reference to daily rhythm, sunning, absence of “panics”, nesting (with possible value of the birds' own guano for cement), fighting, and courtship (with details of ground and aerial displays), copulation (apparently rare) and the behaviour of parents and young. On Ascension the birds nest on small cliff-ledges, elsewhere they also use trees, both kinds of sites being relatively inaccessible to ground predators, unlike the majority of terns, which nest on the ground. As with the Kittiwake, another cliff-nesting species which has diverged from ground-nesting stock, a number of the Black Noddy's peculiarities can be related directly or indirectly to its nesting site. Indeed the occurrence in just these two species of two features absent in their ground-nesting relatives indicates that there are some cliff-nesting adaptations which had previously been overlooked. The Black Noddy's adaptation in this respect can be summarised as follows: Cryptic adaptations lost. Cryptic plumage of nestling and juvenile. Dispersal of egg-shells and droppings. Adaptations acquired, and their consequences. Shortage of nest-sites, focussing territory on nest, with consequent increasing, though not exclusive, importance of Bridling as “song”. Allopreening between mates. More elaborate nest-building, with gathering of material (if available), which is cemented together with guano; side-stepping and depressing tail when shaping nest. Young intolerant of others; crouch if threatened instead of running; peck at feet of intruders, or beak-hide to appease attacks, precocious development of adult (threat?) plumage. The Brown Noddy is the closest relative of the Black, and what little is known of its breeding biology suggests it is less specialised for nesting above the ground, and that its ability to nest on the ground as well as in bushes and trees allows it to breed in localities where the Black Noddy could not.

Adjacent communities and ecosystems often differ in underlying productivity but are connected by flows of nutrients, energy, and matter. Pacific salmon (Oncorhynchus spp.) transport substantial quantities of nutrients from marine ecosystems to coastal freshwater habitats when they return to spawn and die. Nutrients from their carcasses are initially concentrated in spawning streams and lakes, but are subsequently dispersed by abiotic (floods, hyporheic flow) and biotic processes (predators and scavengers). In southwest Alaska, mobile avian scavengers (gulls; Larus spp.) breed on small islands within salmon nursery lakes and consume large quantities of spawning salmon during the chick-rearing period. However the role of birds as vectors of salmon-derived nutrients remains unknown. We examined how gulls - by transporting salmon tissues to their chicks - create hotspots of biological productivity in the aquatic habitats surrounding their nesting colonies. We found that algal production was ~ 10 x higher at islands with high gull densities compared to islands without nesting gulls, but was concentrated within 40 m of island shorelines. Carbon stable isotopes (51 C) confirmed that gulls enhance primary production in local benthic communities and demonstrated that this production was transferred up the food web to grazers (snails) and carnivores (blackfish). Nitrogen stable isotopes (515N) confirmed that salmon dominated the diet of gulls and that nutrients from gull guano were incorporated into algae and passed up the food web. By relocating and concentrating salmon-derived nutrients into new and distant locations, gulls alter and magnify production in local aquatic communities. We offer the first evidence that the avian community can move salmon-derived nutrients great distances, enriching otherwise isolated habitats.

The Florida Everglades is an oligotrophic wetland system with tree islands as one of its most prominent landscape features. Total soil phosphorus concentrations on tree islands can be 6 to 100 times greater than phosphorus levels in the surrounding marshes and sloughs, making tree islands nutrient hotspots. Several mechanisms are believed to redistribute phosphorus to tree islands: subsurface water flows generated by evapotranspiration of trees, higher deposition rates of dry fallout, deposition of guano by birds and other animals, groundwater upwelling, and bedrock mineralization by tree exudates. A conceptual model is proposed, in which the focused redistribution of limiting nutrients, especially phosphorus, onto tree islands controls their maintenance and expansion. Because of increased primary production and peat accretion rates, the redistribution of phosphorus can result in an increase in both tree island elevation and size. Human changes to hydrology have greatly decreased the number and size of tr...

Species-specific primers for portions of the mitochondrial DNA 16S ribosomal subunit gene were designed to identify bats of the family Vespertilionidae (Mammalia). Two fragments, each of 190 base pairs, were found to contain sufficient genetic variability to consistently resolve 10 of the 14 Pacific Northwest species included in this study. The remaining four species could be resolved into two sets of paired species that were unique from the other 10 species. These primers are demonstrably useful for purposes of surveys designed for species identification of bats, including using DNA extracted from guano collected from roost sites when bats are absent.

More than 100 radiocarbon dates of penguin guano and remains, shells and seal skin afford ages for raised beaches adjacent to Terra Nova Bay, Antarctica. These dates permit construction of a new relative sea-level curve that bears on the timing of deglaciation. Recession of the Ross Sea ice-sheet grounding line from Terra Nova Bay occurred no earlier than 7200 14C yr (8000 cal. yr) BP. Retreat along the Victoria Land coast may have been rapid, possibly contributing to eustatic sea-level rise centred at ca. 7600 cal. yr BP. The presence of a significant amount of ice remaining in the Ross Sea Embayment in Holocene time lessens the chance that Antarctica contributed significantly to meltwater pulse 1A several thousand years earlier. Copyright © 2004 John Wiley & Sons, Ltd.

Southern fur seals Arctocephalus australis in Peru have declined gradually over the past decade, and declined dramatically (72%) as a result of low food availability during the severe El Nino in 1997–98. In 1999, seals abandoned some historically important breeding sites. This is particularly alarming because new sites were not colonized. Our objective was to examine how habitat features and human disturbance influenced whether sites were currently used, abandoned or apparently not used in the past by fur seals for breeding. Data were collected on 14 variables at 70 potential breeding sites at three guano reserves in Peru. Discriminant analysis revealed significant multivariate differences among sites currently used for breeding, abandoned sites and unused sites (F=5.97, P

The corn–guano model is the simplest model with exhaustible resources. On the constant rate of profit hypothesis, a change of the numeraire from corn to corn-and-labour affects the price trajectories. In particular, the corn price at the exhaustion date is no longer equal to its long-term level. The system of intertemporal prices admits one degree of freedom. Only one path, called the natural path, admits a positive price, wage and royalty for any number of periods before exhaustion. But, for a given exhaustion date, the non-natural paths close to it are also associated with positive prices, wages and royalties. In this paper we study the characteristics of the natural path and the properties of the non-natural paths. The results are partly extended to multisector models.

Allochthonous marine input is a key component of the dynamics of islands and terrestrial coastal ecosystems on islands in the Gulf of California, Mexico, where an unproductive desert juxtaposes a highly productive ocean. In this area, seabirds are a major conduit bringing marine productivity to land. Seabirds leave guano on roosting and nesting islands, and carrion on nesting islands. We analyze how seabirds affect the abundance of a dominant consumer group, tene- brionid beetles, on 25 islands in the Gulf of California over a three-year period. Tenebrionid densities vary by three orders of magnitude among islands. Beetles are most abundant in areas influenced by seabirds: they are approximately five times more dense on nesting and roosting islands than on other islands and on mainland sites, and approximately six times more dense inside vs. outside colonies. Path analyses show that seabirds significantly affect beetles by two distinct pathways. On roosting islands, effects are mainly indirect: guano, a fertilizing nutrient, significantly enhances plant productivity; beetles eat plant detritus. On nesting islands, significant effects are primarily direct: beetles eat seabird carrion; the indirect pathway (guano and plants) contributes little on these islands. By providing energy and nutrients to fuel a diverse array of consumer populations, seabirds are central to the dynamics of these island ecosystems. Scavengers and avian parasites directly increase by eating seabird products. Likewise, populations of consumers that eat detritus, plant tissues, and seeds are enhanced indirectly via the fertilizing effects of guano on plants. Increases in these primary consumers indirectly facilitate high densities of many predators. Thus, consumer populations on these islands are deeply subsidized by substantial input from seabirds. Because consumers cannot influence the renewal rate of their allochthonous resources, the dynamics of these consumers and their food webs are largely donor controlled.

Surface sediment, speleothems, and bat guano from two adjacent, topographically different cave sites in eastern Spain have been studied palynologically to elucidate the potential of cave sediments for palaeoenvironmental reconstruction. A cave opening with a large entrance and constant width presented far fewer problems of alteration in the pollen assemblages than a sac‐like cave opening with a constricted entrance. Pollen concentration is linked primarily to the amount of pollen input rather than to the result of post‐depositional alteration. Sampling should be undertaken away from parietal and rear areas and avoiding moisture zones. Lateral differences in the pollen spectra indicate that sampling should be on the basis of a multiple‐profile approach and selection of dry rather than wet sediments. If these procedures are followed, within‐cave sediments can realistically reflect not only local but also regional vegetation of the site. Fil: Navarro Camacho, C.. Universidad de Murcia; España Fil: Carrión, J. S.. Universidad de Murcia; España Fil: Navarro, J.. Universidad de Murcia; España Fil: Munuera, M.. Universidad de Murcia; España Fil: Prieto, Aldo Raul. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Mar del Plata. Instituto de Investigaciones Marinas y Costeras. Universidad Nacional de Mar del Plata. Facultad de Ciencias Exactas y Naturales. Instituto de Investigaciones Marinas y Costeras; Argentina

Most of the old-growth redwood (Sequoia sempervirens) in California has been cut; regenerating forests willprobably never resemble those that were harvested, and what old growth remains on private land occurs in small, isolated remnant patches. The landscapes in which these stands occur differ so markedly from their original condition that their value as habitat to many species of wildlife, including bats, is un- known. Previous research in unfragmented redwoodforests demonstrated that bats use basal hollows in old- growth redwoods as roosts. We sought to determine whether bats use similar old-growth trees as roosts when they occur in small, remnant patches of isolated old growth on commercialforest land. We compared bat oc- currence in remnant and contiguous stands by collecting guano in traps suspended in hollows and by moni- toring flight activity with ultrasonic bat detectors. Hollows in trees within the remnant stands had signifi- cantly more guano deposited per tree than the trees within the contiguous forest. The mean numbers of bat passes per night were statistically indistinguishable between the two treatments, although mean flight activity in the remnant stands was greater than in the contiguous forest. Bats frequently used basal hollows in small (

Larvae of Lepidoptera are mainly herbivores, and only the larval stages of Bradypodicola and Cryptoses pyralid moths are known to be specialized coprophages. Here, we report coprophagy as a main feeding habit for the larvae of another pyralid moth, Aglossa pinguinalis, on the basis of observational and experimental evidence. The larvae of this moth require two years to complete development, and construct silk tubes connecting the food source with a shelter chamber. Larvae of A. pinguinalis were mostly found inside caves, where they were about 700 times more abundant than in the open field. Inside the caves, observational results indicate that 98% of the larvae fed mostly on excrement and only 2% were recorded as eating other kinds of detritus (decayed mushrooms). No larvae were recorded eating plant detritus in the censuses nor in experimental detritus patches. The larvae did not show preferences for different types of excrement in the caves. Experimental dog and sheep excrement corroborated this result, showing that the abundance of larvae did not differ between these two excrement types over the entire study. We suggest that habitat selection of A. pinguinalis is ecologically similar to the trophic specialization of Bradypodicola and Cryptoses, in spite of the differences in natural history traits among these moths. Feeding on dry excrement inside caves, or in buried sloth dung, may reflect the constraints for Lepidoptera successfully to colonize excrement: in more exposed excrement, they cannot compete with other coprophagous specialists owing to the lack of parental care (e.g. resource relocation) and slow growth rates. We hypothesize that coprophagous Lepidoptera will be restricted to competition- and predation-free habitats and resources, feeding on excrement inside caves and animal burrows.

Bird guano has been shown to alter the structure and function of ecological communities. Here we characterize the effects of vulture guano on the phylogenetic structure, taxa richness, and abundance in soil bacterial communities within an African savanna. By altering soil chemistry and nutrient status, vulture guano appears to play a role in influencing the structure of soil bacterial communities. DNA was extracted from soil collected under twenty trees: five African white-backed vulture (Gyps africanus, WBV) nesting sites, five lappet-faced vulture (Torgos tracheliotos, LFV) nesting sites and ten control sites where no sign of vulture activity was detected. Using a high-density phylogenetic microarray (PhyloChip G2), we identified 1,803 bacterial Operational Taxonomic Units (OTUs) in the twenty samples. Analysis of beta-diversity using the Unifrac distance metric demonstrated that WBV nesting sites were phylogenetically distinct from both control trees and LFV nesting sites. We detected a higher degree of phylogenetic clustering in soil bacterial communities associated with both WBV and LFV nesting sites compared to control sites, suggesting that the deposition of guano increases the strength of habitat filtering in these communities. Canonical correspondence analysis revealed that variation in OTU intensity (a measure of relative abundance) could be related to variations in pH, electrical conductivity and total nitrogen content. WBV sites explained 10% to 22% of the variation in OTU intensity. The elevated total nitrogen and lower pH characteristic of soils associated with vultures may favor Proteobacteria and suppress Firmicutes, particularly Clostridia and Bacilli. Acidic aggregations of vulture guano may be unlikely to support long-term survival of spore-forming Firmicute pathogens and thus may limit the role that vultures play as potential disease vectors.

The production rates of intertidal algae at seven study sites in the Saklanha Bay area. South Africa, were measured monthly for 15 months. Algae grew rapidly and formed extensive, permanent mats on shores that were enriched with nutrients from the dissolved guano of seabirds. The most significant correlate of limpet (Patella granulans) growth was the rate of algal production; more rapid limpet growth on nutrient-rich shores, combined with the effects of predation by African Black Oystercatchers Haematopus moquini on limpets, led to differences in the size structures and life-history patterns of populations of limpets on enriched and unenricheci shores. The rate of primary production affected relationships between intertidal organisms at all trophic levels and influenced patterns of organization within the community.

By far the commonest consituents of insular phosphate deposits are calcium phosphates of the apatite series of minerals, especially varieties that contain structural carbonate in their crystal lattices, e.g. francolite and dahllite. This reflects the fact that the vast majority of described insular phosphate deposits occur in low latitudes, where they have formed, in a tropical environment, by metasomatic replacement of carbonate in coral substrates by phosphate derived either from avian excrement or from phosphorus-enriched lagoonal/lacustrine waters. Thin, recently-formed guano accumulations on islands of the subantarctic Bounty group differ radically in that they consist principally of struvite, a hydrated ammonium magnesium phosphate that is better known as a component of human/mammalian urinary stones and enteroliths, and of bat guano in sheltered speleothems. Apatite occurs only in an accessory role in the Bounty Islands guano. This contrast in mineralogy, and the somewhat anomalous survival of soft, unstable struvite in a totally exposed environment on the Bounty Islands, are attributed to climatic factors and to the nature of the granodioritic rock substrate.

Densely nesting Guanay Cormorants (Phalacrocorax bougainvillii), Peruvian Booby (Sula variegate), and Peruvian Brown Pelican (Pelecanus occidentalis thagus) deserted eggs and young in response to high densities of argasid ticks (Ornithodoros amblus). Alternative causes of desertion such as human disturbance, predation, disease, heat, and food shortage did not fit the evidence. Tick predators were less effective at controlling ticks than were frequent human harvestings of guano. A review of the avian and medical entomological literature indicates that ticks and other ectoparasites may be strong determinants of nesting success and of nest structure and duration of use.

SUMMARY Ectomycorrhizas have been discovered on the root system of Pisoniagrandis R. Br. (Nyctaginaceae) collected from two islands in the Capricorn group of the Great Barrier Reef, Australia. These mycorrhizas are unusual in that the Hartig net is poorly developed but instead transfer cells develop in the epidermis and cortex of the host root. The wall labyrinth is located precisely on host wall regions abutting the fungus. It is suggested that the development of wall protuberances, with an accompanying amplification of host membrane surface area exactly in the region where transfer between the two symbionts is most likely to occur, is an alternative strategy to Hartig net formation. Pisonia grandis is usually found in locations colonized by seabirds as nesting sites and therefore rich in guano. It is unusual to find an ectomycorrhiza in such a situation, where there is a high input of organic nitrogen and phosphorus at least once a year in addition to a permanently high calcium level, and the nutritional aspects of this association deserve further study.

Arixenia esau (Dermaptera) feeds on the skin of bats of the genus Cheiromeles. The food material is collected by the maxillae, lifted through the cibarium by the hypopharynx and then pushed into the oesophagus by the orally directed setae on the mandibles. Arixenia jacobsoni has less specialised mouthparts and feeds upon insects inhabiting bat guano in a similar way to many carnivorous labidurid earwigs. Hemimerus talpoides (Hemimerina), which feeds upon the skin of the giant rats of the genus Cricetotnys, cuts off food particles from the host with the mandibles and then passes the food material through the cibarium into the oesophagus by means of the maxillae. The form of the head and neck of Arixenia can be derived from a pygidicranid or labidurid type and the apparently “primitive” features of the head of Arixenia can be explained as the result of a change of feeding habits from being carnivorous insects to external parasites of mammals. The head of Hemimerus so differs from those of the dermapteran types that this, together with consideration of other anatomical features, resulted in the Hemimerina having been regardod as a separate order of insects.