Insect communities under skyglow: diffuse night-time illuminance induces spatio-temporal shifts in movement and predation.

Artificial light at night (ALAN) is predicted to have far-reaching consequences for natural ecosystems given its influence on organismal physiology and behaviour, species interactions and community composition. Movement and predation are fundamental ecological processes that are of critical importance to ecosystem functioning. The natural movements and foraging behaviours of nocturnal invertebrates may be particularly sensitive to the presence of ALAN. However, we still lack evidence of how these processes respond to ALAN within a community context. We assembled insect communities to quantify their movement activity and predation rates during simulated Moon cycles across a gradient of diffuse night-time illuminance including the full range of observed skyglow intensities. Using radio frequency identification, we tracked the movements of insects within a fragmented grassland Ecotron experiment. We additionally quantified predation rates using prey dummies. Our results reveal that even low-intensity skyglow causes a temporal shift in movement activity from day to night, and a spatial shift towards open habitats at night. Changes in movement activity are associated with indirect shifts in predation rates. Spatio-temporal shifts in movement and predation have important implications for ecological networks and ecosystem functioning, highlighting the disruptive potential of ALAN for global biodiversity and the provision of ecosystem services. This article is part of the theme issue 'Light pollution in complex ecological systems'.

Artificial light at night decreases plant diversity and performance in experimental grassland communities.

Artificial light at night (ALAN) affects many areas of the world and is increasing globally. To date, there has been limited and inconsistent evidence regarding the consequences of ALAN for plant communities, as well as for the fitness of their constituent species. ALAN could be beneficial for plants as they need light as energy source, but they also need darkness for regeneration and growth. We created model communities composed of 16 plant species sown, exposed to a gradient of ALAN ranging from 'moonlight only' to conditions like situations typically found directly underneath a streetlamp. We measured plant community composition and its production (biomass), as well as functional traits of three plant species from different functional groups (grasses, herbs, legumes) in two separate harvests. We found that biomass was reduced by 33% in the highest ALAN treatment compared to the control, Shannon diversity decreased by 43% and evenness by 34% in the first harvest. Some species failed to establish in the second harvest. Specific leaf area, leaf dry matter content and leaf hairiness responded to ALAN. These responses suggest that plant communities will be sensitive to increasing ALAN, and they flag a need for plant conservation activities that consider impending ALAN scenarios. This article is part of the theme issue 'Light pollution in complex ecological systems'.

Ecosystem consequences of invertebrate decline.

Human activities cause substantial changes in biodiversity.1,2 Despite ongoing concern about the implications of invertebrate decline,3,4,5,6,7 few empirical studies have examined the ecosystem consequences of invertebrate biomass loss. Here, we test the responses of six ecosystem services informed by 30 above- and belowground ecosystem variables to three levels of aboveground (i.e., vegetation associated) invertebrate community biomass (100%, 36%, and 0% of ambient biomass) in experimental grassland mesocosms in a controlled Ecotron facility. In line with recent reports on invertebrate biomass loss over the last decade, our 36% biomass treatment also represented a decrease in invertebrate abundance (-70%) and richness (-44%). Moreover, we simulated the pronounced change in invertebrate biomass and turnover in community composition across the season. We found that the loss of invertebrate biomass decreases ecosystem multifunctionality, including two critical ecosystem services, aboveground pest control and belowground decomposition, while harvested plant biomass increases, likely because less energy was channeled up the food chain. Moreover, communities and ecosystem functions become decoupled with a lower biomass of invertebrates. Our study shows that invertebrate loss threatens the integrity of grasslands by decoupling ecosystem processes and decreasing ecosystem-service supply.

Microhabitat conditions remedy heat stress effects on insect activity.

Anthropogenic global warming has major implications for mobile terrestrial insects, including long-term effects from constant warming, for example, on species distribution patterns, and short-term effects from heat extremes that induce immediate physiological responses. To cope with heat extremes, they either have to reduce their activity or move to preferable microhabitats. The availability of favorable microhabitat conditions is strongly promoted by the spatial heterogeneity of habitats, which is often reduced by anthropogenic land transformation. Thus, it is decisive to understand the combined effects of these global change drivers on insect activity. Here, we assessed the movement activity of six insect species (from three orders) in response to heat stress using a unique tracking approach via radio frequency identification. We tracked 465 individuals at the iDiv Ecotron across a temperature gradient up to 38.7°C. In addition, we varied microhabitat conditions by adding leaf litter from four different tree species to the experimental units, either spatially separated or well mixed. Our results show opposing effects of heat extremes on insect activity depending on the microhabitat conditions. The insect community significantly decreased its activity in the mixed litter scenario, while we found a strong positive effect on activity in the separated litter scenario. We hypothesize that the simultaneous availability of thermal refugia as well as resources provided by the mixed litter scenario allows animals to reduce their activity and save energy in response to heat stress. Contrary, the spatial separation of beneficial microclimatic conditions and resources forces animals to increase their activity to fulfill their energetic needs. Thus, our study highlights the importance of habitat heterogeneity on smaller scales, because it may buffer the consequences of extreme temperatures of insect performance and survival under global change.

The iDiv Ecotron-A flexible research platform for multitrophic biodiversity research.

Across the globe, ecological communities are confronted with multiple global environmental change drivers, and they are responding in complex ways ranging from behavioral, physiological, and morphological changes within populations to changes in community composition and food web structure with consequences for ecosystem functioning. A better understanding of global change-induced alterations of multitrophic biodiversity and the ecosystem-level responses in terrestrial ecosystems requires holistic and integrative experimental approaches to manipulate and study complex communities and processes above and below the ground. We argue that mesocosm experiments fill a critical gap in this context, especially when based on ecological theory and coupled with microcosm experiments, field experiments, and observational studies of macroecological patterns. We describe the design and specifications of a novel terrestrial mesocosm facility, the iDiv Ecotron. It was developed to allow the setup and maintenance of complex communities and the manipulation of several abiotic factors in a near-natural way, while simultaneously measuring multiple ecosystem functions. To demonstrate the capabilities of the facility, we provide a case study. This study shows that changes in aboveground multitrophic interactions caused by decreased predator densities can have cascading effects on the composition of belowground communities. The iDiv Ecotrons technical features, which allow for the assembly of an endless spectrum of ecosystem components, create the opportunity for collaboration among researchers with an equally broad spectrum of expertise. In the last part, we outline some of such components that will be implemented in future ecological experiments to be realized in the iDiv Ecotron.

Ecotrons: Powerful and versatile ecosystem analysers for ecology, agronomy and environmental science.

Ecosystems integrity and services are threatened by anthropogenic global changes. Mitigating and adapting to these changes require knowledge of ecosystem functioning in the expected novel environments, informed in large part through experimentation and modelling. This paper describes 13 advanced controlled environment facilities for experimental ecosystem studies, herein termed ecotrons, open to the international community. Ecotrons enable simulation of a wide range of natural environmental conditions in replicated and independent experimental units while measuring various ecosystem processes. This capacity to realistically control ecosystem environments is used to emulate a variety of climatic scenarios and soil conditions, in natural sunlight or through broad-spectrum lighting. The use of large ecosystem samples, intact or reconstructed, minimizes border effects and increases biological and physical complexity. Measurements of concentrations of greenhouse trace gases as well as their net exchange between the ecosystem and the atmosphere are performed in most ecotrons, often quasi continuously. The flow of matter is often tracked with the use of stable isotope tracers of carbon and other elements. Equipment is available for measurements of soil water status as well as root and canopy growth. The experiments ran so far emphasize the diversity of the hosted research. Half of them concern global changes, often with a manipulation of more than one driver. About a quarter deal with the impact of biodiversity loss on ecosystem functioning and one quarter with ecosystem or plant physiology. We discuss how the methodology for environmental simulation and process measurements, especially in soil, can be improved and stress the need to establish stronger links with modelling in future projects. These developments will enable further improvements in mechanistic understanding and predictive capacity of ecotron research which will play, in complementarity with field experimentation and monitoring, a crucial role in exploring the ecosystem consequences of environmental changes.

Invertebrate Decline Leads to Shifts in Plant Species Abundance and Phenology.

Climate and land-use change lead to decreasing invertebrate biomass and alter invertebrate communities. These biotic changes may affect plant species abundance and phenology. Using 24 controlled experimental units in the iDiv Ecotron, we assessed the effects of invertebrate decline on an artificial grassland community formed by 12 herbaceous plant species. More specifically, we used Malaise traps and sweep nets to collect invertebrates from a local tall oatgrass meadow and included them in our Ecotron units at two different invertebrate densities: 100% (no invertebrate decline) and 25% (invertebrate decline of 75%). Another eight EcoUnits received no fauna and served as a control. Plant species abundance and flowering phenology was observed weekly over a period of 18 weeks. Our results showed that invertebrate densities affected the abundance and phenology of plant species. We observed a distinct species abundance shift with respect to the invertebrate treatment. Notably, this shift included a reduction in the abundance of the dominant plant species, Trifolium pratense, when invertebrates were present. Additionally, we found that the species shifted their flowering phenology as a response to the different invertebrate treatments, e.g. with decreasing invertebrate biomass Lotus corniculatus showed a later peak flowering time. We demonstrated that in addition to already well-studied abiotic drivers, biotic components may also drive phenological changes in plant communities. This study clearly suggests that invertebrate decline may contribute to already observed mismatches between plants and animals, with potential negative consequences for ecosystem services like food provision and pollination success. This deterioration of ecosystem function could enhance the loss of insects and plant biodiversity.

Mobile larynx in Mongolian gazelle: Retraction of the larynx during rutting barks in male Mongolian gazelle (Procapra gutturosa Pallas, 1777).

This study provides the first evidence of pronounced temporary laryngeal descent in a bovid species. An elaborate acoustic display is prominent in male courtship behavior of polygynous Mongolian gazelle. During rut, rounding up of females is accompanied by continuous head-up barking by dominant males. Throughout the rut their evolutionarily enlarged larynx descends to a low mid-neck resting position. In the course of each bark the larynx is additionally retracted toward the sternum by 30% of the resting vocal tract length. A geometric model of active larynx movements was constructed by combining results of video documentation, dissection, skeletonization, and behavioral observation. The considerable distance between resting position and maximal laryngeal descent suggests a backward tilting of the hyoid apparatus and an extension of the thyrohyoid connection during the retraction phase. Return to the resting position is effected by strap muscles and by the elastic recoil of the pharynx and the thyrohyoid connection. An intrapharyngeal inflation of the peculiar palatinal pharyngeal pouch of adult males is inferred from a short-time expansion of the ventral neck region rostral to the laryngeal prominence. The neck of adult dominant males is accentuated by long gray guard hairs during the rut. The passive swinging of the heavy larynx of adult males during locomotion gives the impression of a handicap imposed on rutting males. Apparently, this disadvantage becomes outweighed by the profits for reproductive success.

Nordic rattle: the hoarse vocalization and the inflatable laryngeal air sac of reindeer (Rangifer tarandus).

Laryngeal air sacs have evolved convergently in diverse mammalian lineages including insectivores, bats, rodents, pinnipeds, ungulates and primates, but their precise function has remained elusive. Among cervids, the vocal tract of reindeer has evolved an unpaired inflatable ventrorostral laryngeal air sac. This air sac is not present at birth but emerges during ontogenetic development. It protrudes from the laryngeal vestibulum via a short duct between the epiglottis and the thyroid cartilage. In the female the growth of the air sac stops at the age of 2-3 years, whereas in males it continues to grow up to the age of about 6 years, leading to a pronounced sexual dimorphism of the air sac. In adult females it is of moderate size (about 100 cm3), whereas in adult males it is large (3000-4000 cm3) and becomes asymmetric extending either to the left or to the right side of the neck. In both adult females and males the ventral air sac walls touch the integument. In the adult male the air sac is laterally covered by the mandibular portion of the sternocephalic muscle and the skin. Both sexes of reindeer have a double stylohyoid muscle and a thyroepiglottic muscle. Possibly these muscles assist in inflation of the air sac. Head-and-neck specimens were subjected to macroscopic anatomical dissection, computer tomographic analysis and skeletonization. In addition, isolated larynges were studied for comparison. Acoustic recordings were made during an autumn round-up of semi-domestic reindeer in Finland and in a small zoo herd. Male reindeer adopt a specific posture when emitting their serial hoarse rutting calls. Head and neck are kept low and the throat region is extended. In the ventral neck region, roughly corresponding to the position of the large air sac, there is a mane of longer hairs. Neck swelling and mane spreading during vocalization may act as an optical signal to other males and females. The air sac, as a side branch of the vocal tract, can be considered as an additional acoustic filter. Individual acoustic recognition may have been the primary function in the evolution of a size-variable air sac, and this function is retained in mother-young communication. In males sexual selection seems to have favoured a considerable size increase of the air sac and a switch to call series instead of single calls. Vocalization became restricted to the rutting period serving the attraction of females. We propose two possibilities for the acoustic function of the air sac in vocalization that do not exclude each other. The first assumes a coupling between air sac and the environment, resulting in an acoustic output that is a combination of the vocal tract resonance frequencies emitted via mouth and nostrils and the resonance frequencies of the air sac transmitted via the neck skin. The second assumes a weak coupling so that resonance frequencies of the air sac are lost to surrounding tissues by dissipation. In this case the resonance frequencies of the air sac solely influence the signal that is further filtered by the remaining vocal tract. According to our results one acoustic effect of the air sac in adult reindeer might be to mask formants of the vocal tract proper. In other cervid species, however, formants of rutting calls convey essential information on the quality of the sender, related to its potential reproductive success, to conspecifics. Further studies are required to solve this inconsistency.

Directionality of dog vocalizations.

The directionality patterns of sound emission in domestic dogs were measured in an anechoic environment using a microphone array. Mainly long-distance signals from four dogs were investigated. The radiation pattern of the signals differed clearly from an omnidirectional one with average differences in sound-pressure level between the frontal and rear position of 3-7 dB depending from the individual. Frequency dependence of directionality was shown for the range from 250 to 3200 Hz. The results indicate that when studying acoustic communication in mammals, more attention should be paid to the directionality pattern of sound emission.