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Atlas of the Hoverflies of Greece

(Diptera: Syrphidae)

Ante Vujić, Martin Speight, Michael Edwin de Courcy Williams, Santos Rojo, Gunilla Ståhls, Snežana Radenković, Laura Likov, Marija Miličić, Celeste Pérez-Bañón, Steven Falk and Theodora Petanidou

The Atlas of the Hoverflies of Greece is the first of a kind within the Mediterranean region. It is the result of decades of research, many travels into the fascinating habitats of Greece (a biodiversity hotspot), visits to world museums, and many people’s passion for hoverflies.
The Atlas is a concise presentation of all 418 hoverfly species for Greece known so far. The species are documented with photos and distribution GIS-maps and they are preceded by a general introduction on the hoverflies and Greek nature, and a generic key.
The Atlas of the Hoverflies of Greece is a handbook for insect aficionados, students and teachers, everyone interested in nature, and managers and conservationists aiming at raising public awareness of a nature nowadays threatened more than ever.

Nitzan Segev, Efrat Gavish-Regev and Oded Berger-Tal

Antlions (Neuroptera) are a group of sit-and-wait predator insects, with some species further specializing in digging conical pit-traps in the ground in order to catch prey. Studies on antlions’ predators are scarce with only few generalist predators known to feed on them. Here we report for the first time on field observations of antlions’ predation by three scorpion genera. We suggest that scorpions may be common predators of antlions, at least in the hyper-desert environment of southern Israel. The effects of predation risk on the behavior of sit-and-wait and particularly on trap-building predators received little attention in the literature. In light of our observations, we posit that predation risk must be taken into serious consideration in future research of antlions in particular, and sit-and-wait predators in general.

Colin M. Wright, James L.L. Lichtenstein, Lauren P. Luscuskie, Graham A. Montgomery, Sara Geary, Jonathan N. Pruitt, Noa Pinter-Wollman and Carl N. Keiser

Social spiders are thought to predominantly receive information about their environment through vibrational cues. Thus, group living introduces the challenge of distinguishing useful vibrational information from the background noise of nestmates. Here we investigate whether spatial proximity between colony-mates may allow social spiders (Stegodyphus dumicola) to reduce background noise that might obstruct vibrational information from prey. To do so, we constructed experimental colonies and measured whether the number of spiders in proximity to one another whilst resting could predict the number of spiders that participated in prey capture. Additionally, we exposed spider colonies to five different simulated vibrational cues mimicking prey to determine which cue types spiders were most responsive to. We found that the number of spiders huddled together prior to foraging trials was positively correlated with the number of spiders participating in collective foraging. Furthermore, colonies responded more quickly to pulsed vibrational cues over other types of vibrational patterns. Together these data reveal that both social interactions and prey cues shape how social sit-and-wait predators experience and respond to their environment.

Vesna Klokočovnik, Eva Veler and Dušan Devetak

The study focused on the behaviour of sit-and-wait antlion larvae in interspecific interactions. Antlion larvae usually occur in clusters with a high density of individuals; therefore, competition can be intense. We observed two abundant antlion species, E. nostras and M. formicarius, which co-occur in some habitats. In a simple habitat choice experiment where substrates differed according to sand particle size, we found that E. nostras exhibited dominance over M. formicarius. Most E. nostras larvae remained in the more suitable substrate and constructed pits, while all the M. formicarius larvae moved out of the suitable area, and did not build pits. In the second experiment, we observed the characteristics of the pit-fall traps and scored the occurrence of larval relocation in relation to interactions and in the control group, where larvae were kept in containers separately. In interactions, the larvae of E. nostras constructed smaller pits, but pit enlargement was greater in comparison to the control group. M. formicarius larvae constructed similar sized pits in both groups; however, enlargement was greater in the control group. Relocation of larvae occurred only during interactions. In direct interactions, we found 15 behavioural patterns, which are described in detail for the first time. In the presence of a competitor, larvae showed intense territorial behaviour. We recorded several behavioural patterns during larval confrontation, and interestingly, intraguild predation rarely occurred. In most cases, E. nostras larvae outcompete M. formicarius, which was evident from the larger pits and the rate of pit-construction.

Sean J. Blamires

Traps are rarely used by animals, despite the plausible benefits of broadening the number and diversity of prey that sit-and-wait foragers might be able to capture. The most well-known trap building sit-and-wait foragers are among the invertebrates, i.e. antlions, wormlions, glow worms, caddisflies, and spiders. A plausible hypothesis for the paucity of trap building by other animals is that biomechanical limitations render them inefficient or ineffective at catching sufficient prey. Here I examined the literature to make a valued judgement about the validity of this hypothesis. It appears that antlion and wormlion pit traps cannot catch and retain the largest prey they might expect to encounter. Arachnacampa glowworm traps are functionally efficient, facilitated by the animal’s bioluminescence. Nevertheless they only function in very moist or humid conditions. Caddisfly traps rely on flowing water to be able to capture their prey. Spiders are exceptional in developing a wide range of prey trapping strategies, from webs with dry adhesives, to sticky orb webs, to modified orb webs, e.g. elongated “ladder” webs, to webs with additional structures, and web aggregations. Some spiders have even redesigned their webs to minimize the high prey escape rates associated with web two dimensionality. These webs nevertheless are constructed and used at specific costs. While hard data across all of the invertebrate predators is lacking, there seems to be credence in the hypothesis that the biomechanical limitations placed on trap functionality can explain their limited use among animals.

Maja Peryga and Krzysztof Miler

Some terrestrial leeches mate by entwining the anterior ends of their bodies and then copulating. Here, we report first observations of a similar behavioral pattern in Haemadipsa picta terrestrial leeches from Malaysian Borneo. However, because the observed pattern can be easily induced artificially with no clear evidence of copulation, we suggest that it may serve another function, particularly in H. picta. We hypothesize that the wrestling behavior, as we term it, may be a ritualized aggressive display driven by competition for ambush location. Haemadipsid fauna of the region is poorly studied, therefore our observations extend limited knowledge about these leeches and open interesting research avenues for the study of the wrestling behavior.

Aaron M. Ellison

Carnivorous plants are pure sit-and-wait predators: they remain rooted to a single location and depend on the abundance and movement of their prey to obtain nutrients required for growth and reproduction. Yet carnivorous plants exhibit phenotypically plastic responses to prey availability that parallel those of non-carnivorous plants to changes in light levels or soil-nutrient concentrations. The latter have been considered to be foraging behaviors, but the former have not. Here, I review aspects of foraging theory that can be profitably applied to carnivorous plants considered as sit-and-wait predators. A discussion of different strategies by which carnivorous plants attract, capture, kill, and digest prey, and subsequently acquire nutrients from them suggests that optimal foraging theory can be applied to carnivorous plants as easily as it has been applied to animals. Carnivorous plants can vary their production, placement, and types of traps; switch between capturing nutrients from leaf-derived traps and roots; temporarily activate traps in response to external cues; or cease trap production altogether. Future research on foraging strategies by carnivorous plants will yield new insights into the physiology and ecology of what Darwin called “the most wonderful plants in the world”. At the same time, inclusion of carnivorous plants into models of animal foraging behavior could lead to the development of a more general and taxonomically inclusive foraging theory.

Inon Scharf, Yehonatan Samocha and Noa Katz

When choosing among several potential habitats, animals should strive to choose the habitat that provides the highest fitness. When animals choose habitats that do not provide the best possible fitness, there is a mismatch between habitat preference and performance. A common reason is that of limited information or perceptual range. Sit-and-wait predators are particularly deficient in information, due to spending most of their time in ambushing prey and, when they do travel, they cover only short distances. Here, we studied the effect of placing pit-building wormlions at a short distance from their preferred microhabitats, on the likelihood of them choosing it. When placed on the border between deep and shallow sand, fine and coarse sand, or dry and wet sand, wormlions chose the former in each case in vast proportions. However, the ability of wormlions to recognize and/or choose favorable habitats decreased sharply with distance, suggesting that they are limited in their perceptual range. We next examined whether wormlions relocate their pits following a continuous disturbance arriving always from the same side. Although they did so, their movement direction was unrelated to the direction of the disturbance. The preference of wormlions for a favorable, shaded microhabitat depended greatly on the location of a conspecific competitor: the favorable microhabitat was more attractive for wormlions when a competitor was more distant from it than when the competitor was located on the border between the favorable and unfavorable microhabitats. Our findings support previous studies indicating the limited perceptual range of pit-building predators.

Alejandro G. Farji-Brener and Sabrina Amador-Vargas

The physical structures built by animals are considered extended phenotypes that reflect how organisms make decisions and deal with changes in their biotic and abiotic environment. We summarize the results of several studies on Myrmeleon crudelis, a neuropteran larva that digs pit-traps in the soil to capture small arthropods (mostly ants) in the tropical dry forests of Costa Rica. Specifically, we showed how this species responds to varying biotic and abiotic conditions with changes in the design and/or location of its pit traps. Several experiments and field comparisons indicate that: 1) antlions adjust the pit design according to the abundance and type of prey. When prey is scarce, antlions increased trap diameter, an architectural adjustment that enhances the probability of prey encounter. Antlions that experienced high prey abundance, but the prey easily escaped, then increased pit depth, an adjustment that increases the chance of prey retention; 2) soil compaction strongly reduced pit-trap size and abundance; 3) antlions preferred soils with high proportion of fine-particle size to build pits. In fine-grained soil, pit-traps are larger and more efficient to capture prey than traps in coarse-grained soils; and 4) pit-traps may also be affected through indirect effects of soil structure and vegetation cover. Areas with fine-soil presented less plant cover, and plant cover could be beneficial for antlions because it acts as a shelter against direct sunlight and rainfall, or it may represent a cost because it is a source of leaflitter falling in the pits. The works summarized here how trap-building predators can exhibit considerable flexibility in trap construction in response to various biotic and abiotic factors, emphasizing how the study of extended phenotypes can be a useful approach to better understand the flexibility of foraging behaviors.

Dulce Rodríguez-Morales, Helena Ajuria-Ibarra, Laura T. Hernández-Salazar, Víctor Rico-Gray, José G. García-Franco and Dinesh Rao

Predation is one of the main interactions between organisms and one of the primary selective agents for their survival. Both prey and predators have developed different strategies and characteristics that allow them to be conspicuous or go undetected. In sit and wait predators, their shape and appearance are important factors that allow them to remain undetected by their potential prey. Sit and wait predators such as crab spiders are difficult to identify when they sit on flowers or areas of flowers with colors similar to the color of their bodies. In this study, we aimed to determine if insects can recognize the morphology and color polymorphism of crab spiders by evaluating the response of flower visitors. We quantified the visits and approaches of floral visitors to the flowerheads of Palafoxia lindenii with spider morphology and color polymorphism treatments. Our results show that insects in general, and bees in particular, avoid visiting flowers with a real spider or a spider model and visit vacant flowers more frequently. In the case of the color polymorphism, insects approached flowerheads with spiders with a similar frequency independently of the color of the spiders, but did not visit them. Insects appeared to identify spiders through their morphological characteristics rather than their color characteristics, since flower visitors did not discriminate between the evaluated spider colors (white, lilac, and purple). This study emphasizes the differential response of different insect prey to the presence, color, and morphology of sit and wait predators.