Cowards or clever guys: an alternative nest defence strategy employed by shrikes against magpies.

Red-backed shrikes (Lanius collurio) show a substantial variability in their nest defence behaviour, which usually follows the rules of optimal parental behaviour, vigorously attacking egg and chick predators and only passively guarding against harmless animals. Nevertheless, shrikes hesitate to attack the Eurasian magpie (Pica pica), which specializes in plundering passerine nests. Our previous studies have suggested that this behaviour may be the result of an alternative defence strategy, relying on nest crypsis. To test this hypothesis, at the shrike nests, we presented a magpie dummy associated with playbacks drawing the predators' attention to the presence of the nest. We predicted that the presentation of a magpie dummy associated with shrike alarm calls moves the parents to action, causing them to chase the magpie away from the nest. We showed that the presence of a magpie dummy associated with shrike alarm calls elicits a significantly more active response in shrike parents compared to a magpie dummy associated with neutral song. Parents actively moved around the dummy and produced alarm calls; nevertheless, most of the tested pairs hesitated to attack the dummy. We may conclude that the low nest defence activity of shrike parents towards magpie dummy was partly the result of an alternative strategy, which may be cancelled out by alerting the predator to the location of the nest; nevertheless, shrikes seem to be afraid of the magpie and hesitate to attack it physically.

A kestrel without hooked beak and talons is not a kestrel for the red-backed shrike (Lanius collurio).

Birds are strongly motivated to recognize various predators to secure survival and reproductive success. Thus, predator recognition provides a useful tool for the investigation of the process of discrimination and categorization in non-trained animals. The most important role in this process is usually attributed to the prominent unique features shared by all members within the wider category of predators (sharp teeth or beaks, claws, talons and conspicuous eyes). However, birds are also able to discriminate between particular predator species according to their species-specific features (based mainly on colouration). We manipulated general raptor salient features (hooked beak, talons with claws and eyes with supraorbital ridges) and/or species-specific plumage colouration on the dummy of a Eurasian kestrel (Falco tinnunculus) to investigate their importance in the recognition process of the red-backed shrikes (Lanius collurio) during nest defence. We showed that the presence of general raptor salient features is necessary for the recognition of a predator. The kestrel dummy with natural species-specific colouration with altered raptor salient features was treated by shrikes as a harmless bird. Nevertheless, pronounced changes in colouration also prevented successful recognition of the dummy as a kestrel, even when raptor salient features of a raptor were present.

Object categorization by wild-ranging birds in nest defence.

Despite object categorization being an important ability for the survival of wild animals, the principles behind this ability have been only scarcely studied using wild-ranging, untrained animals. Reiterating our previous study undertaken with wild-ranging titmice on winter feeders (Nováková et al. Behav Process 143:7-12, 2017), we aimed to test two hypotheses of object recognition proposed by animal psychology studies: the particulate feature theory and recognition by components in the methodological paradigm of nest defence. We tested whether the parents of the red-backed shrike (Lanius collurio) recognize the dummies of the common kestrel (Falco tinnunculus), which is a potential predator of large chicks or fledglings, as a threat in case when their body parts are scrambled. The kestrel dummy was presented with the head at the top, in the middle, and at the bottom of the body. We showed that the shrikes did not consider dummies of a kestrel with an inappropriately placed head as a threat to the nest and attacked it equally scarcely as the harmless control. These results support the theory of recognition by components, presuming that the mutual position of body parts is essential for appropriate recognition of the object. When the body parts were scrambled, most of shrikes were not able to identify the kestrel in such an object despite all local features (eye, beak, colouration, and claws) being present. Nevertheless, shrikes did not consider the scrambled dummies as completely harmless, because they fed their chicks in their presence significantly less often than in the presence of harmless control.

A Novel Class of Defensive Compounds in Harvestmen: Hydroxy-γ-Lactones from the Phalangiid Egaenus convexus.

When threatened, the harvestman Egaenus convexus (Opiliones: Phalangiidae) ejects a secretion against offenders. The secretion originates from large prosomal scent glands and is mainly composed of two isomers of 4-hydroxy-5-octyl-4,5-dihydro-3H-furan-2-one (1), a ß-hydroxy-γ-lactone. The compounds were characterized by GC-MS of their microreaction derivatives, HRMS, and NMR. After the synthesis of all four possible stereoisomers of 1, followed by their separation by chiral-phase GC, the absolute configurations of the lactones in the Egaenus secretion was found to be (4S,5R)-1 (90%) and (4S,5S)-1 (10%). Hydroxy-γ-lactones represent a new class of exocrine defense compounds in harvestmen.

Subtle Chemical Variations with Strong Ecological Significance: Stereoselective Responses of Male Orchid Bees to Stereoisomers of Carvone Epoxide.

Different enantiomers of chiral compounds within floral perfumes usually trigger distinct responses in insects; however, this has frequently been neglected in studies investigating semiochemicals in plant-pollinator interactions. Approximately 1000 neotropical plants produce floral perfumes as the only reward for pollinators, i.e. male euglossine bees. The chiral compound carvone epoxide is a key component of the scent bouquet of many perfume-rewarding plants that are pollinated by males of Eulaema. Here, we tested the biological activity of the four carvone epoxide stereoisomers to four Eulaema species occurring in the Atlantic Rainforest of NE-Brazil. We determined the stereochemistry of carvone epoxide in the floral scent of several Catasetum species, tested whether the antennae of bees respond differentially to these stereoisomers and investigated if there is a behavioural preference for any of the stereoisomers. We found that 1) Catasetum species emit only the (-)-trans-stereoisomer of carvone epoxide, 2) for E. atleticana and E. niveofasciata antennal responses to the (-)-trans-carvone epoxide were significantly stronger than those to (-)-cis-carvone epoxide, 3) the strength and pattern of antennal responses to all 4 stereoisomers (separately tested) did not differ among Eulaema species, and 4) there were significant differences in attractiveness of the four stereoisomers to the bees species with the (-)-trans-stereoisomer being particularly attractive. We assume (-)-trans-carvone epoxide to be the dominant isomer in perfume-rewarding plants pollinated by Eulaema. The universal occurrence of carvone epoxide in Catasetum species pollinated by Eulaema, suggests that this compound has evolved in perfume-rewarding as a specific attractant for Eulaema bees as pollinators.

The effect of social learning on avoidance of aposematic prey in juvenile great tits (Parus major).

Social learning plays an important role in acquiring new foraging skills and food preferences in many bird species but its potential role in learning to avoid aposematic prey has never been studied. We tested the effect of social learning on the acquisition of avoidance of aposematic insect prey (firebug Pyrrhocoris apterus; Heteroptera) in juvenile, hand-reared great tits (Parus major). Behaviour towards aposematic prey was compared between two groups of birds: (1) the observers that were, prior to encounter with firebugs, allowed to watch the experienced conspecific demonstrator repeatedly refuse to attack the prey, and (2) the control birds that lacked this opportunity. Observing an experienced demonstrator was not sufficient for learning complete avoidance, because birds from both groups attacked at least the first firebug they had encountered in avoidance training. However, the opportunity to observe the avoidance behaviour of another bird significantly increased the rate of subsequent individual learning of observers in comparison with control birds. Social learning also decreased mortality of firebugs killed by the birds during the avoidance learning. Socially enhanced learning to avoid aposematic prey might be a mechanism important especially for naive juvenile birds learning from their parents, but it could also enhance learning in adults from their more experienced flock mates. Because social learning of avoidance may also lead to decreased mortality of aposematic prey, its effect should be taken into account in scenarios considering evolution and maintenance of prey warning signals.

Trophoblast subtype-specific EGFR/ERBB4 expression correlates with cell cycle progression and hyperplasia in complete hydatidiform moles.

STUDY QUESTION: Do trophoblast subtypes differ in their expression of erythroblastic leukaemia viral oncogene homologue (ERBB) receptor family members and responsiveness towards specific growth factor ligands? SUMMARY ANSWER: Our data reveal a reciprocal expression pattern of epidermal growth factor receptor (EGFR)/ERBB4 in proliferative and ERBB2/ERBB3 in invasive trophoblast subtypes, as well as a restricted responsiveness to epidermal growth factor (EGF) and heparin-binding (HB)-EGF. WHAT IS KNOWN ALREADY: EGFR is expressed by villous cytotrophoblasts (vCTBs), but absent from extravillous trophoblasts (EVTs), which specifically up-regulate ERBB2. STUDY DESIGN, SIZE, DURATION: Tissue samples of human first trimester placentae (n = 50) and deciduae (n = 5) obtained from elective pregnancy terminations were used to study trophoblast subtype-specific ERBB receptor expression and responsiveness to recombinant human EGF and HB-EGF. Age-matched complete hydatidiform mole (CHM) placentae (n = 12) were assessed for EGFR and ERBB4 expression in proliferation-competent regions. PARTICIPANTS/MATERIALS, SETTING, METHODS: ERBB receptor expression was analysed in primary trophoblast cell isolates by means of microarray, quantitative real-time PCR and western blotting, as well as immunofluorescence stainings of placental and decidual tissue sections. EGF and HB-EGF were tested for their potential to activate ERBB receptors in purified EGFR(+) and HLA-G(+) trophoblasts. 5-Ethynyl-2'-deoxyuridine incorporation assays were performed to study the effect of both ligands on the proliferative capacity of primary trophoblasts as well as of vCTBs and proximal cell column trophoblasts (pCCTs) in placental floating explants. Finally, the average number of EGFR(+) vCTB and pCCT layers was determined in CHM placentae and compared with healthy age-matched controls. MAIN RESULTS AND THE ROLE OF CHANCE: Proliferative vCTBs and pCCTs co-express EGFR and ERBB4, but are devoid of ERBB2 and ERBB3. In contrast, HLA-G(+) trophoblast subtypes exhibit an EGFR/ERBB4(-) and ERBB2/ERBB3(+) phenotype. EGF and HB-EGF activate EGFR, ERBB4, AKT and extracellular signal-regulated kinase 1/2 in EGFR(+) primary trophoblasts; however, they do not show an effect on HLA-G(+) EVTs. Both ligands strongly induce cell cycle progression in primary trophoblasts (P < 0.05) and placental explant-associated vCTBs (P < 0.05) and pCCTs (P < 0.05). Notably, EGFR(+) vCTB (P < 0.0001) and pCCT (P < 0.0001) layers are significantly expanded in CHM placentae when compared with healthy controls. LIMITATIONS, REASONS FOR CAUTION: Cells were removed from their physiological context and may therefore respond differently to various stimuli. WIDER IMPLICATIONS OF THE FINDINGS: In this study we define EGFR as a marker for proliferative trophoblast subtypes within the human placenta. Manipulation of EGFR signalling might thus offer a promising therapeutic avenue for the treatment of molar pregnancies associated with trophoblast hyperplasia. STUDY FUNDING/COMPETING INTERESTS: This study was supported by the Austrian Science Fund (grant P-25187-B13 to J.P.). There are no competing interests to declare.

Surface texture and priming play important roles in predator recognition by the red-backed shrike in field experiments.

We compared the responses of the nesting red-backed shrikes (Lanius collurio) to three dummies of a common nest predator, the Eurasian jay (Garrulus glandarius), each made from a different material (stuffed, plush, and silicone). The shrikes performed defensive behaviour including attacks on all three dummies. Nevertheless, the number of attacks significantly decreased from the stuffed dummy through the plush dummy and finally to the silicone dummy. Our results show that wild birds use not only colours but also other surface features as important cues for recognition and categorization of other bird species. Moreover, the silicone dummy was attacked only when presented after the stuffed or plush dummy. Thus, we concluded that the shrikes recognized the jay only the stuffed (with feathered surface) and plush (with hairy surface) dummies during the first encounter. Recognition of the silicon dummy (with glossy surface) was facilitated by previous encounters with the more accurate model. This process resembles the effect of perceptual priming, which is widely described in the literature on humans.

The role of key features in predator recognition by untrained birds.

The most important role in the recognition and categorization of predators (as well as other animals) is usually attributed to so-called key features. Under laboratory conditions, we tested the role of yellow eyes (specific for the genus Accipiter in European raptors) and hooked beak (common for all European birds of prey) in the recognition of the sparrowhawk (Accipiter nisus) by untrained great tits (Parus major) caught in the wild. Using wooden dummies, we interchanged either one of these potential key features or the body of the sparrowhawk (predator) and domestic pigeon (harmless bird). The tested tits showed three types of behaviour in the presence of the dummies: fear, interest without fear, and lack of interest. Eye interchange lowered fear of the sparrowhawk, but did not cause fear of the pigeon. Beak interchange did not lower fear of the sparrowhawk. Eye interchange caused increased interest in both species. Thus, a specific sparrowhawk feature is necessary for correct sparrowhawk dummy recognition but a general raptor feature is not. On the other hand, a specific sparrowhawk feature on a pigeon dummy is not enough to prompt sparrowhawk recognition. Thus, key features play an important, but not exclusive, role in predator recognition. An increased interest in some of the modified dummies implies that the tits have a general concept of a sparrowhawk. The individual variability in behaviour of tits is discussed.

Red-backed shrike (Lanius collurio) versus common cuckoo (Cuculus canorus): An example of ineffective cuckoo-hawk mimicry.

The red-backed shrike (Lanius collurio) used to be one of the most common hosts of the common cuckoo (Cuculus canorus). Nevertheless, during the last 30 years, there is increasing evidence from Central Europe that the occurrence of cuckoo chicks in shrike nests has become scarcer, and that in some locations they have disappeared completely. Multiple hypotheses have been suggested to explain this abandonment. Here, we test the hypothesis that shrikes vigorously attack adult cuckoos, potentially resulting in ineffective parasitism. Adult common cuckoos resemble in appearance the Eurasian sparrowhawk (Accipiter nisus), a common predator of small passerines. One hypothesis presumes that the cuckoo has evolved this mimicry to avoid attack by small passerines when searching for their nests. Our results show that shrikes defending their nests attacked cuckoos very vigorously, more often, and more intensively than they did sparrowhawks. In the presence of a sparrowhawk dummy, parent shrikes only produced alarm calls and flew over the dummy. This suggests that cuckoo-hawk mimicry is ineffective in the case of shrikes and that they attack them much more often than they do any other presented intruder. Therefore, this activity could possibly result in the abandonment of shrikes as potential hosts for cuckoos.

Local Insect Availability Partly Explains Geographical Differences in Floral Visitor Assemblages of Arum maculatum L. (Araceae).

Geographical variation in abundance and composition of pollinator assemblages may result in variable selection pressures among plant populations and drive plant diversification. However, there is limited knowledge on whether differences in local visitor and pollinator assemblages are the result of site-specific strategies of plants to interact with their pollinators and/or merely reflect the pollinator availability at a given locality. To address this question, we compared locally available insect communities obtained by light-trapping with assemblages of floral visitors in populations of Arum maculatum (Araceae) from north vs. south of the Alps. We further investigated whether and how the abundance of different visitors affects plants' female reproductive success and examined the pollen loads of abundant visitors. Local insect availability explained inter-regional differences in total visitor abundance, but only partly the composition of visitor assemblages. Northern populations predominantly attracted females of Psychoda phalaenoides (Psychodidae, Diptera), reflecting the high availability of this moth fly in this region. More generalized visitor assemblages, including other psychodid and non-psychodid groups, were observed in the south, where the availability of P. phalaenoides/Psychodidae was limited. Fruit set was higher in the north than in the south but correlated positively in both regions with the abundance of total visitors and psychodids; in the north, however, this relationship disappeared when visitor abundances were too high. High pollen loads were recorded on both psychodids and other Diptera. We demonstrate for the first time that the quantitative assessment of floral visitor assemblages in relation to locally available insect communities is helpful to understand patterns of geographical variation in plant-pollinator interactions. This combined approach revealed that geographical differences in floral visitors of A. maculatum are only partly shaped by the local insect availability. Potential other factors that may contribute to the geographical pattern of visitor assemblages include the region-specific attractiveness of this plant species to flower visitors and the population-specific behavior of pollinators.

Untrained birds' ability to recognise predators with changed body size and colouration in a field experiment.

BACKGROUND: During recognition process, multiple parameters of the encountered stimulus may play a role. Previous studies with wild birds identified the importance of several salient features (e.g., eyes, beak, prominent elements of colouration) which birds use to recognise other bird species, such as predators or nest parasites. In the present study, we observed the responses of passerines visiting winter feeders to stimuli in the form of dummies of Eurasian sparrowhawk which were modified in body size and/or colouration but always carried the salient features of raptors (hooked beak, talons) and one species-specific feature of the sparrowhawk (yellow eyes). In the vicinity of a feeder, we placed a dummy of an unmodified sparrowhawk, life-sized sparrowhawk with pigeon, great tit, or robin colouration, a small, great tit-sized sparrowhawk dummy with unmodified or pigeon colouration, or an unmodified pigeon dummy, which functioned as a harmless control. Then we measured how it affected the number of visits. RESULTS: We found that birds were less afraid of small dummies regardless of their colouration than they were of life-sized raptor dummies or even the pigeon dummy. This contrasts with the results of a previous laboratory experiment where great tits' reaction to small dummies was comparably fearful to their response to life-size dummies. In our experiment, birds were also not afraid of life-sized dummies with modified colouration except for a robin-coloured dummy, which caused an equally significant fear reaction as an unmodified sparrowhawk dummy. It is likely that this dummy resembled the colouration of a male sparrowhawk closely enough to cause this effect. CONCLUSIONS: Based on our observations, we conclude that birds use contextual features to evaluate the size of other birds. Distance and familiar reference points seem to play an important part in this process.

Predation by avian predators may have initiated the evolution of myrmecomorph spiders.

Myrmecomorphy is a strategy utilized by a variety of species, among which spiders are the most common. It is supposed that myrmecomorphy tends to be selected by predator avoidance of preying on ants rather than by blind ant workers. To date, this hypothesis has been tested mainly on invertebrate predators (mantises and spiders). We are the first to test whether an imperfect myrmecomorph spider (Phrurolithus festivus) gains protection against avian predators (wild adult great tits-Parus major) through its appearance. In a set of preferential trials, we showed that the ant model and the myrmecomorph spider are equally well protected against attack, though the attacked myrmecomorphs are usually eaten. This suggests that the mimicry of the myrmecomorph spiders is effective against avian predators and works in a Batesian manner. In this study, we have provided evidence toward the evolution of myrmecomorphy in response to selective pressure elicited by visually-oriented predators like birds.

Macroclimatic conditions as main drivers for symbiotic association patterns in lecideoid lichens along the Transantarctic Mountains, Ross Sea region, Antarctica.

Lecideoid lichens as dominant vegetation-forming organisms in the climatically harsh areas of the southern part of continental Antarctica show clear preferences in relation to environmental conditions (i.e. macroclimate). 306 lichen samples were included in the study, collected along the Ross Sea coast (78°S-85.5°S) at six climatically different sites. The species compositions as well as the associations of their two dominant symbiotic partners (myco- and photobiont) were set in context with environmental conditions along the latitudinal gradient. Diversity values were nonlinear with respect to latitude, with the highest alpha diversity in the milder areas of the McMurdo Dry Valleys (78°S) and the most southern areas (Durham Point, 85.5°S; Garden Spur, 84.5°S), and lowest in the especially arid and cold Darwin Area (~ 79.8°S). Furthermore, the specificity of mycobiont species towards their photobionts decreased under more severe climate conditions. The generalist lichen species Lecanora fuscobrunnea and Lecidea cancriformis were present in almost all habitats, but were dominant in climatically extreme areas. Carbonea vorticosa, Lecidella greenii and Rhizoplaca macleanii were confined to milder areas. In summary, the macroclimate is considered to be the main driver of species distribution, making certain species useful as bioindicators of climate conditions and, consequently, for assessing the consequences of climate change.

Floral Scents of a Deceptive Plant Are Hyperdiverse and Under Population-Specific Phenotypic Selection.

Floral scent is a key mediator in plant-pollinator interactions. However, little is known to what extent intraspecific scent variation is shaped by phenotypic selection, with no information yet in deceptive plants. In this study, we collected inflorescence scent and fruit set of the deceptive moth fly-pollinated Arum maculatum L. (Araceae) from six populations north vs. five populations south of the Alps, accumulating to 233 samples in total, and tested for differences in scent, fruit set, and phenotypic selection on scent across this geographic barrier. We recorded 289 scent compounds, the highest number so far reported in a single plant species. Most of the compounds occurred both north and south of the Alps; however, plants of the different regions emitted different absolute and relative amounts of scent. Fruit set was higher north than south of the Alps, and some, but not all differences in scent could be explained by differential phenotypic selection in northern vs. southern populations. This study is the first to provide evidence that floral scents of a deceptive plant are under phenotypic selection and that phenotypic selection is involved in shaping geographic patterns of floral scent in such plants. The hyperdiverse scent of A. maculatum might result from the imitation of various brood substrates of its pollinators.

Tits use amodal completion in predator recognition: a field experiment.

Amodal completion enables an animal to perceive partly concealed objects as an entirety, and to interact with them appropriately. Several studies, based upon either operant conditioning or filial imprinting techniques, have shown that various animals (both mammals and birds) can perform amodal completion. Before this study, the use of amodal completion by untrained animals in the recognition of objects had not been considered. Using two feeders, we observed in a field experiment the reaction of tits to the torso of a sparrowhawk (partly occluded or an 'amputated' dummy) in two different treatments (sparrowhawk torso vs. complete dummy pigeon; and torso vs. complete dummy sparrowhawk). It is clear that the birds considered the two torso variants as predators and kept away from both of them when the second feeder offered a 'pigeon' instead. On the other hand, when a 'complete sparrowhawk' was present on the second feeder, the number of visits to the occluded torso remained low; while the number of visits to the amputated one increased threefold. Birds risked perching near what was clearly an amputated torso; while the fear of a "hiding" (occluded) torso remained unchanged, when the second feeder did not provide a safe alternative. Such discrimination between torsos requires the ability for amodal completion. Our results demonstrate that in their recognition process, the birds not only use simple sign stimuli, but also complex cognitive functions.

Effect of the exploratory behaviour on a bird's ability to categorize a predator.

Despite the encountering of a predator always being extremely threatening, there is a significant plasticity among individuals in how they cope with such a situation. In laboratory experiments with wild-caught great tits (Parus major), we tested the effect of exploratory behaviour (performance in novel food, object and environment test, startle test) on the ability of individual birds to assess the threat represented by a predator. We presented a wooden dummy of the European sparrowhawk (Accipiter nisus), an extremely dangerous predator, and its visual modifications (chimeras), changing the beak or head to be non-threatening (those of a pigeon - Columba livia f. domestica). We showed that the differences between 'slow' and 'fast explorers' are not very distinct, but that 'slow explorers' generally tended to be more cautious in the presence of an unmodified sparrowhawk dummy, while the 'fast explorers' tended to observe the dummy. On the contrary, 'slow explorers' tended to treat both chimaeras (and the pigeon dummy as well) as less-threatening than 'fast explorers'. Since 'slow explorers' are usually considered to be more sensitive to environmental cues, it came as no surprise that most of them correctly assessed the unmodified sparrowhawk dummy as threatening, while they probably subjected the chimeras to a detailed inspection and were not confused by the presence of sparrowhawk features and assessed them as non-threatening.

Incomplete synchrony of inflorescence scent and temperature patterns in Arum maculatum L. (Araceae).

In many Araceae both scent and heat production are known to temporally vary throughout anthesis, and in several species strong scents are released for pollinator attraction when thermogenesis is also strong. However, it is not known whether the temporal patterns of both scent emission and temperature are strictly synchronous and, for example, reach their maxima at the same time. We studied Arum maculatum, a brood-site deceptive species attracting its moth fly pollinators with strong fetid scents, to study temporal patterns in scent emission and temperature during anthesis. Inflorescence scents were collected and analysed by dynamic headspace and gas chromatography-mass spectrometry (GC-MS) or by proton-transfer-reaction-time-of-flight mass spectrometry (PTR-TOFMS), and the temperature of the appendix, which is the heating osmophore during pollinator attraction, was recorded by a thermocouple. We overall found that scent emission and temperature patterns were strongly correlated. However, in none of the seven studied individuals was the highest amount of scent released at times with the maximum temperature difference. Thus, patterns of scent emission and temperature are somewhat asynchronous suggesting that high scent emission rates and temporal scent patterns in plants with thermogenesis cannot be solely explained by temperature patterns. This calls for more in-depth studies to better understand the interplay between scent emission and thermogenesis.

Object categorization by wild ranging birds-Winter feeder experiments.

The object categorization is only scarcely studied using untrained wild ranging animals and relevant stimuli. We tested the importance of the spatial position of features salient for categorization of a predator using wild ranging birds (titmice) visiting a winter feeder. As a relevant stimulus we used a dummy of a raptor, the European sparrowhawk (Accipiter nisus), placed at the feeding location. This dummy was designed to be dismantled into three parts and rearranged with the head in the correct position, in the middle or at the bottom of the dummy. When the birds had the option of visiting an alternative feeder with a dummy pigeon, they preferred this option to visiting the feeder with the dummy sparrowhawk with the head in any of the three positions. When the birds had the option of visiting an alternative feeder with an un-rearranged dummy sparrowhawk, they visited both feeders equally often, and very scarcely. This suggests that the titmice considered all of the sparrowhawk modifications as being dangerous, and equally dangerous as the un-rearranged sparrowhawk. The position of the head was not the most important cue for categorization. The presence of the key features was probably sufficient for categorization, and their mutual spatial position was of lower importance.

Do predator energy demands or previous exposure influence protection by aposematic coloration of prey?

Growing evidence exists that aposematic and toxic prey may be included in a predator's diet when the predator experiences physiological stress. The tree sparrow Passer montanus is known to have a significant portion of aposematic and toxic ladybirds in its natural diet. Here, we present experiments testing the attack and eating rate of the tree sparrow toward the invasive aposematic harlequin ladybird Harmonia axyridis. We wondered whether the sparrow's ability to prey on native ladybirds predisposes them to also prey on harlequin ladybirds. We compared the attack and eating rates of tree sparrows of particular age and/or experience classes to test for any changes during ontogeny (hand-reared × young wild-caught ×adult wild-caught) and with differing perceived levels of physiological stress (summer adult × winter adult). Winter adult tree sparrows commonly attacked and ate the offered ladybirds with no evidence of disgust or metabolic difficulties after ingestion. Naïve and wild immature tree sparrows attacked the ladybirds but hesitated to eat them. Adult tree sparrows caught in the summer avoided attacking the ladybirds. These results suggest that tree sparrows are able to cope with chemicals ingested along with the ladybirds. This pre-adaptation enables them to include ladybirds in their diet; though they commonly do this only in times of shortage in insect availability (winter). Young sparrows showed avoidance toward the chemical protection of the ladybirds.