Effects of defoliation and girdling on fruit production in Ilex aquifolium

1. The effects of defoliation and girdling on fruit production in European holly Ilex aquifolium were studied in a northern Spanish population. Three defoliation treatments (control, 50% and 100% leaf removal) were applied at branch level in 10 trees. Six branches were used per treatment in each tree and half of these branches were girdled (a ring of bark and cambium was removed from the branch base).
2. Leaf removal from ungirdled branches had little effect on fruit set, fruit production and reproductive allocation at branch level. However, these variables decreased as the extent of defoliation increased on girdled branches.
3. Fruit production in undefoliated branches did not differ between girdled and ungirdled branches, showing that girdled branches were autonomous for fruit production when undamaged. Mean fruit mass of girdled-100% defoliated branches averaged 8% of fruit mass produced by control branches.
4. Fruit production of ungirdled-100% defoliated branches did not differ from control branches, showing the importance of resource importation from other branches. Hence the branches may be regarded as only partially autonomous for fruit production.
5. Fruit production of girdled-50% defoliated branches was 42% of fruit production of control branches, while ungirdled-50% defoliated branches amounted to 70% of controls. These results also demonstrated resource importation but did not support the existence of photosynthetic compensatory mechanisms.
6. The ability of resource importation developed by fruiting branches might be a mechanism to reduce the effect of reproductive costs in this species.     

Carnivorous bats?

Only large bats can take large prey but size alone does not identify 'carnivorous bats' (those including small terrestrial vertebrates in their diets). Morphological data, including body mass, aspect ratio and relative wing loading, along with information about orientation and foraging strategies can be used to characterize a suite of features which identifies carnivorous bats. We use the available data to make predictions about which large Microchiroptera will be found to be carnivorous. A combination of morphological features including body mass (^0.017 kg), low aspect ratio (<6.3), and low relative wing loading (<36) significantly identifies carnivorous species from among other animal-eating forms. Some carnivorous species use short, low intensity, high frequency, broadband echolocation cells but rely on prey generated cues to locate their targets. Other carnivorous species are facultative echolocators. The available data lead to the prediction that Phyllostomus hastatus and Hipposideros diadema are not regularly carnivorous, while Otonycteris hemprichi may be. Large species with echolocation calls adapted for flutter detection (rhinolophids and hipposiderids) or those with long narrowband calls and high aspect ratio wings with high relative wing loading (for example molossids, some emballonurids and some vespertilionids) chase airborne prey in the open; neither of these approaches involves prey other than arthropods.     

Ethanol ingestion affects flight performance and echolocation in Egyptian fruit bats

Ethanol, a potential toxin for vertebrates, is present in all fleshy fruits and its content increases as the fruit ripens. Previously, we found that the marginal value of food for Egyptian fruit bats, Rousettus aegyptiacus, decreases when its ethanol content exceeds 1%. Therefore, we hypothesized that, if ingested, food containing >1% ethanol is toxic to these bats, probably causing inebriation that will affect flight and echolocation skills. We tested this hypothesis by flying Egyptian fruit bats in an indoor corridor and found that after ingesting ethanol-rich food bats flew significantly slower than when fed ethanol-free food. Also, the ingestion of ethanol significantly affected several variables of the bats’ echolocation calls and behavior. We concluded that ethanol can be toxic to fruit bats; not only does it reduce the marginal value of food, but it also has negative physiological effects on their ability to fly competently and on their calling ability.     

Experimental transmission of the parasitic flagellates Trypanosoma cruzi and Trypanosoma rangeli between triatomine bugs or mice and captive neotropical bats

Trypanosoma cruzi and Trypanosoma rangeli-like trypanosomes have been found in a variety of neotropical bat species. In this study, bats (Artibeus lituratus, Carollia perspicillata, Desmodus rotundus, Glossophaga soricina, Molossus molossus, Phyllostomus hastatus) were maintained under controlled conditions, and experiments were conducted to determine how they might become infected naturally with trypanosomes. All bats were first screened for existing infections by hemoculture and the examination of blood smears, and only apparently uninfected animals were then used in the experiments. Proof was obtained that the triatomine bug Rhodnius prolixus would readily feed upon some of the bats, and two species became infected after being bitten by bugs infected with T. rangeli. Some bats also became infected by ingesting R. prolixus carrying T. cruzi, or following subcutaneous or intragastic inoculation with fecal suspensions of R. prolixus containing T. cruzi. P. hastatus became infected after ingesting mice carrying T. cruzi. All of the bats studied inhabit roosts that may be occupied by triatomine bugs and, with the exception of D. rotundus, all also feed to at least some extent upon insects. These findings provide further evidence of how bats may play significant roles in the epidemiology of T. cruzi and T. rangeli in the New World tropics.     

The evolution of echolocation in bats

Recent molecular phylogenies have changed our perspective on the evolution of echolocation in bats. These phylogenies suggest that certain bats with sophisticated echolocation (e.g. horseshoe bats) share a common ancestry with non-echolocating bats (e.g. Old World fruit bats). One interpretation of these trees presumes that laryngeal echolocation (calls produced in the larynx) probably evolved in the ancestor of all extant bats. Echolocation might have subsequently been lost in Old World fruit bats, only to evolve secondarily (by tongue clicking) in this family. Remarkable acoustic features such as Doppler shift compensation, whispering echolocation and nasal emission of sound each show multiple convergent origins in bats. The extensive adaptive radiation in echolocation call design is shaped largely by ecology, showing how perceptual challenges imposed by the environment can often override phylogenetic constraints.     

The Evolution of Bat Vestibular Systems in the Face of Potential Antagonistic Selection Pressures for Flight and Echolocation

The vestibular system maintains the body’s sense of balance and, therefore, was probably subject to strong selection during evolutionary transitions in locomotion. Among mammals, bats possess unique traits that place unusual demands on their vestibular systems. First, bats are capable of powered flight, which in birds is associated with enlarged semicircular canals. Second, many bats have enlarged cochleae associated with echolocation, and both cochleae and semicircular canals share a space within the petrosal bone. To determine how bat vestibular systems have evolved in the face of these pressures, we used micro-CT scans to compare canal morphology across species with contrasting flight and echolocation capabilities. We found no increase in canal radius in bats associated with the acquisition of powered flight, but canal radius did correlate with body mass in bat species from the suborder Yangochiroptera, and also in non-echolocating Old World fruit bats from the suborder Yinpterochiroptera. No such trend was seen in members of the Yinpterochiroptera that use laryngeal echolocation, although canal radius was associated with wing-tip roundedness in this group. We also found that the vestibular system scaled with cochlea size, although the relationship differed in species that use constant frequency echolocation. Across all bats, the shape of the anterior and lateral canals was associated with large cochlea size and small body size respectively, suggesting differential spatial constraints on each canal depending on its orientation within the skull. Thus in many echolocating bats, it seems that the combination of small body size and enlarged cochlea together act as a principal force on the vestibular system. The two main groups of echolocating bats displayed different canal morphologies, in terms of size and shape in relation to body mass and cochlear size, thus suggesting independent evolutionary pathways and offering tentative support for multiple acquisitions of echolocation.     

Scale‐dependent effects of forest restoration on Neotropical fruit bats

Neotropical fruit bats (family Phyllostomidae) facilitate forest regeneration on degraded lands by dispersing shrub and tree seeds. Accordingly, if fruit bats can be attracted to restoration sites, seed dispersal could be enhanced. We surveyed bat communities at 10 sites in southern Costa Rica to evaluate whether restoration treatments attracted more fruit bats if trees were planted on degraded farmlands in plantations or island configurations versus natural regeneration. We also compared the relative influence of tree cover at local and landscape spatial scales on bat abundances. We captured 68% more fruit bat individuals in tree plantations as in controls, whereas tree island plots were intermediate. Bat activity also responded to landscape tree cover within a 200‐m radius of restoration plots, with greater abundance but lower species richness in deforested landscapes. Fruit bat captures in controls and tree island plots declined with increasing landscape tree cover, but captures in plantations were relatively constant. Individual species responded differentially to tree cover measured at different spatial scales. We attribute restoration effects primarily to habitat structure rather than food resources because no planted trees produced fruits regularly eaten by bats. The magnitude of tree planting effects on fruit bats was less than previous studies have found for frugivorous birds, suggesting that bats may play a particularly important role in dispersing seeds in heavily deforested and naturally regenerating areas. Nonetheless, our results show that larger tree plantations in more intact landscapes are more likely to attract diverse fruit bats, potentially enhancing seed dispersal.     

Chromosomal homologies among vampire bats revealed by chromosome painting (phyllostomidae, chiroptera)

Substantial effort has been made to elucidate karyotypic evolution of phyllostomid bats, mostly through comparisons of G-banding patterns. However, due to the limited number of G-bands in respective karyotypes and to the similarity of non-homologous bands, an accurate evolutionary history of chromosome segments remains questionable. This is the case for vampire bats (Desmodontinae). Despite several proposed homologies, banding data have not yet provided a detailed understanding of the chromosomal changes within vampire genera. We examined karyotype differentiation of the 3 species within this subfamily using whole chromosomal probes from Phyllostomus hastatus (Phyllostominae) and Carollia brevicauda (Carolliinae). Painting probes of P. hastatus respectively detected 22, 21 and 23 conserved segments in Diphylla ecaudata, Diaemus youngi, and Desmodus rotundus karyotypes, whereas 27, 27 and 28 were respectively detectedwith C. brevicauda paints. Based on the evolutionary relationships proposed by morphological and molecular data, we present probable chromosomal synapomorphies for vampire bats and propose chromosomes that were present in the common ancestor of the 5 genera analyzed. Karyotype comparisons allowed us to relate a number of conserved chromosomal segments among the 5 species, providing a broader database for understanding karyotype evolution in the family.     

Vocal learning by greater spear-nosed bats.

Vocal learning is well known among passerine and psittacine birds, but most data on mammals are equivocal. Specific benefits of vocal learning are poorly understood for most species. One case where vocal learning should be favoured by selection is where calls indicate group membership and group mates are unrelated. Female greater spear-nosed bats, Phyllostomus hastatus, live in stable groups of unrelated bats and use loud, broadband calls to coordinate foraging movements of social group mates. Bats benefit from group foraging. Calls differ between female social groups and cave colonies, and playback experiments demonstrate that bats perceive these acoustic differences. Here I show that the group distinctive structure of calls arises through vocal learning. Females change call structure when group composition changes, resulting in increased similarity among new social group mates. Comparisons of transfers with age-matched half-sibs indicate that call changes are not simply due to maturation, the physical environment or heredity. These results suggest that studies testing vocal learning in mammals could profit by focusing on vocalizations that signify group membership.     

The role of fruit traits of bird-dispersed plants in invasiveness and weed risk assessment

Aim  Birds play a major role in the dispersal of seeds of many fleshy-fruited invasive plants. The fruits that birds choose to consume are influenced by fruit traits. However, little is known of how the traits of invasive plant fruits contribute to invasiveness or to their use by frugivores. We aim to gain a greater understanding of these relationships to improve invasive plant management.
Location  South-east Queensland, Australia.
Methods  We measure a variety of fruit morphology, pulp nutrient and phenology traits of a suite of bird-dispersed alien plants. Frugivore richness of these aliens was derived from the literature. Using regressions and multivariate methods, we investigate relationships between fruit traits, frugivore richness and invasiveness.
Results  Plant invasiveness was negatively correlated to fruit size, and all highly invasive species had quite similar fruit morphology [smaller fruits, seeds of intermediate size and few (< 10) seeds per fruit]. Lower pulp water was the only pulp nutrient trait associated with invasiveness. There were strong positive relationships between the diversity of bird frugivores and plant invasiveness, and in the diversity of bird frugivores in the study region and another part of the plants' alien range.
Main conclusions  Our results suggest that weed risk assessments (WRA) and predictions of invasive success for bird-dispersed plants can be improved. Scoring criteria for WRA regarding fruit size would need to be system-specific, depending on the fruit-processing capabilities of local frugivores. Frugivore richness could be quantified in the plant's natural range, its invasive range elsewhere, or predictions made based on functionally similar fruits.     

食鱼蝙蝠形态和行为特化研究

总结了食鱼蝙蝠种类、分布 ,及其形态结构、回声定位功能和捕食行为的研究成果。比较食鱼蝙蝠与近水面“拖网式”食虫蝙蝠在形态、回声定位信号及捕食行为方面的异、同 ,推测食鱼蝙蝠起源于“拖网式”食虫蝙蝠类 ;体形和回声定位信号的几种特异性是捕食行为进化压力 ,而环境是决定因素。     

Neural coding of echo-envelope disparities in echolocating bats

The effective use of echolocation requires not only measuring the delay between the emitted call and returning echo to estimate the distance of an ensonified object. To locate an object in azimuth and elevation, the bat’s auditory system must analyze the returning echoes in terms of their binaural properties, i.e., the echoes’ interaural intensity and time differences (IIDs and ITDs). The effectiveness of IIDs for echolocation is undisputed, but when bats ensonify complex objects, the temporal structure of echoes may facilitate the analysis of the echo envelope in terms of envelope ITDs. Using extracellular recordings from the auditory midbrain of the bat, Phyllostomus discolor, we found a population of neurons that are sensitive to envelope ITDs of echoes of their sonar calls. Moreover, the envelope-ITD sensitivity improved with increasing temporal fluctuations in the echo envelopes, a sonar parameter related to the spatial statistics of complex natural reflectors like vegetation. The data show that in bats envelope ITDs may be used not only to locate external, prey-generated rustling sounds but also in the context of echolocation. Specifically, the temporal fluctuations in the echo envelope, which are created when the sonar emission is reflected from a complex natural target, support ITD-mediated echolocation.     

Fruit characteristics and factors affecting fruit removal in a Panamanian community of strangler figs

We describe fruiting characteristics for 12 species in a community of strangler figs (Moraceae: Urostigma) studied in Panama. We quantify diurnal and nocturnal removal rates and proportions of fruits removed, and relate them to the activities of the main dispersers of the figs: bats and birds. These results combined with previous studies show that there are clear differences between fig species with fruit that ripen red and those with fruit that remain green(ish). In the red-fruited species, the fruit are small, ripen asynchronously over relatively long periods, produce little scent, and are mainly taken during the day by birds. In contrast, in the green(ish)-fruited species, the fruits are larger, span a range of sizes, ripen relatively synchronously, produce very distinctive aromas, and are mainly taken at night by bats. This dichotomy in fruiting characteristics suggests coadaptive links between groups of dispersers and different species within the genus Ficus. All fig species produce a range of fruit crop sizes (10–155 fuits/m² canopy area) of which a high proportion were removed by seed dispersers (>80%). Removal rates (fruit removed per day) were positively correlated with crop size, suggesting that trees with large crop size attract more frugivores. Removal rates of green-fruited figs were significantly lower and persistence and abortion of ripe fruit were significant higher around full moon, apparently due to the reduced activity of bats. We further estimate the number of bats that are sustained by a tree fruit crop and account for the observed fruit removal. We then discuss the evidence for coadaptation between different groups of figs and their seed dispersers, Finally, we consider the conservation implications for figs as keystone resources in tropical forests. Received: 26 April 1999 / Accepted: 10 January 2000     

Discrimination of Infant Isolation Calls by Female Greater Spear-Nosed Bats, Phyllostomus hastatus

In colonial species, recognition of offspring should be under strong selection. For accurate identification to occur offspring must emit individually distinctive signals and parents must be able to discriminate between signals. Female greater spear-nosed bats (Phyllostomus hastatus) roost in stable social groups and use infant vocalizations, termed isolation calls, to locate and identify their young. In this study, we investigate both the production and perception of isolation calls in P. hastatus. First, we measured acoustic features and found that after controlling for ontogenetic effects, sufficient variation exists between pups for isolation calls to function as individual signatures. Moreover, pups from the same social group emit calls with more similar spectral and spectro-temporal features than pups from different social groups, indicating that these features are likely heritable. We used psychoacoustic experiments in the laboratory to determine if adult females could discriminate between calls from pups in the same or different social group. Females discriminated between pups when faced with a template-matching task and their performance was correlated with the salience of spectral and spectro-temporal features. We found no difference in performance when females had to discriminate between pups from the same and different social groups. These results indicate that females should be able to accurately identify their young using isolation calls.     

The Aerodynamic Cost of Head Morphology in Bats: Maybe Not as Bad as It Seems

At first sight, echolocating bats face a difficult trade-off. As flying animals, they would benefit from a streamlined geometric shape to reduce aerodynamic drag and increase flight efficiency. However, as echolocating animals, their pinnae generate the acoustic cues necessary for navigation and foraging. Moreover, species emitting sound through their nostrils often feature elaborate noseleaves that help in focussing the emitted echolocation pulses. Both pinnae and noseleaves reduce the streamlined character of a bat’s morphology. It is generally assumed that by compromising the streamlined charactered of the geometry, the head morphology generates substantial drag, thereby reducing flight efficiency. In contrast, it has also been suggested that the pinnae of bats generate lift forces counteracting the detrimental effect of the increased drag. However, very little data exist on the aerodynamic properties of bat pinnae and noseleaves. In this work, the aerodynamic forces generated by the heads of seven species of bats, including noseleaved bats, are measured by testing detailed 3D models in a wind tunnel. Models of Myotis daubentonii, Macrophyllum macrophyllum, Micronycteris microtis, Eptesicus fuscus, Rhinolophus formosae, Rhinolophus rouxi and Phyllostomus discolor are tested. The results confirm that non-streamlined facial morphologies yield considerable drag forces but also generate substantial lift. The net effect is a slight increase in the lift-to-drag ratio. Therefore, there is no evidence of high aerodynamic costs associated with the morphology of bat heads.     

Click-based echolocation in bats: not so primitive after all

Echolocating bats of the genus Rousettus produce click sonar signals, using their tongue (lingual echolocation). These signals are often considered rudimentary and are believed to enable only crude performance. However, the main argument supporting this belief, namely the click’s reported long duration, was recently shown to be an artifact. In fact, the sonar clicks of Rousettus bats are extremely short, ~50–100 μs, similar to dolphin vocalizations. Here, we present a comparison between the sonar systems of the ‘model species’ of laryngeal echolocation, the big brown bat (Eptesicus fuscus), and that of lingual echolocation, the Egyptian fruit bat (Rousettus aegyptiacus). We show experimentally that in tasks, such as accurate landing or detection of medium-sized objects, click-based echolocation enables performance similar to laryngeal echolocators. Further, we describe a sophisticated behavioral strategy for biosonar beam steering in clicking bats. Finally, theoretical analyses of the signal design—focusing on their autocorrelations and wideband ambiguity functions—predict that in some aspects, such as target ranging and Doppler-tolerance, click-based echolocation might outperform laryngeal echolocation. Therefore, we suggest that click-based echolocation in bats should be regarded as a viable echolocation strategy, which is in fact similar to the biosonar used by most echolocating animals, including whales and dolphins.     

Variation in leaf longevity of Pistacia lentiscus and its relationship to sex and drought stress inferred from leaf δ13C

1. Leaf formation, loss, retention, longevity and biomass on male branches of the evergreen mediterranean shrub Pistacia lentiscus , L. correlated strongly with water-use efficiency inferred from leaf δ¹³C across a gradient of precipitation on the island of Mallorca, Spain.
2. The correlations suggest that the leaf phenology is under control of drought-induced constraints on the carbon balance.
3. In fruiting female branches, the correlations between the inferred water-use efficiency and number of formed and retained leaves, leaf biomass and leaf longevity were non-significant. Leaf formation was strongly reduced by fruiting and the females compensated the reduced photosynthetic capacity by retaining older leaves for a longer time than male plants.
4. It is suggested that leaf longevity in females is under strong control of resource allocation to fruit formation which is 'overlaid' on the drought-induced carbon stress, which led to the observed longer leaf longevity in females than in males.     

The relative influence of competition and prey defences on the trophic structure of animalivorous bat ensembles

Deterministic filters such as competition and prey defences should have a strong influence on the community structure of animals like animalivorous bats which have life histories characterized by low fecundity, low predation risk, long life expectancy and stable populations. We investigated the relative influence of these two deterministic filters on the trophic structure of animalivorous bat assemblages in South Africa. We used null models to test if patterns of dietary overlap were significantly different from patterns expected by chance and multivariate analyses to test the correlations between diet and phenotype (body size, wing morphology and echolocation). We found little evidence that competition structured the trophic niche of coexisting bats. Contrary to predictions from competition, dietary overlap between bats of ensembles and functional groups (open-air, clutter-edge, and clutter foragers) were significantly higher than expected by chance. Instead, we found support for the predictions of the allotonic frequency hypothesis: there were significant relationships between peak echolocation frequency and the proportion of moths in the diets of bats at local and regional scales, and peak echolocation frequency was the best predictor of diet even after we controlled for the influence of body size and phylogeny. These results suggest that echolocation frequency and prey hearing exert more influence on the trophic structure of sympatric animalivorous bats than competition. Nonetheless, differential habitat use and sensory bias may also be major determinants of trophic structure because these are also correlated with frequencies of bat calls.     

Old World fruit bats can be long-distance seed dispersers through extended retention of viable seeds in the gut

Seed dispersal and pollination by animals play a crucial role in the maintenance of forest ecosystems worldwide. Frugivorous bats are important pollen and seed dispersers in both the Palaeo- and Neotropics, and at least 300 plant species are known to rely on Old World fruit bats (Megachiroptera, Pteropodidae) for their propagation. However, rapid food transit times (generally less than 30 minutes) in frugivorous bats have been thought to limit their ability to disperse seeds to just a few tens of kilometres. Here we demonstrate regular daytime (greater than 12 hours) retention of food and viable fig seeds (Ficus, Moraceae) in the gut of the Old World fruit bat Cynopterus sphinx: a behaviour not previously reported for any frugivorous bat. Field observations indicate that this behaviour also occurs in other genera. Old World fruit bats are highly mobile and many species undertake considerable foraging and migration flights. Our findings indicate that Old World fruit bats have the potential to disperse small seeds hundreds of kilometres. This necessitates a reappraisal of their importance in transporting zoochorous seeds to remote areas and facilitating gene flow between isolated populations of plants, both within mainlands and across ocean barriers.     

Auditory sensitivity and frequency selectivity in greater spear-nosed bats suggest specializations for acoustic communication

We investigated the relationship between auditory sensitivity, frequency selectivity, and the vocal repertoire of greater spear-nosed bats (Phyllostomus hastatus). P. hastatus commonly emit three types of vocalizations: group-specific foraging calls that range from 6 to 11 kHz, low amplitude echolocation calls that sweep from 80 to 40 kHz, and infant isolation calls from 15 to 100 kHz. To determine if hearing in P. hastatus is differentially sensitive or selective to frequencies in these calls, we determined absolute thresholds and masked thresholds using an operant conditioning procedure. Both absolute and masked thresholds were lowest at 15 kHz, which corresponds with the peak energy of isolation calls. Auditory and masked thresholds were higher at sound frequencies used for group-specific foraging calls and echolocation calls. Isolation calls meet the requirements of individual signatures and facilitate parent-offspring recognition. Many bat species produce isolation calls with peak energy between 10 and 25 kHz, which corresponds with the frequency region of highest sensitivity in those species for which audiogram data are available. These findings suggest that selection for accurate offspring recognition exerts a strong influence on the sensory system of P. hastatus and likely on other species of group-living bats.