The unusual inheritance pattern of the courtship songs in closely related grasshopper species of the Chorthippus albomarginatus-group (Orthoptera: Gomphocerinae).

Sibling species of the Chorthippus albomarginatus-group are exceptional among all European grasshopper species because they have an extremely elaborate courtship behaviour. Here, we present a genetic analysis of the courtship song differences between two closely related grasshopper species Ch. albomarginatus and Ch. oschei. Measurements of seven courtship song characters and one parameter of courtship visual display were compared among parentals, F(1) and F(2) hybrids, and backcrosses. Variation in one morphological character, the number of stridulatory pegs, was studied as well. The distributions of song phenotypes were more consistent with a type-III genetic architecture, which involves complementary or duplicate pairs of loci. We suggest that this type of architecture may provide the novel elements of the hybrid songs, even more complex than the parental ones, which may offer a new starting point for sexual selection. One parameter of the visual display appeared to be controlled by a single locus, but differences in other characters of visual display could be explained by a more complex inheritance pattern. Inheritance of the peg number differences was consistent with a simple polygenic additive model. Sex linkage and/or maternal effects were not detected for any trait. We found a disproportionate influence of the Ch. albomarginatus parent on most hybrid song characteristics and the visual display, and this may support a previously suggested idea that Ch. albomarginatus is an ancestral form and Ch. oschei is a derived species.

Echolocation range and wingbeat period match in aerial-hawking bats.

Aerial-hawking bats searching the sky for prey face the problem that flight and echolocation exert independent and possibly conflicting influences on call intervals. These bats can only exploit their full echolocation range unambiguously if they emit their next call when all echoes from the preceding call would have arrived. However, not every call interval is equally available. The need to reduce the high energetic costs of echolocation forces aerial-hawking bats to couple call emission to their wingbeat. We compared the wingbeat periods of 11 aerial-hawking bat species with the delays of the last-expected echoes. Acoustic flight-path tracking was employed to measure the source levels (SLs) of echolocation calls in the field. SLs were very high, extending the known range to 133 dB peak equivalent sound pressure level. We calculated the maximum detection distances for insects, larger flying objects and background targets. Wingbeat periods were derived from call intervals. Small and medium-sized bats in fact matched their maximum detection range for insects and larger flying targets to their wingbeat period. The tendency to skip calls correlated with the species' detection range for background targets. We argue that a species' call frequency is at such a pitch that the resulting detection range matches their wingbeat period.

Object recognition by echolocation: a nectar-feeding bat exploiting the flowers of a rain forest vine.

In the bat-pollinated vine Mucuna holtonii only the first visit to a flower is rewarded with a substantial amount of nectar, which is released when a bat lands on the flower and triggers an explosion mechanism. During later visits the bats receive only small amounts of nectar. Nevertheless, the inflorescences as a whole remain attractive, since further buds successively open during the night. Nectar-feeding bats Glossophaga commissarisi selectively visit unexploded, "virgin" flowers. They can discriminate buds, virgin and exploded flowers using echolocation. In field experiments bats exploited virgin flowers, the vexillum of which had been replaced by a same-sized triple mirror or by an artificial vexillum. Such flowers were frequently inspected, but not as often exploited as natural flowers. In two-alternative-forced-choice experiments the bats learned to discriminate between replicas of the vexillum and triple mirrors. The recognition distance was between 15 and 50 cm. Echoes of the three flowering stages differ in their spectral composition, which changes in dependence of the sound incidence angle in a characteristic way. We conclude that glossophagine bats are able to recognize small motionless structures like flowers and to accurately adjust their landing manoeuvres by using their echolocation system alone.

Conflicting molecular phylogenies of European long-eared bats (Plecotus) can be explained by cryptic diversity.

Conflicting phylogenetic signals of two data sets that analyse different portions of the same molecule are unexpected and require an explanation. In the present paper we test whether (i) differential evolution of two mitochondrial genes or (ii) cryptic diversity can better explain conflicting results of two recently published molecular phylogenies on the same set of species of long-eared bats (genus Plecotus). We sequenced 1714bp of three mitochondrial regions (16S, ND1, and D-loop) of 35 Plecotus populations from 10 European countries. A likelihood ratio test revealed congruent phylogenetic signals of the three data partitions. Our phylogenetic analyses demonstrated that the existence of a previously undetected Plecotus lineage caused the incongruities of previous studies. This lineage is differentiated on the species level and lives in sympatry with its sister lineage, Plecotus auritus, in Switzerland and Northern Italy. A molecular clock indicates that all European Plecotus species are of mid or late Pliocene origin. Plecotus indet. was previously described as an intergrade between P. auritus and Plecotus austriacus since it shares morphological characters with both. It is currently known from elevations above 800 m a.s.l. in the Alps, the Dinarian Alps and the Pindos mountains in Greece. Since we could demonstrate that incongruities of two molecular analyses simply arose from the mis-identification of one lineage, we conclude that molecular phylogenetic analyses do not free systematists from a thorough inclusion of morphological and ecological data.

Cryptic diversity in European bats.

Different species of bat can be morphologically very similar. In order to estimate the amount of cryptic diversity among European bats we screened the intra- and interspecific genetic variation in 26 European vespertilionid bat species. We sequenced the DNA of subunit 1 of the mitochondrial protein NADH dehydrogenase (ND1) from several individuals of a species, which were sampled in a variety of geographical regions. A phylogeny based on the mitochondrial (mt) DNA data is in good agreement with the current classification in the family. Highly divergent mitochondrial lineages were found in two taxa, which differed in at least 11% of their ND1 sequence. The two mtDNA lineages in Plecotus austriacus correlated with the two subspecies Plecotus austriacus austriacus and Plecotus austriacus kolombatovici. The two mtDNA lineages in Myotis mystacinus were partitioned among two morphotypes. The evidence for two new bat species within Europe is discussed. Convergent adaptive evolution might have contributed to the morphological similarity among distantly related species if they occupy similar ecological niches. Closely related species may differ in their ecology but not necessarily in their morphology. On the other hand, two morphologically clearly different species (Eptesicus serotinus and Eptesicus nilssonii) were found to be genetically very similar. Neither morphological nor mitochondrial DNA sequence analysis alone can be guaranteed to identify species.

Cryptic mammalian species: a new species of whiskered bat (Myotis alcathoe n. sp.) in Europe.

The analysis of morphological, behavioural and genetic characters of whiskered bats revealed a new European bat species within the family Vespertilionidae. We describe the morphology, karyology, genetic similarity, ecology and distribution of Myotis alcathoe n. sp. It closely resembles Myotis mystacinus, Myotis brandtii and Myotis ikonnikovi in morphology, but all four species show clear genetic differences in two mitochondrial genes (ND1 and 12S rRNA). Myotis alcathoe n. sp. is the smallest species among the European whiskered bats and uses the highest-frequency echolocation calls of all the European Myotis species. It prefers to hunt in small valleys with deciduous trees and flowing water, which is an endangered habitat. Records from Greece and Hungary indicate a distribution range in south-eastern Europe.

Song pattern recognition in the grasshopper Chorthippus biguttulus: the mechanism of syllable onset and offset detection.

The male song of the duetting grasshopper Chorthippus biguttulus consists of syllables alternating with noisy pauses. The syllable-pause structure is important for song recognition by the female. Using playback experiments we investigated the mechanism by which intensity modulations within the song pattern are used to detect syllable onsets and offsets. We varied the relative onset level (level of the syllable beginning relative to the noisy pause) and the relative offset level (level of the noisy pause relative to the syllable end) independently in different experiments. For all females, an increase in intensity defining the syllable onset was necessary to evoke responses. Syllable offset cues were not always necessary: some females responded to continuous noise stimuli wherein only syllable onsets were marked by short pulses of high intensity. Those females that did not require syllable offset cues did not, however, lack a functional pause detection mechanism, since their responses to model songs containing silent pauses were restricted to a given range of pause durations. We propose that syllable-pause detection involves two independent processes: (1) syllable onset detection by a phasic neuronal unit that can be re-activated only after a short pause, and (2) the rejection of unacceptably long pauses by a second unit.

Sulphur-containing "perfumes" attract flower-visiting bats.

We tested the attractiveness of individual scent compounds of bat-pollinated flowers to their pollinators, small flower-visiting bats of the genus Glossophaga (Phyllostomidae). Twenty compounds belonging to four different chemical substance classes were tested, both in the laboratory and in the field. In the laboratory, the bats (Glossophaga soricina) approached odour sources spontaneously and without preceding experience. Without ever receiving any reward they preferred the scent of a sulphur-containing compound, dimethyl disulphide, to several other odour components emitted by bat-pollinated flowers, and to scentless controls. In the field, at La Selva station in the tropical lowland rain forest of Costa Rica, G. commissarisi were attracted by two sulphur-containing compounds, dimethyl disulphide and 2,4-dithiapentane, to visit artificial flowers filled with sugar water. Thus, in nectarivorous bats the sense of smell obviously plays an important role in searching for and localising food sources, and even single components of the scent bouquets of bat-pollinated flowers are attractive. The preference for sulphur-containing odours seems to be innate.

Listening for bats: the hearing range of the bushcricket Phaneroptera falcata for bat echolocation calls measured in the field.

The hearing range of the tettigoniid Phaneropterafalcata for the echolocation calls of freely flying mouseeared bats (Myotis myotis) was determined in the field. The hearing of the insect was monitored using hook electrode recordings from an auditory interneuron, which is as sensitive as the hearing organ for frequencies above 16 kHz. The flight path of the bat relative to the insect's position was tracked by recording the echolocation calls with two microphone arrays, and calculating the bat's position from the arrival time differences of the calls at each microphone. The hearing distances ranged from 13 to 30 m. The large variability appeared both between different insects and between different bat approaches to an individual insect. The escape time of the bushcricket, calculated from the detection distance of the insect and the instantaneous flight speed of the bat, ranged from 1.5 to more than 4s. The hearing ranges of bushcrickets suggest that the insect hears the approaching bat long before the bat can detect an echo from the flying insect.

The energy cost of flight: do small bats fly more cheaply than birds?

Flapping flight is one of the most expensive activities in terms of metabolic cost and this cost has previously been considered equal for the two extant vertebrate groups which evolved flapping flight. Owing to the difficulty of obtaining accurate measurements without disturbing flight performance, current estimates of flight cost within the group of small birds and bats differ by more than a factor of five for given body masses. To minimize the potential problem that flight behaviour may be affected by the measurements, we developed an indirect method of measuring flight energy expenditure based on time budget analysis in which small nectar-feeding bats (Glossophaginae) could continue their natural rhythm of flying and resting entirely undisturbed. Estimates of metabolic flight power based on 172 24-h time and energy budget measurements were obtained for nine individual bats from six species (mass 7-28 g). Metabolic flight power (PF) of small bats was found to increase with body mass following the relation PF = 50.2 M0.771 (r2 = 0.96, n = 13, PF in W, M in kg). This is about 20-25% below the majority of current predictions of metabolic flight cost for small birds. Thus, either the flight cost of small birds is significantly lower than has previously been thought or, contrary to current opinion, small bats require less energy to fly than birds.

Gas exchange during hovering flight in a nectar-feeding bat Glossophaga soricina.

Glossophagine nectar-feeding bats exploit flowers while hovering in front of them. Aerodynamic theory predicts that power output for hovering flight in Glossophaga soricina is 2.6 times higher than during horizontal flight. We tested this prediction by measuring rates of gas exchange during hover-feeding. Five individuals of Glossophaga soricina (mean mass 11.7 g) were trained to feed from a nectar dispenser designed as a flow-through respirometry mask. Single hover-feeding events lasted for up to 4.5 s. Measured rates of gas exchange varied as a function of hovering duration. O2 and CO2 during short hovering events (up to 1 s) were 20.5+/-6.7 ml g-1 h-1 (N=55) and 21.6+/-5.6 ml g-1 h-1 (N=39) (means +/- S.D.), respectively. These values are in the range of a previous estimate of the metabolic power input for level forward flight (23.8 ml O2 g-1 h-1). However, during hovering events lasting longer than 3 s, oxygen uptake was only 16.7+/-3.5 ml g-1 h-1 (N=73), which is only 70 % of the value expected for forward flight. Thus, bats reduced their rate of oxygen uptake during longer periods of hovering compared with level forward flight. This result is in contrast to the predicted hovering cost derived from aerodynamic theory. The exact metabolic power input during hovering remains uncertain. During longer hovering events, bats were probably not in respiratory steady state, as indicated by the deviation of the respiratory exchange ratio from the expected value of 1 (oxidization of nectar sugar) to the measured value of 0.8.

Headspace analysis of volatile flower scent constituents of bat-pollinated plants.

The flower scents of 11 bat-pollinated plant species have been investigated using the 'closed-loop-stripping'-adsorption technique. GC and GC/MS analyses of the headspace samples resulted in the identification of 49 compounds, comprising amongst others sulphur compounds, terpenoids and aliphatic compounds.

The cost of hovering and forward flight in a nectar-feeding bat, Glossophaga soricina, estimated from aerodynamic theory.

Energy expenditure during flight in animals can best be understood and quantified when both theoretical and empirical approaches are used concurrently. This paper examines one of four methods that we have used to estimate the cost of flight in a neotropical nectar-feeding bat Glossophaga soricina (Phyllostomidae), namely the use of kinematic and morphological data and aerodynamic theory to estimate the mechanical power requirements (power output) for hovering and horizontal forward flight. A hot-wire anemometer was used to measure induced velocity (the velocity of air accelerated by the wings) during hovering in order to estimate induced power. Our estimate of aerodynamic power (the sum of induced, profile and parasite powers) required for a 0.0105 kg G. soricina to hover is 0.15 W and our estimate of the inertial power (the power required to oscillate the wings) is 0.19 W. Thus, the total mechanical power for hovering is 0.34 W or 32.4 W kg-1. The mechanical power required for horizontal forward flight, near the minimum power flight speed (4.2 ms-1) for a 0.0117 kg bat is 0.14 W (12.3 W kg-1), of which 0.10 W is aerodynamic power and 0.042 W is inertial power. Comparison with our results on metabolic power requirements estimated from nectar intake gives a mechanical efficiency of 0.15 for hovering flight and 0.11 for forward flight near the minimum power speed.

'Switching-off' of an auditory interneuron during stridulation in the acridid grasshopper Chorthippus biguttulus L.

In freely moving grasshoppers of the species Chorthippus biguttulus compound potentials were recorded from the neck connectives with chronically implanted hook electrodes. The spikes of one large auditory interneuron, known as the G-neuron (Kalmring 1975a, b) were clearly distinguishable in the recordings and the neuron was identified by its physiology and morphology. In quiescent grasshoppers the G-neuron responds to auditory and vibratory stimuli, but responses to both stimuli are suppressed during stridulation in males. When a male's wings were removed so that the stridulatory movements of its hindlegs produced no sound, the suppression of the G-neuron response still occurred. When proprioceptive feedback from the hindlegs was reduced, by forced autotomy of the legs, the switching-off remained incomplete (production of stridulatory patterns was inferred on the basis of electromyograms from the relevant thoracic musculature). Imposed movement of the hindlegs, on the other hand, suppressed the G-neuron response in a graded fashion, depending on the frequency of the movement. These experiments suggest that the switching-off is brought about by a combination of proprioceptive feedback and central efferences. The switching-off phenomenon could either protect the grasshopper's auditory pathway from undesired effects of overloading by its own intense song (e.g. self-induced habituation as described by Krasne and Wine 1977) and should therefore apply for most auditory neurons. Alternatively it could prevent escape reflexes from being triggered by stridulatory self-stimulation and consequently might apply only for neurons involved in such networks (as the G-neuron might be).