Defining the Impact of Non‐Native Species

Non‐native species cause changes in the ecosystems to which they are introduced. These changes, or some of them, are usually termed impacts; they can be manifold and potentially damaging to ecosystems and biodiversity. However, the impacts of most non‐native species are poorly understood, and a synthesis of available information is being hindered because authors often do not clearly define impact. We argue that explicitly defining the impact of non‐native species will promote progress toward a better understanding of the implications of changes to biodiversity and ecosystems caused by non‐native species; help disentangle which aspects of scientific debates about non‐native species are due to disparate definitions and which represent true scientific discord; and improve communication between scientists from different research disciplines and between scientists, managers, and policy makers. For these reasons and based on examples from the literature, we devised seven key questions that fall into 4 categories: directionality, classification and measurement, ecological or socio‐economic changes, and scale. These questions should help in formulating clear and practical definitions of impact to suit specific scientific, stakeholder, or legislative contexts. Definiendo el Impacto de las Especies No‐Nativas     

Fishing and echolocation behavior of the greater bulldog bat,Noctilio leporinus,in the field

When hunting for fish Noctilio leporinus uses several strategies. In high search flight it flies within 20–50 cm of the water surface and emits groups of two to four echolocation signals, always containing at least one pure constant frequency (CF) pulse and one mixed CF-FM pulse consisting of a CF component which is followed by a frequency-modulated (FM) component. The pure CF signals are the longest, with an average duration of 13.3 ms and a maximum of 17 ms. The CF component of the CF-FM signals averages 8.9 ms, the FM sweeps 3.9 ms. The CF components have frequencies of 52.8–56.2 kHz and the FM components have an average bandwidth of 25.9 kHz. A bat in high search flight reacts to jumping fish with pointed dips at the spot where a fish has broken the surface. As it descends to the water surface the bat shows the typical approach pattern of all bats with decreasing pulse duration and pulse interval. A jumping fish reveals itself by a typical pattern of temporary echo glints, reflected back to the bat from its body and from the water disturbance. In low search flight N. leporinus drops to a height of only 4–10 cm, with body parallel to the water, legs extended straight back and turned slightly downward, and feet cocked somewhat above the line of the legs and poised within 2–4 cm of the water surface. In this situation N. leporinus emits long series of short CF-FM pulses with an average duration of 5.6 ms (CF 3.1 and FM 2.6) and an average pulse interval of 20 ms, indicating that it is looking for targets within a short range. N. leporinus also makes pointed dips during low search flight by rapidly snapping the feet into the water at the spot where it has localized a jumping fish or disturbance. In the random rake mode, N. leporinus drops to the water surface, lowers its feet and drags its claws through the water in relatively straight lines for up to 10m. The echolocation behavior is similar to that of high search flight. This indicates that in this hunting mode N. leporinus is not pursuing specific targets, and that raking is a random or statistical search for surface fishes. When raking, the bat uses two strategies. In directed random rake it rakes through patches of water where fish jumping activity is high. Our interpretation is that the bat detects this activity by echolocation but prefers not to concentrate on a single jumping fish. In the absence of jumping fish, after flying for several minutes without any dips, N. leporinus starts to make very long rakes in areas where it has hunted successfully before (memory-directed random rake). Hunting bats caught a fish approximately once in every 50–200 passes through the hunting area.     

Ecological niche and phylogeny: the highly complex echolocation behavior of the trawling long-legged bat, Macrophyllum macrophyllum

Bats produce echolocation signals that reflect the sensory tasks they perform. In open air or over water, bats encounter few or no background echoes (clutter). Echolocation of such bats is the primary cue for prey perception and varies with the stage of approach to prey, typically comprising search, approach, and terminal group calls. In contrast, bats that glean stationary food from rough surfaces emit more uniform calls without a distinct terminal group. They use echolocation primarily for orientation in space and mostly need additional sensory cues for finding food because clutter echoes overlap strongly with food echoes. Macrophyllum macrophyllum is the only Neotropical leaf-nosed bat (Phyllostomidae) that hunts in clutter-poor habitat over water. As such, we hypothesized that, unlike all other members of its family, but similar to other trawling and aerial insectivorous bats, M. macrophyllum can hunt successfully by using only echolocation for prey perception. In controlled behavioral experiments on Barro Colorado Island, Panamá, we confirmed that echolocation alone is sufficient for finding prey in M. macrophyllum. Furthermore, we showed that pattern and structure of echolocation signals in M. macrophyllum are more similar to aerial and other trawling insectivorous bats than to close phylogenetic relatives. Particularly unique among phyllostomid bats, we found distinct search, approach, and terminal group calls in foraging M. macrophyllum. Call structure, however, consisting of short, multiharmonic, and steep frequency-modulated signals, closely resembled those of other phyllostomid bats. Thus, echolocation behavior in M. macrophyllum is shaped by ecological niche as well as by phylogeny.     

The echolocation and hunting behavior of Daubenton's bat,Myotis daubentoni

Summary The echolocation and hunting behavior of Daubenton's bat (Myotis daubentoni) were studied in the field under completely natural conditions using a multiflash photographic system synchronized with high-speed tape recordings. The hunting behavior of M. daubentoni is separated into four stages. In the search flight stage Daubenton's bat flies with an average speed of 3.4±0.6 m/s SD usually within 30 cm over water surfaces searching for insects. After the detection of potential prey, the approach flight stage occurs, during which the bat approaches the target in a goal-directed flight. The stage tail down indicates that M. daubentoni is close to the potential prey (approximately 10–22 cm) and is preparing for the catch. The insects are caught with the interfemoral membrane, the feet, and sometimes with the additional aid of a wing. In the stage head down, the bat seizes the prey during flight. Immediately afterwards, Daubenton's bat returns to search flight. M. daubentoni shows the typical echolocation behavior of a vespertilionid bat, emitting frequency-modulated (FM) echolocation signals. The three behavioral stages search, approach, and terminal phase (Griffin et al. 1960) are used to describe the pulse pattern of foraging M. daubentoni in the field. The terminal phase (or buzz) of Daubenton's bat is separated into two parts: buzz I and buzz II. Buzz II is distinguished from buzz I by the following characteristics: a sharp drop in terminal frequency, a distinct reduction in the bandwidth of the first harmonic, a continuous high repetition rate throughout the phase in the range 155–210 Hz, very short pulses (0,25–0.3 ms) and interpulse intervals (4.5–5.0 ms) at the end of the phase, and a distinct decrease in duty cycle. A pause in echolocation separates the end of the terminal phase from the ongoing search phase. The reduction in sound duration after the detection of a target and during pursuits with successfull or attempted catches is discussed in relation to the actual distance of the bat to the target at each stage. It is likely that Daubenton's bat reduces sound duration during approach and terminal phase in order to prevent an overlap of an outgoing pulse with the returning echo from the target. It is argued that the minimum detection distance can be estimated from the sound duration during search flight. Estimates of detection and reaction distances of M. daubentoni based upon synchronized photos and echolocation sequences are given to corroborate this hypothesis. An average detection distance of 128 cm and an average reaction distance of 112 cm were determined. Each behavioral stage of foraging M. daubentoni is characterized by a distinct pattern of echolocation signals and a distinct stage in hunting behavior. The approach flight in hunting behavior coincides with the approach phase and with buzz I in echolocation behavior. The stage tail down corresponds to buzz II. The stage head down is correlated with a pause in echolocation. Immediately afterwards, the bat returns into search flight and into the search phase, emitting search signals.     

Fruit detection and discrimination by small fruit-eating bats (Phyllostomidae): echolocation call design and olfaction

We studied the role of echolocation and other sensory cues in two small frugivorous New World leaf-nosed bats (Phyllostomidae: Artibeus watsoni and Vampyressa pusilla) feeding on different types of fig fruit. To test which cues the bats need to find these fruit, we conducted behavioral experiments in a flight cage with ripe and similar-sized figs where we selectively excluded vision, olfaction, and echolocation cues from the bats. In another series of experiments, we tested the discrimination abilities of the bats and presented sets of fruits that differed in ripeness (ripe, unripe), size (small, large), and quality (intact(infested with caterpillars). We monitored the bats' foraging and echolocation behavior simultaneously. In flight, both bat species continuously emitted short (<2 ms), multi-harmonic, and steep frequency-modulated (FM) calls of high frequencies, large bandwidth, and very low amplitude. Foraging behavior of bats was composed of two distinct stages: search or orienting flight followed by approach behavior consisting of exploration flights, multiple approaches of a selected fruit, and final acquisition of ripe figs in flight or in a brief landing. Both bat species continuously emitted echolocation calls. Structure and pattern of signals changed predictably when the bats switched from search or orienting calls to approach calls. We did not record a terminal phase before final acquisition of a fruit, as it is typical for aerial insectivorous bats prior to capture. Both bat species selected ripe over unripe fruit and non-infested over infested fruit. Artibeus watsoni preferred larger over smaller fruit. We conclude from our experiments, that the bats used a combination of odor-guided detection together with echolocation for localization in order to find ripe fruit and to discriminate among them.     

Protected‐Area Boundaries as Filters of Plant Invasions

Abstract: Human land uses surrounding protected areas provide propagules for colonization of these areas by non‐native species, and corridors between protected‐area networks and drainage systems of rivers provide pathways for long‐distance dispersal of non‐native species. Nevertheless, the influence of protected‐area boundaries on colonization of protected areas by invasive non‐native species is unknown. We drew on a spatially explicit data set of more than 27,000 non‐native plant presence records for South Africa's Kruger National Park to examine the role of boundaries in preventing colonization of protected areas by non‐native species. The number of records of non‐native invasive plants declined rapidly beyond 1500 m inside the park; thus, we believe that the park boundary limited the spread of non‐native plants. The number of non‐native invasive plants inside the park was a function of the amount of water runoff, density of major roads, and the presence of natural vegetation outside the park. Of the types of human‐induced disturbance, only the density of major roads outside the protected area significantly increased the number of non‐native plant records. Our findings suggest that the probability of incursion of invasive plants into protected areas can be quantified reliably.     

Dynamic adjustment of biosonar intensity to habitat clutter in the bat Macrophyllum macrophyllum (Phyllostomidae)

Echolocating bats adjust the time–frequency structure such as sweep rate and pulse interval of their sonar calls when they move from open space to vegetation-dense environments. Emitted call intensity is equally important for echolocation, but adjustment of signal intensity to different habitats has never been systematically studied in any bat species. To address this question, we recorded sonar calls of the Neotropical trawling insectivorous bat Macrophyllum macrophyllum (Phyllostomidae) at three sites with different obstacle densities (clutter). We found a clear correlation between emitted intensity and degree of clutter, with intensity proportional to decreasing clutter. In highly cluttered, semicluttered, and open spaces, M. macrophyllum emitted calls with mean source levels (sound pressure level (SPL) 10 cm from the bat’s mouth) of 100, 105, and 111 dB SPL root mean square (rms), respectively. To our knowledge, this is the first documentation of dynamic intensity adjustments in bats. Phyllostomid bats were previously considered silent, but the 111-dB SPL rms emitted by free-ranging M. macrophyllum in open space is comparable to output in aerial insectivorous bats from other families. Our results suggest that the acoustic constraints of habitats are better predictors of call intensity than phylogeny and therefore likely to be major drivers shaping the sonar system of bats in the course of evolution.     

Plasticity in echolocation signals of European pipistrelle bats in search flight: implications for habitat use and prey detection

We studied the echolocation and hunting behavior of three aerial insectivorous species of bats (Vespertilionidae: Pipistrellus) in the field in order to characterize the signals used by the bats and to determine how call structure varies in relation to habitat structure (uncluttered versus cluttered space). We documented free-flying, naturally foraging wild pipistrelles in various habitats using multiflash stereophotography combined with simultaneous sound recordings. Then we reconstructed the bat's flight position in three-dimensional space and correlated it with the corresponding echolocation sequences. In all three species of pipistrelles, signal structure varied substantially. In echolocation sequences of the search phase we found a consistent association of signal types with habitat types. In uncluttered habitats (obstacles more than 5 m from the bat) pipistrelles emitted almost exclusively narrowband signals with bandwidths less than 15 kHz. In cluttered habitats (obstacles less than 5 m from the bat) they switched to signals with bandwidths of more than 15 kHz. Wideband signals were also used when the bats were turning in cluttered and uncluttered spaces and for an instant after turning away from obstacles. Prey detection occured only when the outgoing signal did not overlap with the returning echo from potential prey. The bats also avoided overlap of echoes from potential prey and obstacles. Based on the results of this study, we propose an overlap-free window within which pipistrelles may detect potential prey and which allows predictions of minimum distances to prey and clutter-producing objects. Correspondence to: E.K.V. Kalko     

Echolocation and foraging behavior of the lesser bulldog bat, Noctilio albiventris : preadaptations for piscivory?

We studied variability in foraging behavior of Noctilio albiventris (Chiroptera: Noctilionidae) in Costa Rica and Panamá and related it to properties of its echolocation behavior. N. albiventris searches for prey in high (>20 cm) or low (<20 cm) search flight, mostly over water. It captures insects in mid-air (aerial captures) and from the water surface (pointed dip). We once observed an individual dragging its feet through the water (directed random rake). In search flight, N. albiventris emits groups of echolocation signals (duration 10–11 ms) containing mixed signals with constant-frequency (CF) and frequency-modulated (FM) components, or pure CF signals. Sometimes, mostly over land, it produces long FM signals (duration 15–21 ms). When N. albiventris approaches prey in a pointed dip or in aerial captures, pulse duration and pulse interval are reduced, the CF component is eliminated, and a terminal phase with short FM signals (duration 2 ms) at high repetition rates (150–170 Hz) is emitted. Except for the last pulses in the terminal phase N. albiventris avoids overlap between emitted signals and echoes returning from prey. During rakes, echolocation behavior is similar to that in high search flight. We compare N. albiventris with its larger congener, N. leporinus, and discuss behavioral and morphological specializations that can be interpreted as preadaptations favoring the evolution of piscivory as seen in N. leporinus. Prominent among these specializations are the CF components of the echolocation signals which allow detection and evaluation of fluttering prey amidst clutter-echoes, high variability in foraging strategy and the associated echolocation behavior, as well as morphological specializations such as enlarged feet for capturing prey from the water surface. Received: 21 April 1997 / Accepted after revision: 12 January 1998     

Echolocation behavior and signal plasticity in the Neotropical bat Myotis nigricans (Schinz, 1821) (Vespertilionidae): a convergent case with European species of Pipistrellus?

We used both field and flight cage observations to investigate the echolocation and foraging behavior of the seldom studied, small, aerial insectivorous bat Myotis nigricans (Vespertilionidae) in Panama. In contrast to its temperate congeners, M. nigricans foraged extensively in open space and showed an echolocation behavior well adapted to this foraging habitat. It broadcast narrowband echolocation signals of 7 ms duration that enhance the chance of prey detection in open space. Because of rhythmical alternations of signal amplitude from signal to signal in our sound recordings of search signals in open space, we conclude that the bats scanned their environment with head movements, thereby enlarging their search volume. In edge-and-gap situations, and in the flight cage, M. nigricans introduced an initial broadband component to its search calls. In the field and in the flight cage, M. nigricans hawked for prey in aerial catches; gleaning was never observed. M. nigricans demonstrates call structures, such as narrow bandwidth and rather long signals adapted to foraging predominantly in open space. Moreover, call structure is highly plastic, allowing M. nigricans to forage in edge-and-gap situations also. These adaptations in call structure and plasticity have evolved convergently at least twice within the genus Myotis. Finally, M. nigricans echolocation and foraging behavior parallels that of the small, aerial, insectivorous pipistrelle bats (Vespertilionidae), which are not closely related to M. nigricans but forage in similar habitats.