Crossbow‐netting: a new method for capturing shorebirds

Capturing shorebirds during the non‐breeding season can be challenging because they are usually scattered over wide‐open intertidal areas while foraging and are sensitive to human disturbance at roosts where they gather during high tide in large vigilant flocks. Several techniques are available for capturing shorebirds, but, for a study of stopover ecology, we needed a method that would allow us to capture Dunlins (Calidris alpina) on a regular basis at high‐tide roosts during the day (ruling out mist‐nets), did not require the use of gun‐powder (ruling out cannon‐nets), and that would deploy a net faster than clap nets, whoosh nets, and wilsternets. Therefore, we developed a new method to capture shorebirds where a crossbow is used to pull a mist‐net over flocks of roosting birds. We tested this technique in four habitats (saltpans, salt marshes, beaches, and mudflats) in the Tagus estuary, Portugal, and captured over 380 birds representing eight different species. Advantages of this technique compared to other methods (e.g., mist‐nets, clap‐ and whoosh nets, and cannon‐nets) include (1) portability, (2) ease of set up, (3) minimal disturbance of birds near the capture area, and (4) no explosive materials are needed. Our results suggest that crossbow‐netting is a safe and useful capture technique, especially for studies requiring the capture of small numbers of birds on a regular basis.     

Assessing the short‐term effects of capture,handling and tagging of sandgrouse

Capturing and marking free‐living birds permits the study of important aspects of their biology but may have undesirable effects. Bird welfare should be a primary concern, so it is necessary to evaluate and minimize any adverse effects of procedures used. We assess short‐term effects associated with the capture, handling and tagging with backpack‐mounted transmitters of Pin‐tailed Pterocles alchata and Black‐bellied Pterocles orientalis Sandgrouse, steppe birds of conservation concern. There was a significantly higher mortality (15%) during the first week after capture than during the following weeks (< 2.5%) in Pin‐tailed Sandgrouse, but no significant temporal mortality pattern in Black‐bellied Sandgrouse. In Pin‐tailed Sandgrouse, mortality rate during the first week increased with increasing relative transmitter and harness weight regardless of season, and with increasing handling time during the breeding season. There were no significant differences in mortality rate between study areas, type of tag, sex or age or an effect of restraint time. These results suggest the use of lighter transmitters (< 3% of the bird's weight) and a reduction of handling time (< 20 min), particularly during the breeding season, as essential improvements in procedure to reduce the mortality risk associated with the capture, handling and tagging of these vulnerable species.     

Capture myopathy in little bustards after trapping and marking

Four little bustards (Tetrax tetrax) (one adult and three juvenile males), captured with leg nooses and fitted with a backpack radiotag, died after capture. The first bird was found after 16 days with its left foot caught in the harness and died after 1 day. The other birds showed symptoms of capture myopathy after release, such as the difficulty or inability to fly and/or walk. They died after 5, 6, and 8 days, respectively. At necropsy, muscles affected in all cases were those from the legs, and these were diffusely pale and dull, with a soft friable texture. Microscopically these muscles had multiple foci of myofiber fragmentation, loss of striation, and necrosis; a mononuclear cell infiltrate was observed in muscle from two birds. These findings suggest the little bustard is susceptible to capture myopathy and that caution should be exercised during its capture and handling.     

Estimating mortality rates among passerines caught for ringing with mist nets using data from previously ringed birds

Mist netting is the most commonly used method for catching birds for scientific ringing, but despite decades of use, there have been few attempts to quantify the associated potential risks to the individuals caught. Any incidence of mortality through capture and handling, however low, is of potential ethical concern and may also introduce biases into the data. We estimate the mortality rate associated with capture of previously ringed (recaptured) passerines from the British and Irish Ringing Scheme (c. 1.5 million records) caught using mist nets. The importance of species, age, mass, month, time, previous captures, and an index of predator occurrence was tested using generalized linear mixed‐effects models. The average mortality rate was 0.0011, most of which was reported to occur before the individuals had been extracted from the nets (c. 70% of incidents). Juveniles appeared to be at higher risk and the incidence of predation from mist nets was seasonal, with increased risk during the winter. Species differed in their reported mortality rates with the apparent risk being greatest for Chiffchaff Phylloscopus collybita (0.0029) and Bullfinch Pyrrhula pyrrhula (0.0027). To improve our understanding (and hence minimize risk in future), we recommend collecting more complete data on incidences of mortality, and also injuries; exercising increased care when the species we have identified as being at greater risk are likely to be captured, and ensuring there are robust procedures for the checking of nets (as most reported incidents of mortality occur before handling). We also recommend that all Ringing Schemes should collate and make available data on capture‐related mortality. Overall rates of mortality associated with capture, although, were low and support the use of mist netting as a safe capture technique, without undue bias from mortality, when used by appropriately trained individuals.     

Trapping initiates stress response in breeding and non-breeding house sparrows Passer domesticus: implications for using unmonitored traps in field studies

In response to a variety of unpredictable conditions, birds secrete the steroid hormone corticosterone, which has numerous effects on physiology and behavior. A standardized protocol of handling and restraint has been demonstrated to elicit a robust corticosterone response in many species of birds. In contrast, comparatively little is known about the effects of capture technique on corticosterone secretion in wild birds. Setting up multiple live traps checked at regular intervals allows field researchers to capture many birds in a short period of time. However, one potential drawback of this technique is that birds may spend unknown lengths of time in traps prior to sampling. Many birds appear to remain calm and/or feed during this period, potentially leading researchers to assume that corticosterone secretion is unaffected by trapping. We assessed the impact of being left in traps for up to 30 minutes on baseline corticosterone and subsequent corticosterone responses to restraint in non‐breeding and breeding house sparrows Passer domesticus. Traps were baited with seed, and birds were either removed immediately after entry (controls), or left in the trap undisturbed for 15 or 30 min. Upon removal, birds were subjected to a standardized handling/restraint protocol in which blood samples were collected within 3 min, and again at regular intervals for 60 min. Analysis of blood samples revealed that both non‐breeding and breeding sparrows that were held in the traps had significantly higher baseline corticosterone than controls, and showed no further increase in corticosterone secretion in response to handling. However, corticosterone responses to trapping differed seasonally. Our study indicates that although birds did not exhibit prolonged escape behavior while trapped, entry into a walk‐in trap initiated a robust stress response. Taken together with data from a previous study, our data suggest that ornithologists should consider species‐specific and stage‐specific effects of trapping on physiology when designing field studies.     

Handling stress of female common eiders during avian cholera outbreaks

Researchers often consider the importance of minimizing holding time during research activities; however, the long-term costs of such handling stress is rarely measured explicitly. As part of an ongoing study of common eiders (Somateria mollissima) at a breeding colony in East Bay, Southampton Island, Nunavut, we recorded duration of restraint for females captured during avian cholera epizootics (2007 and 2008) and monitored female fates (breeding probability, onset of laying, and survival) relative to holding time. Probability of death increased with holding time in 2007 from an estimated 0.05 for females held 20 min to 0.33 for females held for 150 min. In 2008, we responded by limiting holding time to <90 min and mortality was no longer positively correlated with holding time, although total mortality was greater due to increased severity of avian cholera. In both years, longer restraint durations delayed onset of egg-laying after capture by 0.5 days for each 10 min of additional restraint but did not prevent breeding. This delay of nest initiation did not enhance survival probability. Our results show that prolonged holding time can exacerbate mortality during epizootics and emphasize the importance of minimizing restraint time in wild birds, especially in the presence of diseases. © 2011 The Wildlife Society.     

Improving our understanding of demographic monitoring: avian breeding productivity in a tropical dry forest

The ratio of juvenile to adult birds in mist‐net samples is used to monitor avian productivity, but whether it is a “true” estimate of per capita productivity or an index proportional to productivity depends on whether capture probability is not age‐dependent (true estimate) or age difference in capture probability is consistent among years (index). Better understanding of the processes affecting age‐ and year‐specific capture probabilities is needed to advance the application of constant‐effort mist‐netting for monitoring and conservation, particularly in many tropical settings where capture rates are often low. We ranked members of the avian community by capture frequencies, determined if temporary emigration influenced the availability of birds to be captured, and assessed the distribution of birds relative to mist‐nets and the parity between capture‐based productivity estimates and number of fledglings in nest plots in a tropical dry forest in Puerto Rico in 2009 and 2010. Few captures characterized the community of 25 resident species and, when estimable, capture probabilities were low, particularly for juveniles (typically < 0.1). Negative trends in capture probability, temporary emigration, and the distribution of birds suggest that avoidance of mist‐nets influenced capture rates in our study. Increasing mist‐net coverage or moving mist‐nets between sampling periods could increase capture rates. The number of fledglings observed in nest plots (25 ha/plot) did not correlate well with capture‐derived estimates (20 ha/net stations), suggesting the presence of immigrants or failure to find all nests. Our results suggest that indices of breeding productivity from mist‐netting data may track temporal changes in productivity, but such data likely do not reflect “true” productivity in most cases unless age‐specific differences in capture probability are incorporated into estimates. Pilot studies should be conducted to evaluate capture rates and the spatial extent sampled by mist‐nets to improve sampling design and inferences before informing decisions.     

Intraspecific and interspecific competition induces density‐dependent habitat niche shifts in an endangered steppe bird

Interspecific competition is a dominant force in animal communities that induces niche shifts in ecological and evolutionary time. If competition occurs, niche expansion can be expected when the competitor disappears because resources previously inaccessible due to competitive constraints can then be exploited (i.e., ecological release). Here, we aimed to determine the potential effects of interspecific competition between the little bustard (Tetrax tetrax) and the great bustard (Otis tarda) using a multidimensional niche approach with habitat distribution data. We explored whether the degree of niche overlap between the species was a density‐dependent function of interspecific competition. We then looked for evidences of ecological release by comparing measures of niche breadth and position of the little bustard between allopatric and sympatric situations. Furthermore, we evaluated whether niche shifts could depend not only on the presence of great bustard but also on the density of little and great bustards. The habitat niches of these bustard species partially overlapped when co‐occurring, but we found no relationship between degree of overlap and great bustard density. In the presence of the competitor, little bustard's niche was displaced toward increased use of the species' primary habitat. Little bustard's niche breadth decreased proportionally with great bustard density in sympatric sites, in consistence with theory. Overall, our results suggest that density‐dependent variation in little bustard's niche is the outcome of interspecific competition with the great bustard. The use of computational tools like kernel density estimators to obtain multidimensional niches should bring novel insights on how species' ecological niches behave under the effects of interspecific competition in ecological communities.     

Handling-related tail loss in an endangered skink: incidence, correlates and a possible solution

Caudal autotomy (tail loss) during capture and handling is widely reported among several families of lizards. Autotomy causes elevated stress levels in lizards, and imposes a significant fitness cost on autotomized individuals. Despite these detrimental impacts, conservation and ethical issues associated with handling-related tail loss have received little attention. We assessed the incidence and correlates of tail autotomy during capture and handling in an endangered skink, the alpine she-oak skink Cyclodomorphus praealtus . A significant proportion (9.3%) of lizards autotomized their tails during capture and handling. Medium-sized lizards were more likely to lose their tails during handling, and this effect was exacerbated at intermediate body temperatures. Probability of autotomy had a complex relationship with cumulative observer experience, independent of other risk factors. Based on the modelled relationship of autotomy with body temperature, we propose that alpine she-oak skinks be cooled immediately after capture to reduce rates of autotomy during subsequent handling.     

Isodars unveil asymmetric effects on habitat use caused by competition between two endangered species

In order for competing species to coexist, segregation on some ecological niche component is required and is often mediated by differential habitat use. When unequal competitors are involved, the dominant species tends to displace the subordinate one to its less preferred habitat. Here, we use habitat isodars, an approach which reflects evolutionary stable strategies of habitat selection, to evaluate whether interspecific competition between two competing species with distinct habitat preferences, the little bustard Tetrax tetrax and the great bustard Otis tarda, modulates their habitat use. Field data on these endangered species demonstrate that unequal competitors can coexist without completely segregating on their preferred habitats. The negatively sloped isodar of the subordinate little bustard unveils its competition with the dominant great bustard. Interference from great bustards in secondary cereal habitats reinforces use of preferred natural habitat by little bustards. Studies of density‐dependent habitat selection by a single‐species can thus aid in identifying the effects of competition on community composition, and guide the conservation of at‐risk species. Isodars, in particular, represent a promising method to gain clear knowledge on interspecific competition for species in which experimental manipulations are not feasible.     

Running out of time? The great Indian bustard Ardeotis nigriceps—status,viability, and conservation strategies

The endemic great Indian bustard (GIB) is evolutionarily trapped between open nesting and k-selection that endangers its persistence under prevailing levels of habitat loss and hunting. A global population of about 300 birds is further fragmented into eight populations in the states of Rajasthan (shared with Pakistan), Maharashtra, Andhra Pradesh, Gujarat, Karnataka, and Madhya Pradesh in India. The largest population of 100–125 birds exists in Jaisalmer, Barmer, and Bikaner districts of Rajasthan. Remaining populations number less than 35 birds each. Prevalent GIB conservation strategies use legislation to (a) secure traditional breeding areas by declaring small Protected Areas (PA) or (b) protect vast areas with varied human land uses. The vagrant nature of GIB reduces the benevolent effect of small PAs, while large reserves alienate people by curbing legitimate subsistence rights through strict legislation. These factors along with ill-informed habitat management challenge the current PA approach, even causing local extinctions. Population viability analysis shows that GIB populations of ≤35 birds can persist only under unrealistic conditions of first year mortality ≤40%, and no human caused mortality of adult birds. Even the largest population (≥100 birds) is sensitive to additional loss of adult birds to human causes. With current levels of hunting in Pakistan, extinction is a real threat. A landscape conservation strategy using conservation/community reserve concept that includes controlled traditional land uses with GIB-friendly infrastructural development is needed. The declining rate of GIB populations calls for immediate commencement of ex situ conservation breeding programs.     

Training Large Macaws for Artificial Insemination Procedures

For some endangered parrot species, captive breeding may be the only insurance for their survival. However, many individuals in captivity do not reproduce. Artificial insemination (AI) may help overcome reproductive failures or geographic distance. For semen collection in birds, massage is the most commonly used method. However, this process, which usually requires capture and restraint, involves risk of stress and injuries. The aim of this experiment was to train large macaws to accept the physical manipulations of their body parts needed for the artificial insemination process. Within 15 weeks, a male and a female Buffon's macaw (Ara ambiguus) learned to accept handling without apparent stress. A pair of green-winged macaws (Ara chloropterus) progressed more slowly and displayed some signs of stress. This stress highlights the need to monitor the birds' possible signs of discomfort during the training in order to adapt the working protocol. These results demonstrate that it is possible to train the AI behaviors, thus avoiding the capture, restraint, and anesthesia. An added benefit to this is the potential for the training to provide a form of behavioral enrichment.     

Effects of helicopter net-gunning on survival of bighorn sheep

Wildlife capture, and the data collection associated with it, has led to major advancements in ecology that are integral to decision making pertaining to wildlife conservation. Capturing wildlife, however, can cause lethal and non-lethal risks to animals. Understanding the factors that contribute to the level of risk involved in wildlife capture is therefore important for the development and implementation of the safest and most effective methodologies. We used data from 736 animal captures of 389 individuals for 2 subspecies of female bighorn sheep (Rocky Mountain bighorn [Ovis canadensis canadensis], Sierra Nevada bighorn sheep [O. c. sierrae]) in Wyoming and California, USA, in 2002–2020 to evaluate the degree and extent of time that capture via helicopter net-gunning affects survival. We compared pre- and post-capture survival during a 10-week window centered on a capture event, and post-capture survival between captured animals and animals that were monitored but not captured during the 10-week window. Additionally, we evaluated the effects of handling techniques (number of times captured, season of capture event, handling time, chase time, and body temp) and biological factors (age and nutritional condition) on probability of capture mortality. Mean daily survival was 0.9992 during a 5-week pre-capture window, dropped to 0.9864 on the day of capture, and rebounded within 3 days of capture to pre-capture levels and that of sheep that were not captured. Overall, direct mortality resulting from capture was 1.36%, with 0.54% mortality occurring within the 3 days following a capture event for an overall 1.90% capture-related mortality. The only handling and biological metrics that influenced the probability of capture mortality were rectal temperature and nutritional condition; high initial rectal temperatures and poor body condition were associated with increased risk of mortality in the days following capture. Overall, helicopter net-gunning imposed low and short-term risk to survival of female bighorn sheep. To reduce bias in survival estimates, we recommend using a 3-day censorship window for post-capture mortalities as opposed to the common practice of a 2–5-week censor window. Helicopter net-gunning, including annual or seasonal recaptures, remains an effective and comparatively safe technique for capture and associated data collection of bighorn sheep.     

Effect of isolation and conspecific presence in a novel environment on corticosterone concentrations in a social avian species, the zebra finch (Taeniopygia guttata)

Zebra finches are a highly social and monogamous avian species. In the present study, we sought to determine the effect of social isolation (separation from the flock) in a novel environment with and without a conspecific present on the adrenocortical activity of paired and unpaired individuals of this species. With regard to paired birds, we hypothesized that the presence of the mate during isolation from the group would act as a social buffer against the stressful effects of isolation. We observed that 10 but not 30 minutes of social isolation resulted in elevated concentrations of corticosterone in unpaired and paired male zebra finches in comparison to baseline concentrations of corticosterone. Furthermore, the presence of a mate during isolation in a novel environment did not have a buffering effect against increases in corticosterone concentrations. Additionally, to compare concentrations of corticosterone in response to isolation (in a novel environment) to a previously well-established stressor, we subjected groups of birds to restraint. We observed that 10 or 30 minutes of restraint led to significantly higher concentrations of corticosterone as compared to baseline. Finally, to rule out the possibility that merely handling a bird would result in significantly elevated concentrations of corticosterone as compared to baseline samples, we measured corticosterone concentrations 10 or 30 minutes after handling involving capture and release only. Our results suggest that handling alone might have contributed to the elevation of corticosterone in birds exposed to 10 minutes but not 30 minutes of restraint. Handling by itself did not account, however, for the elevated corticosterone in birds socially isolated for 10 minutes.     

A new method for catching cavity‐nesting birds during egg laying and incubation

ABSTRACT The physiological condition of female birds during the egg‐laying and incubation periods is of considerable interest and yet is relatively understudied in wild birds, primarily due to the difficulty of catching birds during this period without causing nest desertion. We therefore developed a box‐net to capture cavity‐nesting birds using sections of a mist‐net placed around a metal cubic frame. We captured female Great Tits (Parus major) as they left nest boxes during the egg‐laying and incubation periods and measured desertion rates. Using box‐nets, we captured 108 of 119 (90%) females during egg laying and 10 of 12 (83%) during incubation. Our recapture rate over two consecutive days during incubation was 50% (5 of 10). Females not captured left nest boxes before we attempted to capture them, escaped through a hole in the mist‐net, or remained in nest boxes for more than 2 h, after which we ended capture attempts. Overall, 22% of egg‐laying females deserted, with desertion rates highest early in the egg‐laying period. Desertion rates of females captured using box‐nets did not differ from those of undisturbed females. One of 10 females captured in a box‐net deserted during the incubation period. Box‐nets are portable, can be set up and taken down quickly and easily, and could potentially be used with nest boxes or natural cavities at any height. Box‐nets are easy to construct and adaptable for use with an array of cavity‐nesting birds, and can be an important tool for studying female physiology during egg laying and incubation.     

Effects of repeated investigator handling of Leach's Storm‐Petrel chicks on growth rates and the acute stress response

Disturbance during development may have lasting effects on the growth rates and stress physiology of birds. Although repeated handling by researchers is often necessary, the possible effects of such handling on the development of semi‐altricial young are unclear. We examined the effect of daily handling on growth rates and plasma corticosterone levels of Leach's Storm‐Petrel (Oceanodroma leucorhoa) chicks on Kent Island in the Bay of Fundy, New Brunswick, Canada, during the 2011 nesting season. From post‐hatch day 7 to post‐hatch days 14–36, birds in the experimental group were extracted from burrows and measured (wing, tarsus, and mass) for ～3 min every day, whereas birds in the control group were left undisturbed. After the treatment period, blood was collected from birds in both groups within 3 min of initially reaching into burrows (baseline) and after a 30‐min restraint stress test to assess the effect of early life disturbance on programming of the hypothalamic‐pituitary‐adrenal (HPA) axis. A second acute restraint stress test was conducted three weeks after the end of the treatment period to investigate possible longer term effects of early life disturbance. Growth rates of wings and tarsi were similar for handled chicks (N = 18) and non‐handled control chicks (N = 21), as were baseline and 30‐min acute restraint stress‐induced corticosterone levels. As also reported in previous studies of other altricial and semi‐altricial species, older chicks (42–64 d old) had higher plasma corticosterone levels than younger chicks (21–43 d old) after acute restraint stress tests, reflecting delayed development of the HPA axis. The age‐related increase in HPA axis sensitivity observed prior to fledging could facilitate foraging and predator avoidance behaviors while minimizing exposure to high levels of corticosterone earlier in development. Overall, we found no evidence that repeated disturbance influenced either growth rates or HPA axis programming of Leach's Storm‐Petrel chicks.     

DIFFERENCES IN THE FORAGING OF JUVENILE AND ADULT BIRDS: THE IMPORTANCE OF DEVELOPMENTAL CONSTRAINTS

Juvenile birds differ from conspecific adults in their diet and methods of prey capture and prey handling. Juvenile-adult differences in foraging result from (1) immaturity of the beak, skeleto-muscular and neurological systems and (2) the time required to learn foraging skills. These conclusions are largely based on field observations. More experimental studies to assess the relative importance of the various constraints are needed.
Juvenile birds appear to be under strong selection to reach adult form and function as rapidly as possible. Remaining differences between juveniles and adults are largely attributable to constraints. In contrast to many other taxa there are few examples in which juvenile-adult differences in foraging have been accentuated by selection on the juvenile behaviours.     

Midazolam as an adjunctive therapy for capture myopathy in Bar-tailed Godwits (Limosa lapponica baueri) with prognostic indicators

Capture myopathy is a complication of capture and handling in many species of birds and mammals. Muscular necrosis leads to ataxia, paralysis, and pain, whereas metabolic disturbances can result in death. We conducted an opportunistic clinical trial on Bar-tailed Godwits (Limosa lapponica baueri) that developed capture myopathy after a cannon-net capture in New Zealand in October 2008. We assessed the beneficial effects of midazolam, a benzodiazepine with the effects of anxiolysis, muscle relaxation, and sedation, in the adjunctive treatment of capture myopathy. Physical and biochemical parameters were analyzed retrospectively for their potential as indicators for survival until release. Birds (n=16) were treated with subcutaneous fluid therapy, a nonsteroidal anti-inflammatory (meloxicam), gavage feeding, and sling therapy twice daily. The treatment group (n=8) was treated twice daily with intramuscular midazolam injections, 1.5 mg/kg. Surviving godwits were released over 1-9 days, with 6 of 8 treated birds (75%) surviving to release, compared with 3 of 8 controls (38%). Inability to counteract weight loss in captivity was the most significant problem for both groups. Lack of waterproofing and predation were contributing causes of death for at least two godwits after release. Birds treated with midazolam showed subjective benefits including improved tolerance of handling and sling therapy. Clinical parameters (change in body mass, packed cell volume [PCV], plasma creatine kinase [CK], aspartate aminotransferase [AST], total protein, and uric acid [UA] over time) were not statistically different between groups, although peak average values for CK, AST, and UA were lower in the treatment group. Decline in body mass (%), PCV, final plasma UA, and peak plasma CK were the most useful prognostic indicators. Midazolam shows potential as an ancillary treatment for capture myopathy in birds and is worthy of continued study and use.     

Prey handling time and ingestion probability for Mesocyclops sp. predation on small cladoceran species Bosmina longirostris, Bosminopsis deitersi, and Scapholeberis mucronata

We compared the vulnerability of small cladoceran species (Bosminopsis deitersi, Bosmina longirostris, and Scapholeberis mucronata) to predation by Mesocyclops sp. in the laboratory based on prey handling time and ingestion probability. We also estimated the effects of ontogenetic changes on handling time by testing prey of various body sizes. All tested prey species showed an increasing pattern of handling time along with growth (increase of body size). For juveniles smaller than 0.3mm, the prey handling time was similar for B. deitersi and B. longirostris, but markedly longer for S. mucronata. Mesocyclops sp. rejected large (>0.35mm) B. longirostris and S. mucronata soon after capture and consequently showed a low probability of successful ingestion, whereas most large individuals of B. deitersi were consumed. The shorter handling time and higher ingestion probability of small juveniles indicate that the copepods ingest small juveniles more efficiently than larger individuals. Thus, it seems that the vulnerability of small cladocerans to copepod predation differs ontogenetically, with the greatest vulnerability being during the juvenile period. The ontogenetic reduction in the vulnerability of B. longirostris and S. mucronata was shown in the rapidly reduced ingestion probability as well as by the increase in prey handling time for large individuals. The results show that the vulnerability to predation by copepods of B. longirostris and S. mucronata reduces more quickly with increasing size than does that of B. deitersi, and it seems that the former two species are more resistant to copepod predation.