Alarm cue induces an antipredator morphological defense in juvenile Nicaragua cichlids Hypsophrys nicaraguensis

Olfactory cues that indicate predation risk elicit a number of defensive behaviors in fishes, but whether they are sufficient to also induce morphological defenses has received little attention. Cichlids are characterized by a high level of morphological plasticity during development, and the few species that have been tested do exhibit defensive behaviors when exposed to alarm cues released from the damaged skin of conspecifics. We utilized young juvenile Nicaragua cichlids Hypsophrys nicaraguensis to test if the perception of predation risk from alarm cue （conspecific skin extract） alone induces an increased relative body depth which is a defense against gape-limited predators. After two weeks of exposure, siblings that were exposed to conspecific alarm cue increased their relative body depth nearly double the amount of those exposed to distilled water （control） and zebrafish Danio rerio alarm cue. We repeated our measurements over the last two weeks （12 and 14） of cue exposure when the fish were late-stage juveniles to test if the rate of increase was sustained; there were no differences in final dimensions between the three treatments. Our results show that 1） the Nicaragua cichlid has an innate response to conspecific alarm cue which is not a generalized response to an injured fish, and 2） this innate recognition ultimately results in developing a deeper body at a stage of the life history where predation risk is high [Current Zoology 56 （1）： 36-42, 2010].

Antipredator Behavioral Responses of Native and Exotic Tadpoles to Novel Predator

Factors related to the invasion process, such as high abundance of invaders, residence time, and functional distinctiveness, are well documented, but less attention has been given to the effects of antipredator strategy of invasive species during colonization. In this study, we explored the antipredator strategy of an introduced species by comparing the predator avoidance behaviors of two native anuran species and one introduced （“exotic”） species in the presence of different predators. The two native anuran species used in the study were Black-spotted Pond FrogRana nigromaculata and Terrestrial Frog Rana limnocharis. The introduced （invasive） species used was American bullfrogLithobates catesbeianus. Chinese pond turtleChinemys reevesii, Red-backed rat snakeElaphe rufodorsata, and Big-headed turtle Platysternon megacephalum were used as predator species. Chinese pond turtles and Red-backed rat snakes are native predators of Black-spotted Pond Frogs and Terrestrial Frogs, while Big-headed turtles are novel （“unfamiliar”） to the two frogs. All three predator species are novel （“unfamiliar”） to the American bullfrog. The results show that tadpoles of the two native species displayed behaviors of recognizing the two native predators, but did not display the capability of identifying the novel predator. Results from our study also suggest that American bullfrog tadpoles exhibited strong antipredator behavioral responses by displaying the capability of identifying “unfamiliar” predators without cohabitation history and prior exposure to them. Such antipredator behavioral responses could have resulted in more favorable outcomes for an invading species during the invasive introductory process.

Same place, different time, head up: Multiple antipredator responses to a recolonizing apex predator

Prey adjust their antipredator behavioral tactics to minimize the risk of an encounter with predators.Spatiotemporal responses of prey to pred-ators have been reported,but the nature of antipredator response is not ubiquitous and it is the object of increasing interest,especially consid-ering the recent recovery of large carnivores in Europe,and the potential for behavioral antipredator responses to elicit consequences at the ecosystem level.We have tested multiple antipredator responses by fallow deer Dama dama to wolf Canis lupus in a Mediterranean protected area recently recolonized by this apex predator.Through intensive camera trapping,we tested for temporal and spatial association between predator and prey,and we have also studied deer vigilance in forest habitats where focal observations are usually impossible.Wolf detection rates were spatially associated with those of fallow deer.Accordingly,no evidence was found for fallow deer avoiding sites with higher predator detection rates.Temporal activity patterns were significantly different between the 2 species,with the wolf being mainly nocturnal whereas fallow deer was active especially during daylight.A comparison with a preliminary study strongly suggests an increase in the diurnal activity of fallow deer along with the stabilization of wolf presence in the area.Both the rate and the duration of vigilance of female fallow deer increased with the local frequency of wolf activity.We suggest an antipredator response based on temporalrather than spatialavoidance,as well as increasedvigilance.

Antipredatory reaction of the leopard gecko Eublepharis macularius to snake predators

Ability to recognize a risk of predation and react with adaptive antipredatory behavior can enhance fitness, but has some costs as well. Animals can either specifically react on the most dangerous predators （threat-sensitive avoidance） or they have safe but costly general wariness avoiding all potential predators. The level of threat may depend on the predator＇s foraging ecology and distri- bution with the prey with sympatric and specialist species being the most dangerous. We used 2 choice trials to investigate antipredatory behavior of captive born and wild-caught leopard geckos confronted with different snake predators from 2 families （Colubridae, Boidae） varying in foraging ecology and sympatric/allopatric distribution with the geckos. Predator-na＇ive subadult individuals have general wariness, explore both chemically and visually, and perform antipredatory postures toward a majority of snake predators regardless of their sympatry/allopatry or food specialization. The most exaggerated antipredatory postures in both subadult and adult geckos were toward 2 sympatric snake species, the spotted whip snake Hemorrhois ravergieri, an active forager, and the red sand boa Eryxjohnii, a subterranean snake with a sit-and-wait strategy. In contrast, also subter- ranean but allopatric the Kenyan sand boa Eryx colubrinus did not elicit any antipredatory reaction. We conclude that the leopard gecko possesses an innate general antipredatory reaction to different species of snake predators, while a specific reaction to 2 particular sympatric species can be observed. Moreover, adult wild caught geckos show lower reactivity compared with the captive born ones, presumably due to an experience of a real predation event that can hardly be simulated under laboratory conditions.

Predatory Cues Influence the Behavioral Responses and Metamorphic Traits of Polypedates maculatus(Anura: Rhacophoridae)

Mechanisms of predator detection and the influence of the presence of nonlethal predators on antipredator defense behavior and metamorphic traits were studied in the Indian tree frog, Polypedates maculatus. Exposure of P. maculatus tadpoles to chemical cues of caged predator（crabs, Barytelphusa spp.） fed with either conspecific or heterogeneric tadpoles, or were starved elicited defense behavior（by avoiding predator zone） in them. Such a behavior was not evident when exposed to predators housed in a glass beaker（visual cues）. Both early（Gosner stage 27–28） and later（Gosner stage 35–36） stage tadpoles when exposed to caged predators（fed with conspecific tadpoles）, prey tadpoles spent less time swimming and remained motionless for longer periods. Yet, the time spent by prey in feeding was unaffected. Further, the predator avoidance behavior exhibited by them was of the same intensity regardless of whether the caged predators were fed or starved implying the influence of predator＇s kairomones. Tadpoles reared with caged predator reached the metamorphic climax stage（MC stage; Gosner stage 42） earlier than those reared without a predator. Size at emergence（Gosner stage 46） was comparable in both the groups. The findings suggest that P. maculatus tadpoles assess predation risk chiefly by sensing kairomones of the predator in eliciting antipredator defense behaviors. Accelerated development and early metamorphosis without any compromise of the size at emergence may be due to their unaltered feeding activity.

The Relationship between Shell Morphology and Crevice Size Affecting Retreat Selection of the Keeled Box Turtle(Cuora mouhotii)

Rock crevices are important retreat sites for many reptiles.Unlike lizards and snakes,crevicedwelling turtles have rigid shells which severely limit the size and shape of crevices they are able to use.Few studies have addressed the relationship between turtle's shell morphology,behavior,and crevice size.Here,we investigate the relationship between morphology and crevice size selection in the Keeled box turtle(Cuora mouhotii).We radio tracked turtles in the field to document crevice dimensions and turtle behavior associated with crevices,and we conducted a controlled experiment in the lab to understand how a turtle's morphology influences its decision to select crevices with different dimensions.Both laboratory and field experiments demonstrate that turtles select deep and moderately wide crevices that are low enough to barely accommodate the carapace.Crevices used by turtles tend to be about 2 times as wide,1.2 times as high,and 3 times longer than the turtle's shell.Moreover,turtles tended to prefer crevices with horizontal openings in the field and position themselves head first in the crevice.We conclude that morphology,especially carapace height,influences the Keeled box turtles' decision to select specific sizes of crevice.We argue that the tendency for turtles to select crevices that come into contact with their carapace contributes to successful defense against predators,as it would be more difficult for a predator to remove a turtle when the crevice is nearly touching the turtles shell.

A test of the species confidence hypothesis in dusky damselfish

Visual cues are important in both interspecific and intraspecific communication.The species confidence hypothesis proposes that animals are more attracted to conspecific colors and repelled by colors,not on their bodies.Studies on terrestrial lizards and birds have tested the species confidence hypothesis and shown that conspecific colors elicit reduced antipredator behavior.To date,the species confidence hypothesis has not been tested in the marine environment,specifically on coral reefs where color communication is of vital importance.We addressed this knowledge gap by measuring flight initiation distance(the distance an individual moves away from an approaching threat)in dusky damselfish(Stegastes nigricans)in response to an approaching disc of 1 of 4 different color treatments:conspecific,blue,yellow,and black.If the species confidence hypothesis explained variation in damselfish flight initiation distance,then we expected individuals to tolerate closer approaches when approached by a conspecific color.In addition,we calculated the color difference between each stimulus and its corresponding back-ground as a potential alternative explanation for flight responses.Damselfish tolerated the closest approach from the conspecific color stimulus;there were no significant differences between other colors and there was no support for the alternative color difference hypothesis.As with similar terrestrial studies,these results are relevant to ecotourists'choice of swimsuit and wetsuit colors because color choice may modify naturalantipredatorbehavior.

Differences in predator-avoidance behavior between two invasive gobies and their native competitors

Globally,fish are frequently introduced beyond their native range.Some,like Ponto-Caspian gobies,are becoming invasive,achieving high colo-nization rates and constituting frequent prey for native predators.However,little is known about the effectiveness of antipredator behaviors of the invaders,which may shape their role in the invaded community and contribute to the invasion success.We compared antipredator behaviors of invasive gobies and native fish species after their detection by the predator,when the danger becomes direct.We studied 2 fish pairs,each consisting of an invasive and native species co-occurring in the environment and belonging to the same prey guild:(1)the racer goby Babka gymnotrachelus versus European bulhead Cottus gobio,(2)the monkey goby Neogobius fluviatilis versus gudgeon Gobio gobio,facing a naive predator(the Eurasian perch Perca fluviatilis).We analyzed behaviors of single prey individuals(escaping,staying in shelter,and activity)and sin-gle predators(activity,searching,following,capturing,and latency to prey consumption).In the predator presence,the bullhead was less active and more often managed to escape after capture than the racer goby.The gudgeon escaped before the capture more often than the monkey goby.The predator succeeded later with the bullhead compared to racer goby,whereas no differences in ingestion time occurred between the gudgeon and monkey goby.The results suggest that,in terms of hunting effort of native predators,the invasive gobies are equivalent to or more profitable prey than their native analogs,which can facilitate the integration of the gobies into local food webs.

Local predation pressure predicts the strength of mobbing responses in tropical birds

Many birds join cooperative mobbing aggregations and collectively harass predators. Individuals participating in these ephemeral associations benefit by deterring the predator, but also incur energetic costs and increased risk of predation. Ex- plaining the evolution of mobbing is challenging because individuals could prevail by selfishly seeking safety while allowing others to mob. An important step in understanding the evolution of mobbing is to identify factors affecting its expression. The ecological constraints model suggests that animals are more likely to cooperate under adverse environmental conditions, such as when local predation pressure is high. We tested this prediction by comparing the mobbing responses of several species of birds to the local abundance of their primary predator, the ferruginous pygmy-owl Glaucidium brasilianum. We used acoustic playback to elicit mobbing responses in environments where owls were common, uncommon, or rare. Stimuli were either the song of a fer- ruginous pygmy-owl or the mobbing calls of three of the owl＇s common prey species. During each playback, we characterized mobbing responses by noting the number of species and individuals that approached the loudspeaker, as well as the closest ap- proach by any bird. Mobbing responses to both stimuli were strong in locations where Ferruginous Pygmy-owls were common, intermediate where owls were uncommon, and weak where they were rare. This pattern persisted even after controlling for dif- ferences in species richness and composition among the three environments. Results support the ecological constraints model and provide strong evidence that intense predation pressure increases the expression of cooperative mobbing in tropical birds [Cur- rent Zoology 58 （5）： 781-790, 2012].

Determinants of vigilance in a reintroduced population of Pere David＇s deer

After being kept in captivity and isolated from natural predators for more than 1,200 years, P6re David＇s deer has been reintroduced in China and now occurs in a reserve where human activity is the only potential threat. Antipredator vigilance i~ an important component of survival for many prey animals in their natural habitat. Do deer still adjust vigilance as a function of risk after such a long period of relaxed predation pressure？ Here, we examined vigilance levels in P6re David＇s deer groups as a function of group size, sex and level of human disturbance. The results showed that individual vigilance significantly decreased with group size in all-female groups but not in all-males or mixed-sex groups. In rutting season, males compete with one another and harass females, and we argue that vigilance is partly aimed at threatening males and that such vigilance increases with group size. This explains why overall vigilance did not vary with group size for males in general and for females in mixed-sex groups. Vigilance increased in more disturbed areas but in in male deer only. The results indicate that despite relaxed predation pressure over centuries, P6re David＇s deer can still adjust antipredator responses as a function of perceived risk. Such information may become useful in the rewilding programme now under way for this species in China [Current Zoology 59 （2）： 265-270, 2013].

Marmots do not consistently use their left eye to respond to an approaching threat but those that did fled sooner

In many vertebrates,the brain's right hemisphere which is connected to the left visual field specializes in the processing of information about threats while the left hemisphere which is connected to the right visual field specializes in the processing of information about conspecifics.This is referred to as hemispheric lateralization.But individuals that are too predictable in their response to predators could have reduced survival and we may expect selection for somewhat unpredictable responses.We studied hemispheric lateralization in yellow-bellied marmots Marmota flaviventer,a social rodent that falls prey to a variety of terrestrial and aerial predators.We first asked if they have lateralized responses to a predatory threat.We then asked if the eye that they used to assess risk influenced their perceptions of risk.We recorded the direction marmots were initially looking and then walked toward them until they fled.We recorded the distance that they responded to our experimental approach by looking,the eye with which they looked at us,and the distance at which they fled (i.e.,flight initiation distance;FID).We found that marmots had no eye preference with which they looked at an approaching threat.Furthermore,the population was not comprised of individuals that responded in consistent ways.However,we found that marmots that looked at the approaching person with their left eye had larger FIDs suggesting that risk assessment was influenced by the eye used to monitor the threat.These findings are consistent with selection to make prey less predictable for their predators,despite underlying lateralization.

Does disruptive camouflage conceal edges and features？

Camouflage is ubiquitous in the natural world and benefits both predators and prey. Amongst the range of conceal- ment strategies, disruptive coloration is thought to visually fragment an animal＇s＇ outline, thereby reducing its rate of discovery. Here, I propose two non-mutually exclusive hypotheses for how disruptive camouflage functions, and describe the visual me- chanisms that might underlie them. （1） The local edge disruption hypothesis states that camouflage is achieved by breaking up edge information. （2） The global feature disruption hypothesis states camouflage is achieved by breaking up the characteristic features of an animal （e.g., overall shape or facial features）. Research clearly shows that putatively disruptive edge markings do increase concealment; however, few tests have been undertaken to determine whether this survival advantage is attributable to the distortion of features, so the global feature disruption hypothesis is under studied. In this review the evidence for global feature disruption is evaluated. Further, I address if object recognition processing provides a feasible mechanism for animals＇ features to influence concealment. This review concludes that additional studies are needed to test if disruptive camouflage operates through the global feature disruption and proposes future research directions [Current Zoology 61 （4）： 708-717, 2015].

Take only pictures, leave only...fear？ The effects of photography on the West Indian anole Anolis cristatellus

Ecotourism encourages an environmentally friendly exploration of the world＇s natural habitats. Tourists often engage in wildlife photography, an activity that is generally not considered disturbing to animals. We investigated the effects of cam- era-related stimuli to determine whether shutter noise and/or flash affected the immediate behavior of female crested anoles Ano- lis cristaellus. Anoles decreased their display rate following stimuli that included shutter noises, but did not change their behavior in response to flash or silence treatments. To determine the relative importance of this response, we observed anole behavior fol- lowing playbacks of calls from kestrels Falco sparverius, a predator, and bananaquits Coereba flaveola, a non-predator. Anoles decreased display rates following kestrel calls when compared to their response to bananaquit calls. Furthermore, anoles spent a greater proportion of time displaying following bananaquit calls compared to both kestrel calls and silence. The magnitude of response to shutter noises was about the same as that to predator calls. This demonstrates that photography may not be as benign as commonly believed, and we should consider whether restrictions on camera noises should be implemented to reduce ani- mal disturbance

Ontogenetic shifts in perceptions of safety along structural complexity gradients in a territorial damselfish

Age and body size can influence predation risk and hence habitat use. Many species undergo ontogenetic shifts in habitat use as individuals grow larger and have different age-specific predation pressures. On coral reefs, a number of fish species are more tolerant of threats in structurally complex habitats that contain more refuges than in less structurally complex habitats. However, we do not know how risk perception varies with age, and whether age interacts with habitat complexity. Adults and juveniles, because of their size, may face different risks in structurally simple versus complex habitats. We used flight initiation distanee as a metric to analyze perceptions of risk in a species of damselfish Stegastes nigricans. All else being equal, fish fleeing at greater distances are inferred to perceive higher risk. We targeted juvenile and adult damselfish to assess whether there are ontogenetic shifts in perceptions of safety in relation to structural complexity, inferred based on percent coral cover and rugosity. We found that adult damselfish tolerated closer approach in more complex habitats as measured by percent coral cover, but not rugosity, whereas juvenile fish always allowed closer approach than adult fish regardless of complexity. This ontogenetic shift in habitat use may result from juvenile fish taking bigger risks to maximize growth, whereas older animals, who are closer to their maximum body size, can afford to take fewer risks and protect their assets.

Together We Stand-Analyzing Schooling Behavior in Naive Newborn Guppies through Biorobotic Predators

A major advantage of animal aggregations concerns cooperative antipredator strategies.Schooling behavior emerges earlier in many fish species,especially in those cannibalizing their offspring.Experience is fundamental for developing schooling behavior.However,the cognitive ability of naive newborn fish to aggregate remains unclear.Herein,Poecilia reticulata,was selected as model organism to investigate how combinations of biomimetic robotic agents and adult conspecific olfactory cues affect collective responses in newborns.The role of white and brown backgrounds in evoking aggregations was also assessed.Olfactory cues were sufficient for triggering aggregations in P.reticulata newborns,although robotic agents had a higher influence on the group coalescence.The combination of robotic agents and olfactory cues increased schooling behavior duration.Notably,schooling was longer in the escape compartment when robotic agents were presented,except for the combination of the male-mimicking robotic fish plus adult guppy olfactory cues,with longer schooling behavior in the exploring compartment.Regardless of the tested cues,newborn fish aggregated preferentially on the brown areas of the arena.Overall,this research provides novel insights on the early collective cognitive ability of newborn fish,paving the way to the use of biomimetic robots in behavioral ecology experiments,as substitutes for real predators.

Consistent individual differences in fathering in threespined stickleback Gasterosteus aculeatus

There is growing evidence that individual animals show consistent differences in behavior. For example, individual threespined stickleback fish differ in how they react to predators and how aggressive they are during social interactions with con- specifics. A relatively unexplored but potentially important axis of variation is parental behavior. In sticklebacks, fathers provide all of the parental care that is necessary for offspring survival; therefore paternal care is directly tied to fitness. In this study, we assessed whether individual male sticklebacks differ consistently from each other in parental behavior. We recorded visits to nest, total time fanning, and activity levels of 11 individual males every day throughout one clutch, and then allowed the males to breed again. Half of the males were exposed to predation risk while parenting during the first clutch, and the other half of the males ex- perienced predation risk during the second clutch. We detected dranlatic temporal changes in parental behaviors over the course of the clutch： for example, total time fanning increased six-fold prior to eggs hatching, then decreased to approximately zero. De- spite these temporal changes, males retained their individually-distinctive parenting styles within a clutch that could not be ex- plained by differences in body size or egg mass. Moreover, individual differences in parenting were maintained when males re- produced for a second time. Males that were exposed to simulated predation risk briefly decreased fanning and increased activity levels. Altogether, these results show that individual sticklebacks consistently differ from each other in how they behave as parents [Current Zoology 58 （1）： 45-52, 2012].

Vivid birds do not initiate flight sooner despite their potential conspicuousness

The distance from an approaching threat at which animals initiate flight - flight-initiation distance （FID） -- is a sensitive metric of variation in risk, but the effects on FID associated with the risk of possessing highly detectable external coloration are unknown. We tested whether variation in the degree of plumage vividness in birds explained variation in flight-initiation distance. After controlling for body mass, the distance at which the experimental approach began, and phylogenetic relatedness, plumage vividness was not a predictor of FID. Contrary to the expectation that vividness affects risk, and therefore risk assess- ment, these results suggest that birds do not compensate for greater visual conspicuousness by fleeing sooner from approaching threats [Current Zoology 61 （4）： 773-780, 2015].

Urban invaders are not bold risk-takers: a study of 3 invasive lizards in Southern California

Biological invasions threaten biodiversity worldwide,and therefore,understanding the traits of successful invaders could mitigate their spread.Many comm only invasive species do well in disturbed habitats,such as urban environments,and their abilities to effectively respond to disturbarices could con tribute to their invasiveness.Yet,there are non invasive species that also do well in disturbed habitats.The question remains whether urban invaders behave differently in urban environments than noninvaders,which could suggest an"urban-exploiting"phenotype.In Southern California,the co-occurrence of in vasive Italia n wall lizards Pod arc is siculus,brown anoles Anolis sagrei,and green anoles A.carolinensis,and native western fence lizards Sceloporus occidentalis offers an opportunity to test whether invasives exhibit consistent differences in risk-taking within human-altered habitats compared with a native species.We predicted that invasive lizards would exhibit more bold behavior by having shorter flight-initiation distances(FIDs)and by being found farther from a refuge(behaviors that would presumably maximize foraging in low-risk environments).Invasive populations had similar or longer FIDs,but were consistently found at distances closer to a refuge.Collectively,invasive lizards in urban habitats were not bolder than a native species.Relianee on nearby refuges might help species successfully invade urban habitats,and if a general pattern,may pose an added challenge in detecting or eliminating them.

Anti-predator behavior along elevational and latitudinal gradients in dark-eyed juncos

Flight-initiation distance(FID),the distance between an individual and experimenter when it begins to flee,can be used to quantify risk-assessment.Among other factors,prior studies have shown that latitude explains significant variation in avian FID:at lower latitudes,individuals and species have longer FIDs than those living at higher latitudes.No prior studies have focused on the effect of elevation on FID.Given the similar patterns of seasonality,climate,and potentially predator density,that covary between latitude and elevation,birds at higher elevations might tolerate closer approaches.We asked whether elevation or latitude would explain more variation in the FID of a common passerine bird species,dark-eyed juncos(Junco hyemalis).Juncos live in a variety of habitats along both latitudinal and elevational gradients.We found that statistical models containing elevation as a variable explained more of the variation in FID than did models containing latitude.We also found,unexpectedly,that birds at higher elevation fled at greater distances.While more predators were sighted per hour at higher elevations than at lower elevations,the frequency of predator sightings did not explain a significant amount of variation in FID.This result questions whether predator density is the main driver of risk perception along elevational gradients.Nonetheless,because elevation explains more variation in FID than latitude in at least one species,these findings have direct implications on how human impacts on birds are managed.Specifically,those designing set-back zones to reduce human impact on birds may consider modifying them based on both latitude and elevation.

Nesting innovations allow population growth in an invasive population of rose-ringed parakeets

Certain traits of recipient environments,such as the availability of limiting resources,strongly determine the establishment success and spread of non-native species.These limitations may be overcome through behavioral plasticity,allowing them to exploit alternative resources.Here,we show how a secondary cavity nester bird,the rose-ringed parakeet Psittacula krameri,innovates its nesting behavior as a response to the shortage of tree cavities for nesting in its invasive range in Tenerife(Canary Islands).We observed that some breeding pairs excavated their own nest cavities in palms,thus becoming primary cavity nester,whereas others occupied nests built with wood sticks by another invasive species,the monk parakeet Myiopsitta monachus.The use of these novel nesting strategies increased the number of breeding pairs by up to 52%over 6 years,contributing to a 128.8%increase of the whole population.Innovative nests were located at greater heights above ground and were more aggregated around conspecifics but did not result in greater breeding success than natural cavities.Occupation of monk parakeet colonies by rose-ringed parakeets also benefited the former species through a protective-nesting association against nest predators.Our results show how an invasive species innovate nesting behaviors and increase nest-site availability in the recipient environment,thus facilitating its population growth and invasion process.Potential behavioral innovations in other invasive rose-ringed parakeet populations may be overlooked,and should be considered for effective management plans.