Survey
* Your assessment is very important for improving the workof artificial intelligence, which forms the content of this project
* Your assessment is very important for improving the workof artificial intelligence, which forms the content of this project
10/16/2012 Spines and such: morphological traits suggest that predation exerts strong selection. Three stages in the process of predation: Detection Attack Major Theme: Behavior enhancing morphology (Why not always be a ball of spines?) Strategies for Predator Avoidance A. Avoid Detection Consumption Let’s look at predator avoidance at each stage. Crypsis (= Camouflage) Cephalopods can change their colors to match a variety of different backgrounds. Pepper moths are cryptic only if they match themselves against background.* * these are all the same moths placed against different backgrounds 1 10/16/2012 Cuttlefish match variety of backgrounds. Cephalopods, like squid or octopus, can tune colors and patterns to background against which they rest. Even though they don’t see in color. Seems cognitively sophisticated. Deception Walking stick insects not only look like twigs but, in the wind, sway like twigs… They are mimicking swaying of twig! Predators may turn the tables. preying mantis floral mimicry 2 10/16/2012 False Head Mimics (cont’d) False Head Mimics Scolopendra, our desert centipede, has a false head that looks like the real one. Predator has <50% chance of grabbing the right end… If it doesn’t, it gets bit. False head at rear of butterfly has ‘antennae’ that move! Attacks to the false head result in a mouthful of scales. Strategies for Predator Avoidance B. Avoid Capture and Consumption Active Defense Many lizards have tails that are built to break off easily. Termed caudal autotomy http://www.youtube.com/watch?v=T sQRzRcQ3dM Broken tail writhes vigorously, distracting the predator from escaping lizard… Grasshoppers and relatives lose legs in the same way... Another example of convergent evolution. Skunk sitting in its own…. pew. Bombardier beetle sprays attacker; Beetle can aim spray with precision; Spray is irritating benzoquinone in boiling water. vinegaroon Chemical defenses are everywhere! 3 10/16/2012 hydrogen peroxide (H2O2) and hydroquinone in ‘reservoir’ Catalases and peroxidases in ‘reaction chamber’ When attacked, reservoir empties into reaction chamber; Enzymes catalyze production of benzoquinone, O2, and HEAT. Elegant ‘design.’ But presumed to have evolved via gradual process of natural selection. There are a variety of species of spraying beetles that provide for a series of intermediate stages. O2 causes boiling solution to propel out of back end! Stealing defenses from prey nudibranchs incorporate stinging cells called cnidocytes from jellyfish into their tissues. They are defended from predation by their prey… Nematocyst discharge occurs in ca. 700 nanoseconds. Force generated equivalent to that of bullet. Warning Coloration (= Aposematism) Some butterflies sequester noxious chemicals from host plants while feeding as caterpillars. Pipevine swallowtail butterfly Poison arrow frogs Monarch butterfly Same strategy… must be independently evolved =CONVERGENT EVOLUTION. (Dendrobates spp.) are toxic, advertising toxicity with warning coloration, also called aposematic coloration. 4 10/16/2012 Spotted skunk Enhancing warning pattern by doing handstands. VIDEO red eft ladybird beetles Do predators learn high-contrast prey better than low-contrast prey? lionfish Predator: Tenodera aridifolia Why be conspicuous? monarch butterfly Prey: Oncopeltus fasciatus Project Leader: Katy Prudic Why not let the predator learn to avoid your cryptic pattern? Prudic, Skemp & Papaj, Behavioral Ecology 2007 10 8 Mean No. Trials to Criteria (+ 1 s.e.) 6 4 2 Toxic bugs painted gray: either conspicuous or cryptic. Bug released at bottom and walked up the ramp to meet its fate. 0 Cryptic Conspicuous 20 15 Mean No. Days to Re10 Sampling (+ 1 s.e.) 5 0 Cryptic Conspicuous Prey Type Mantid first trained to avoid a given morph, then its retention tested. Mantids learned conspicuous prey faster and retained memory of aversion longer. 5 10/16/2012 Monarch butterfly is toxic, owing to cardenolides sequestered as caterpillars from milkweed host plants. Viceroy butterfly is unrelated but also toxic; Jays learn to avoid patterns associated with noxious experience. = Mullerian mimicry mimicry of each other protects both better. Having two species with same pattern may facilitate learning experience. mimic Two “mimicry rings” from tropics coral snake mimic Coral snakes are highly poisonous. Once thought to serve as a model for various harmless mimic species. models & Mullerian mimics Batesian mimics* But how can mimics benefit if predators die from coral snake bites? mimic *Batesian mimics are not harmful. Coral snakes may mimic less poisonous relatives in South America! By mimicking less poisonous snakes, the coral snake benefits from the predator’s learning experience. spider mimicking ant 6 10/16/2012 Dynamic mimicry Indo-Malayan octopus impersonates: • • • • http://www.youtube.com/watch?v=Owelt6QKMKo&NR=1 http://www.youtube.com/watch?v=t-LTWFnGmeg&feature=related poisonous flatfish banded sea snakes lionfish jellyfish mimic can rapidly swap between 'impersonations' http://www.youtube.com/watch?v=H8oQBYw6xxc “Red against yellow can kill a fellow.” “Red against yellow can kill a fellow.” Coral Snake Story Re-Visited Coral Snake Story Re-Visited 1. “Mertensian mimicry”: more poisonous coral snake may mimic less poisonous relatives. 1. “Mertensian mimicry”: more poisonous coral snake may mimic less poisonous relatives. 2. Predators evolve innate avoidance of corallike pattern. 2. Predators evolve innate avoidance of corallike pattern. #2 seems to be more accepted. Ultimate Mimicry of Bad-Tasting Stuff Why #2? Partial answer: Because coral snake dummies attacked more where coral snake is absent. Multiple Strategies for Predator Avoidance Bluff: hiss & look big Hognose snake Backup defenses are common. Bird dropping mimic! Play dead *Look for these swallowtail caterpillars on citrus plants. 7 10/16/2012 All-in-One Strategies Crypticity – Aposematism Tradeoff Black swallowtail butterfly Caterpillar is first a poop mimic But later adopts a pattern which is cryptic from a distance but conspicuous close up. Multimodal Warning Displays Rattlesnakes warn in both visual modality (black & white tail) and acoustic modality (rattle). Why multiple sensory modalities? Rattlesnakes are cryptic from side (to prey) but conspicuous from above (to would-be predators). Multimodal Warning Displays When threatened, a gopher snake mimics rattlesnakes. It has a striped tail, makes its head triangular, and generates a hiss that can sound like a rattle. In other words, it’s a multimodal mimic. Predator Avoidance and Being in a Group Whirlygig beetles and per capita risk of predation Large groups more likely to be attacked by fish. BUT… per capita risk of predation goes down. It is per capita risk that is important to consider. 8 10/16/2012 Active group defense Prey defense is more effective when done by multiple individuals. mobbing behavior Musk oxen form a circle with heads outwards, soldier aphid Sawfly larvae do something similar… 6.13 Communal defense by sawfly larvae Spit and Poop! such that each individual has his back end covered... Dilution of risk prey so locally abundant that local predator fauna is satiated, which reduces per capita risk grunion running at full moon Butterflies puddling on damp soil… monarchs roosting Predation risk decreases with group size Mmm… amino acids… 9 10/16/2012 Confusion effect Predators may have difficulty choosing a target out of a tightly-packed group Zebra stripes make it even more difficult for a predator to focus on an individual. Selfish herd Prey use each others as shields. Animals attempt to move to the center of the group. Leopard seal Adelie penguin Average per capita risk can actually be greater than if animals spread themselves out. sheep fitted with GPS unit http://www.youtube.com/watch?v=o54zO30lnas Sheep show selfish herd behavior in response to a herding dog. 10 10/16/2012 Group vigilance and early detection Vigilance for predators may be facilitated by being in groups. Better advertisement Being in a group may better advertise a warning patter. Standing head to tail helps zebras cover their behinds. Being in a group may provide a more effective learning experience for predator. The ‘many eyes’ strategy permits any one individual to spend more time foraging and less time on vigilance Goshawk has less success as flock gets larger. Group advantage is due to better vigilance. Vervet Alarm Calls Vervets make three different kinds of alarm calls, according to kind of predator. Alarm calls Birds and mammals utter calls that signal predator threat to others Predator Effect on Listeners Belding’s ground squirrel Alarm pheromones Insects and fish emit chemical signals that signal predator threat. Eagle Snake Look up and run to bushes Look down at ground Leopard Run and climb trees Alarm pheromone of honey bee Sometimes signal simply alerts individuals who then flee; sometimes signal elicits a group defense. 11 10/16/2012 Diana’s monkeys have separate alarm calls for leopards and crownedhawk eagles. African hornbills respond to the alarm calls for eagles (which are hornbill predators) but not the alarm call for eagles. Campbell’s monkey males link sounds together, making for a rich repertoire. Hok! given in response to eagle. Hok oo! given in response to almost any disturbance in the canopy, including eagles, neighboring groups of monkeys, squirrel, etc. Figure 1. Spectrographic illustrations of the different loud call types produced by male Campbell's monkeys in different contexts. Ouattara K, Lemasson A, Zuberbühler K (2009) Campbell's Monkeys Use Affixation to Alter Call Meaning. PLoS ONE 4(11): e7808. doi:10.1371/journal.pone.0007808 http://www.plosone.org/article/info:doi/10.1371/journal.pone.0007808 Audio S1 ‘‘Hok’’ calls are almost exclusively associated with crowned eagle presence. Found at: doi:10.1371/journal.pone.0007808.s001 (0.05 MB WAV) Audio S2 ‘‘Hok-oo’’ are given to a range of disturbances within the canopy, including eagles, the presence of neighbouring groups and, on a few occasions, to a flying squirrel. While producing these calls, males adopt a threat posture, combined with flashing their eyelids, sometimes combined with a short dash towards the disturbance. Found at: doi:10.1371/journal.pone.0007808.s002 (0.06 MB WAV) Audio S3 ‘‘Krak’’ calls are exclusively given after detecting a leopard. Found at: doi:10.1371/journal.pone.0007808.s003 (0.09 MB WAV) Audio S4 ‘‘Krak-oo’’ function as a general alert call and can be given to almost any disturbance. Found at: doi:10.1371/journal.pone.0007808.s004 (0.09 MB WAV) Audio S5 ‘‘Wak-oo’’ calls are given to the same events as ‘‘hokoo’’ calls (eagles, other flying animals, Diana monkey eagle alarms), but not to neighbours. Found at: doi:10.1371/journal.pone.0007808.s005 (0.14 MB WAV) Audio S6 ‘‘Boom’’ calls are given to non-predatory contexts, such as a falling branch or tree, to initiate or halt group travel, during disputes with neighbours, and to any unusual vocal excitation with the group. Found at: doi:10.1371/journal.pone.0007808.s006 (0.20 MB WAV) Composition of call sequences in different behavioral contexts. “Alarm” indicates leopard or eagle alarm calls given by sympatric Diana monkeys. Ouattara K et al. PNAS 2009;106:22026-22031 ©2009 by National Academy of Sciences 12