Optimal Predator Risk Assessment by the Sonar-Jamming Arctiine Moth Bertholdia trigona
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{"title"=>"Optimal Predator Risk Assessment by the Sonar-Jamming Arctiine Moth Bertholdia trigona", "type"=>"journal", "authors"=>[{"first_name"=>"Aaron J.", "last_name"=>"Corcoran", "scopus_author_id"=>"28067695600"}, {"first_name"=>"Ryan D.", "last_name"=>"Wagner", "scopus_author_id"=>"57198961215"}, {"first_name"=>"William E.", "last_name"=>"Conner", "scopus_author_id"=>"7102658227"}], "year"=>2013, "source"=>"PLoS ONE", "identifiers"=>{"isbn"=>"1932-6203 (Electronic)\\r1932-6203 (Linking)", "pmid"=>"23671686", "doi"=>"10.1371/journal.pone.0063609", "pui"=>"368853026", "issn"=>"19326203", "sgr"=>"84877092035", "scopus"=>"2-s2.0-84877092035"}, "id"=>"248df0d4-046d-3bf9-ae02-8c50ee2b15de", "abstract"=>"Nearly all animals face a tradeoff between seeking food and mates and avoiding predation. Optimal escape theory holds that an animal confronted with a predator should only flee when benefits of flight (increased survival) outweigh the costs (energetic costs, lost foraging time, etc.). We propose a model for prey risk assessment based on the predator's stage of attack. Risk level should increase rapidly from when the predator detects the prey to when it commits to the attack. We tested this hypothesis using a predator--the echolocating bat--whose active biosonar reveals its stage of attack. We used a prey defense--clicking used for sonar jamming by the tiger moth Bertholdia trigona--that can be readily studied in the field and laboratory and is enacted simultaneously with evasive flight. We predicted that prey employ defenses soon after being detected and targeted, and that prey defensive thresholds discriminate between legitimate predatory threats and false threats where a nearby prey is attacked. Laboratory and field experiments using playbacks of ultrasound signals and naturally behaving bats, respectively, confirmed our predictions. Moths clicked soon after bats detected and targeted them. Also, B. trigona clicking thresholds closely matched predicted optimal thresholds for discriminating legitimate and false predator threats for bats using search and approach phase echolocation--the period when bats are searching for and assessing prey. To our knowledge, this is the first quantitative study to correlate the sensory stimuli that trigger defensive behaviors with measurements of signals provided by predators during natural attacks in the field. We propose theoretical models for explaining prey risk assessment depending on the availability of cues that reveal a predator's stage of attack.", "link"=>"http://www.mendeley.com/research/optimal-predator-risk-assessment-sonarjamming-arctiine-moth-bertholdia-trigona", "reader_count"=>51, "reader_count_by_academic_status"=>{"Unspecified"=>1, "Professor > Associate Professor"=>3, "Researcher"=>11, "Student > Doctoral Student"=>2, "Student > Ph. D. Student"=>7, "Student > Postgraduate"=>1, "Student > Master"=>6, "Other"=>2, "Student > Bachelor"=>17, "Professor"=>1}, "reader_count_by_user_role"=>{"Unspecified"=>1, "Professor > Associate Professor"=>3, "Researcher"=>11, "Student > Doctoral Student"=>2, "Student > Ph. D. Student"=>7, "Student > Postgraduate"=>1, "Student > Master"=>6, "Other"=>2, "Student > Bachelor"=>17, "Professor"=>1}, "reader_count_by_subject_area"=>{"Engineering"=>1, "Unspecified"=>3, "Environmental Science"=>6, "Agricultural and Biological Sciences"=>39, "Medicine and Dentistry"=>1, "Computer Science"=>1}, "reader_count_by_subdiscipline"=>{"Engineering"=>{"Engineering"=>1}, "Medicine and Dentistry"=>{"Medicine and Dentistry"=>1}, "Agricultural and Biological Sciences"=>{"Agricultural and Biological Sciences"=>39}, "Computer Science"=>{"Computer Science"=>1}, "Unspecified"=>{"Unspecified"=>3}, "Environmental Science"=>{"Environmental Science"=>6}}, "reader_count_by_country"=>{"Netherlands"=>1, "United States"=>1, "United Kingdom"=>1, "Italy"=>1}, "group_count"=>5}

Scopus | Further Information

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Figshare

  • {"files"=>["https://ndownloader.figshare.com/files/1053360"], "description"=>"*<p>Units of B<sub>time</sub> are dB•s<sup>−1</sup> for source level and intensity at moth and degrees•s<sup>−1</sup> for approach angle. Units of B<sub>time</sub><sup>2</sup> are dB•s<sup>−2</sup> for source level and intensity at moth and degrees•s<sup>−2</sup> for approach angle.</p>†<p>The variables time and time<sup>2</sup> were added as fixed effects. Attack number was entered as a random effect.</p>", "links"=>[], "tags"=>["Anatomy and physiology", "Neurological system", "Sensory physiology", "ecology", "Behavioral ecology", "Ecophysiology", "Evolutionary ecology", "Physiological ecology", "Evolutionary biology", "Animal behavior", "Model organisms", "Animal models", "neuroscience", "Sensory perception", "Zoology", "Entomology", "Mammalogy", "intensities", "intervals", "clicking"], "article_id"=>697600, "categories"=>["Biological Sciences"], "users"=>["Aaron J. Corcoran", "Ryan D. Wagner", "William E. Conner"], "doi"=>"https://dx.doi.org/10.1371/journal.pone.0063609.t002", "stats"=>{"downloads"=>0, "page_views"=>10, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Myotis_call_intensities_and_pulse_intervals_with_respect_to_time_before_clicking_for_three_Bertholdia_trigona_moths_/697600", "title"=>"<i>Myotis</i> call intensities and pulse intervals with respect to time before clicking for three <i>Bertholdia trigona</i> moths.", "pos_in_sequence"=>0, "defined_type"=>3, "published_date"=>"2013-05-06 02:06:40"}
  • {"files"=>["https://ndownloader.figshare.com/files/1053347"], "description"=>"<p>Predation can be categorized into a series of stages (A). That presented here is modified from <a href=\"http://www.plosone.org/article/info:doi/10.1371/journal.pone.0063609#pone.0063609-Endler1\" target=\"_blank\">[13]</a>. Predator risk is hypothesized to relate to predator stage by a sigmoidal function (B), with a rapid increase occurring from when the predator detects the prey to when it commits to the attack. (C) The bat echolocation attack sequence provides a model for studying risk relating to predator stage of attack, as the phases of bat echolocation reliably indicate the predator's attack stage. See text for explanation of specific predation stages and echolocation phases.</p>", "links"=>[], "tags"=>["Anatomy and physiology", "Neurological system", "Sensory physiology", "ecology", "Behavioral ecology", "Ecophysiology", "Evolutionary ecology", "Physiological ecology", "Evolutionary biology", "Animal behavior", "Model organisms", "Animal models", "neuroscience", "Sensory perception", "Zoology", "Entomology", "Mammalogy", "relates", "predation"], "article_id"=>697592, "categories"=>["Biological Sciences"], "users"=>["Aaron J. Corcoran", "Ryan D. Wagner", "William E. Conner"], "doi"=>"https://dx.doi.org/10.1371/journal.pone.0063609.g001", "stats"=>{"downloads"=>1, "page_views"=>5, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Hypothesized_predator_risk_level_as_it_relates_to_predation_stage_/697592", "title"=>"Hypothesized predator risk level as it relates to predation stage.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2013-05-06 02:06:32"}
  • {"files"=>["https://ndownloader.figshare.com/files/1053349"], "description"=>"<p>The bat searches for prey with directional beams that are not aimed at the prey. After detection (beginning of approach phase) the bat localizes the prey and then locks its sonar beam on the target. The bat then decreases the intensity of emissions to keep echoes returning at a constant level (automatic gain control) through the late approach and buzz phases. Simulation was based on three-dimensional coordinates and bat call intensities of an attack by a <i>Myotis</i> bat. Beam shape and direction was not measured directly but was based on previous literature <a href=\"http://www.plosone.org/article/info:doi/10.1371/journal.pone.0063609#pone.0063609-Ghose1\" target=\"_blank\">[19]</a>, <a href=\"http://www.plosone.org/article/info:doi/10.1371/journal.pone.0063609#pone.0063609-Jakobsen1\" target=\"_blank\">[51]</a>. Sonar beam shapes are depicted as the estimated volume ensonified by at least 90 dB SPL.</p>", "links"=>[], "tags"=>["Anatomy and physiology", "Neurological system", "Sensory physiology", "ecology", "Behavioral ecology", "Ecophysiology", "Evolutionary ecology", "Physiological ecology", "Evolutionary biology", "Animal behavior", "Model organisms", "Animal models", "neuroscience", "Sensory perception", "Zoology", "Entomology", "Mammalogy", "simulation", "sonar", "attacking", "moth", "spectrogram", "echolocation", "two-dimensional"], "article_id"=>697593, "categories"=>["Biological Sciences"], "users"=>["Aaron J. Corcoran", "Ryan D. Wagner", "William E. Conner"], "doi"=>"https://dx.doi.org/10.1371/journal.pone.0063609.g002", "stats"=>{"downloads"=>1, "page_views"=>8, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Three_dimensional_simulation_of_the_sonar_beam_of_a_bat_attacking_a_moth_left_panel_and_a_spectrogram_of_the_bat_echolocation_sequence_with_two_dimensional_plots_of_the_bat_s_echolocation_beam_shape_and_direction_relative_to_the_target_right_panel_/697593", "title"=>"Three-dimensional simulation of the sonar beam of a bat attacking a moth (left panel), and a spectrogram of the bat echolocation sequence with two-dimensional plots of the bat's echolocation beam shape and direction relative to the target (right panel).", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2013-05-06 02:06:33"}
  • {"files"=>["https://ndownloader.figshare.com/files/1053351"], "description"=>"<p>Three infrared cameras recorded bat flight trajectories, positions of tethered and free-flying moths, and positions and directional axes of the ultrasound microphone. An ultraviolet light attracted free-flying moths and foraging bats to the observation area. In field experiment 1 a silent noctuid moth was tethered and suspended along with a miniature (3 mm diameter) microphone from a 10 m telescoping pole. The microphone's close proximity to the tethered moth allowed for estimating bat call intensities emitted in the moth's direction (10 cm from the bat's mouth) and arriving at the moth for attacks on the tethered moth (real attacks) and attacks on nearby free-flying moths (false threats). In field experiment 2, the clicking moth <i>Bertholdia trigona</i> was tethered below the microphone and bat passes that elicited a clicking response were compared to passes that did not elicit clicking.</p>", "links"=>[], "tags"=>["Anatomy and physiology", "Neurological system", "Sensory physiology", "ecology", "Behavioral ecology", "Ecophysiology", "Evolutionary ecology", "Physiological ecology", "Evolutionary biology", "Animal behavior", "Model organisms", "Animal models", "neuroscience", "Sensory perception", "Zoology", "Entomology", "Mammalogy", "schematic", "setup", "attacks", "tethered", "free-flying"], "article_id"=>697594, "categories"=>["Biological Sciences"], "users"=>["Aaron J. Corcoran", "Ryan D. Wagner", "William E. Conner"], "doi"=>"https://dx.doi.org/10.1371/journal.pone.0063609.g003", "stats"=>{"downloads"=>4, "page_views"=>8, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Three_dimensional_schematic_of_field_setup_for_recording_bat_attacks_on_tethered_and_free_flying_moths_/697594", "title"=>"Three-dimensional schematic of field setup for recording bat attacks on tethered and free-flying moths.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2013-05-06 02:06:34"}
  • {"files"=>["https://ndownloader.figshare.com/files/1053352"], "description"=>"<p>Echolocation call intensity and pulse interval impinging upon a tethered focal moth are shown for five attacks on individual noctuid moths (real threat) and nine attacks on nearby, free-flying moths (false threat; see <a href=\"http://www.plosone.org/article/info:doi/10.1371/journal.pone.0063609#pone-0063609-g002\" target=\"_blank\">figure 2</a>). The optimal threshold for discriminating real and false threats was determined using quadratic discriminant analysis with 50% (p = 0.5) and 75% (p = 0.75) likelihood of a call being assigned as a real threat. <i>Bertholdia trigona</i> clicking thresholds (mean ± s. e.) measured from eight moths suspended in a sound chamber matched predicted p = 0.75 thresholds for pulse intervals of 35–100 ms.</p>", "links"=>[], "tags"=>["Anatomy and physiology", "Neurological system", "Sensory physiology", "ecology", "Behavioral ecology", "Ecophysiology", "Evolutionary ecology", "Physiological ecology", "Evolutionary biology", "Animal behavior", "Model organisms", "Animal models", "neuroscience", "Sensory perception", "Zoology", "Entomology", "Mammalogy", "clicking", "thresholds", "echolocation", "pulse-intervals"], "article_id"=>697595, "categories"=>["Biological Sciences"], "users"=>["Aaron J. Corcoran", "Ryan D. Wagner", "William E. Conner"], "doi"=>"https://dx.doi.org/10.1371/journal.pone.0063609.g004", "stats"=>{"downloads"=>2, "page_views"=>10, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Predicted_and_measured_clicking_thresholds_for_different_bat_echolocation_pulse_intervals_in_Bertholdia_trigona_/697595", "title"=>"Predicted and measured clicking thresholds for different bat echolocation pulse-intervals in <i>Bertholdia trigona</i>.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2013-05-06 02:06:35"}
  • {"files"=>["https://ndownloader.figshare.com/files/1053354"], "description"=>"<p>A real threat is a bat attacking the target moth (A, B); a false threat is a bat attacking a nearby moth (C). Real threats are shown (A) aligned by the end of the attack, and (B) aligned by the echolocation call of maximum intensity. The top panels show the bat echolocation intensities 10 cm from the bat (emitted in the moth's direction) and at the moth's position. Middle and bottom panels show how bat pulse intervals and approach angles change over the course of an attack, respectively. The rise in bat call intensity early in attacks in A and B reflects the bat localizing the moth, followed by an immediate decrease in emission intensity, which contributes to the bat's automatic gain control (see text and <a href=\"http://www.plosone.org/article/info:doi/10.1371/journal.pone.0063609#pone-0063609-g002\" target=\"_blank\">Figure 2</a>) <a href=\"http://www.plosone.org/article/info:doi/10.1371/journal.pone.0063609#pone.0063609-Hartley1\" target=\"_blank\">[21]</a>. Bats engage gain control (time = 0 s in B) in the middle of approach phase and then soon turn towards the prey (a decrease in approach angle). False threats are characterized by decreasing call intensities at the moth's position and approach angles that are poorly correlated with time in the attack (see <a href=\"http://www.plosone.org/article/info:doi/10.1371/journal.pone.0063609#pone-0063609-t001\" target=\"_blank\">Table 1</a> for statistics).</p>", "links"=>[], "tags"=>["Anatomy and physiology", "Neurological system", "Sensory physiology", "ecology", "Behavioral ecology", "Ecophysiology", "Evolutionary ecology", "Physiological ecology", "Evolutionary biology", "Animal behavior", "Model organisms", "Animal models", "neuroscience", "Sensory perception", "Zoology", "Entomology", "Mammalogy"], "article_id"=>697596, "categories"=>["Biological Sciences"], "users"=>["Aaron J. Corcoran", "Ryan D. Wagner", "William E. Conner"], "doi"=>"https://dx.doi.org/10.1371/journal.pone.0063609.g005", "stats"=>{"downloads"=>0, "page_views"=>5, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Characterization_of_8220_real_8221_and_8220_false_8221_threats_/697596", "title"=>"Characterization of “real” and “false” threats.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2013-05-06 02:06:36"}
  • {"files"=>["https://ndownloader.figshare.com/files/1053356"], "description"=>"<p>Acoustic characteristics (bat pulse interval <i>vs</i>. bat call intensity) and bat flight characteristics (distance from moth <i>vs</i>. flight trajectory relative to bat-moth vector, or phi) were used to differentiate bat passes that elicited a clicking response and those that did not (Top two rows of graphs). Dashed lines indicate thresholds for discriminating clicking from non-clicking interactions using discriminant function analysis. PPV, positive predictive value, or the percent of clicking interactions correctly classified; NPV, negative predictive value, or the percent of non-clicking interactions classified correctly. Some interactions were included where bats were outside the calibrated volume of the cameras and bat-moth distance (BMD) was>4 m. Exact BMD and bat approach angles were not available for these interactions. Bat call intensities increased and pulse intervals decreased prior to moth clicking (bottom two rows of graphs; see <a href=\"http://www.plosone.org/article/info:doi/10.1371/journal.pone.0063609#pone-0063609-t002\" target=\"_blank\">Table 2</a> for statistics).</p>", "links"=>[], "tags"=>["Anatomy and physiology", "Neurological system", "Sensory physiology", "ecology", "Behavioral ecology", "Ecophysiology", "Evolutionary ecology", "Physiological ecology", "Evolutionary biology", "Animal behavior", "Model organisms", "Animal models", "neuroscience", "Sensory perception", "Zoology", "Entomology", "Mammalogy", "passes", "did", "elicit", "clicking", "tethered"], "article_id"=>697597, "categories"=>["Biological Sciences"], "users"=>["Aaron J. Corcoran", "Ryan D. Wagner", "William E. Conner"], "doi"=>"https://dx.doi.org/10.1371/journal.pone.0063609.g006", "stats"=>{"downloads"=>0, "page_views"=>3, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Discrimination_of_bat_passes_that_did_or_did_not_elicit_clicking_by_tethered_B_trigona_in_the_field_/697597", "title"=>"Discrimination of bat passes that did or did not elicit clicking by tethered <i>B.</i><i>trigona</i> in the field.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2013-05-06 02:06:37"}
  • {"files"=>["https://ndownloader.figshare.com/files/1053357"], "description"=>"<p>Prey cost of fleeing C is dependent on predator distance and prey characteristics (C-low vs. C-high). In the classic model (panel A), benefit of fleeing B is equal to predator risk and increases as the predator gets closer (modified from <a href=\"http://www.plosone.org/article/info:doi/10.1371/journal.pone.0063609#pone.0063609-Ydenberg1\" target=\"_blank\">[1]</a>). The optimal escape distance D* occurs at the intersection of B and C. We propose that this model holds for prey that lack information about predator attack stage. For prey with information about predator attack stage (panel C), B is predicted to be a sigmoidal function with an increase in risk/benefit of fleeing when the predator detects a target and commits to an attack (see <a href=\"http://www.plosone.org/article/info:doi/10.1371/journal.pone.0063609#pone-0063609-g001\" target=\"_blank\">Figure 1</a>). Partial information on predator stage has a risk function that is a combination of predator distance and attack stage (panel B). Increased availability of information on predator stage diminishes the effect of prey cost level (C-high vs. C-low) on optimal escape distance D*.</p>", "links"=>[], "tags"=>["Anatomy and physiology", "Neurological system", "Sensory physiology", "ecology", "Behavioral ecology", "Ecophysiology", "Evolutionary ecology", "Physiological ecology", "Evolutionary biology", "Animal behavior", "Model organisms", "Animal models", "neuroscience", "Sensory perception", "Zoology", "Entomology", "Mammalogy", "varying", "prey"], "article_id"=>697598, "categories"=>["Biological Sciences"], "users"=>["Aaron J. Corcoran", "Ryan D. Wagner", "William E. Conner"], "doi"=>"https://dx.doi.org/10.1371/journal.pone.0063609.g007", "stats"=>{"downloads"=>0, "page_views"=>3, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Cost_benefit_models_of_escape_behavior_with_varying_prey_knowledge_of_predator_attack_stage_/697598", "title"=>"Cost-benefit models of escape behavior with varying prey knowledge of predator attack stage.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2013-05-06 02:06:38"}
  • {"files"=>["https://ndownloader.figshare.com/files/1053359"], "description"=>"*<p>Units of B<sub>time</sub> are dB•s<sup>−1</sup> for source level and intensity at moth and degrees•s<sup>−1</sup> for approach angle. Units of B<sub>time</sub><sup>2</sup> are dB•s<sup>−2</sup> for source level and intensity at moth and degrees•s<sup>−2</sup> for approach angle.</p>†<p>The variables time and time<sup>2</sup> were added as fixed effects. Attack number was entered as a random effect.</p>", "links"=>[], "tags"=>["Anatomy and physiology", "Neurological system", "Sensory physiology", "ecology", "Behavioral ecology", "Ecophysiology", "Evolutionary ecology", "Physiological ecology", "Evolutionary biology", "Animal behavior", "Model organisms", "Animal models", "neuroscience", "Sensory perception", "Zoology", "Entomology", "Mammalogy", "intensities", "angles"], "article_id"=>697599, "categories"=>["Biological Sciences"], "users"=>["Aaron J. Corcoran", "Ryan D. Wagner", "William E. Conner"], "doi"=>"https://dx.doi.org/10.1371/journal.pone.0063609.t001", "stats"=>{"downloads"=>0, "page_views"=>15, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Myotis_call_intensities_and_approach_angles_with_respect_to_time_for_real_and_false_attacks_/697599", "title"=>"<i>Myotis</i> call intensities and approach angles with respect to time for real and false attacks.", "pos_in_sequence"=>0, "defined_type"=>3, "published_date"=>"2013-05-06 02:06:39"}

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Relative Metric

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