研究生: |
陳震邑 Chen, Zhen-Yi |
---|---|
論文名稱: |
雄性二型雞冠細身赤鍬形蟲之異速生長與打鬥評估策略 The allometry and fighting assessment strategy of male dimorphic Cyclommatus mniszechi |
指導教授: |
林仲平
Lin, Chung-Ping |
學位類別: |
碩士 Master |
系所名稱: |
生命科學系 Department of Life Science |
論文出版年: | 2020 |
畢業學年度: | 108 |
語文別: | 英文 |
論文頁數: | 56 |
中文關鍵詞: | 體型 、雄性競爭 、資源佔有潛力 、行為序列分析 、性擇 、武器 |
英文關鍵詞: | Body size, male-male competition, resource holding potential, sequential analysis, sexual selection, weapon |
DOI URL: | http://doi.org/10.6345/NTNU202000271 |
論文種類: | 學術論文 |
相關次數: | 點閱:221 下載:17 |
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雄性鍬形蟲具有誇張的大顎作為打鬥時獲取食物與繁殖資源的武器,大顎武器的形狀與大小可能會影響打鬥的行為與勝負。本研究檢測雞冠細身赤鍬形蟲(Cyclommatus mniszechi)的異速生長關係、打鬥行為序列和打鬥評估策略。首先,大顎與體型之間的異速生長關係的判定可以用來作為決定不同型雄蟲的基礎,再者,藉由行為序列分析來描述依大顎大小隨機配對與對等配對的雄蟲間打鬥的行為序列,最後,從打鬥時間(和激烈程度)與資源佔有潛力(RHP, resource holding potential)的相關性來檢驗不同評估策略(自我、累加和相互評估策略)的預期。異速生長的分析結果顯示大顎長與體長之間存在非線性正異速生長關係,且在體長(翅鞘長)為15.03毫米時為異速生長係數改變的轉折點,可以此將鍬形蟲個體分為大型與小型雄蟲。從打鬥行為序列中可以定義出九種行為單元,包含「觸碰」、「防衛姿勢」、「抬身對峙」、「攻擊」、「推擊」、「纏鬥」、「抓取1」、「抓取2」(抓取於對手胸節或腹節)以及「撤退」。大小型雄蟲具有不同的打鬥行為序列,小型雄蟲傾向持續待在相同打鬥階段(較多在同一個打鬥階段內的行為轉換),相對於大型雄蟲則較容易進入纏鬥階段。大顎長對於打鬥結果有決定性的影響,可以做為代表雞冠細身赤鍬形蟲的資源佔有潛力相關的可靠特徵。在隨機配對打鬥中,打鬥時間與勝者及敗者的大顎長有顯著正相關,顯示自我評估可能為此種鍬形蟲所採用的打鬥評估策略。在對等配對打鬥中,打鬥時間與平均大顎長有顯著正相關,進一步支持鍬形蟲採用自我評估策略。然而,雄蟲在打鬥中出現對手之間的對等行為,呈現多次但很少造成傷害的身體接觸,與單方向的行為階段進程(由低度至高度激烈且鮮少逆向的打鬥階段)。因此,雞冠細身赤鍬形蟲的打鬥可能不僅使用自我評估策略,累加與相互評估策略可能也在其打鬥決策中扮演重要角色。
Male stag beetles (Coleoptera: Lucanidae) possessed exaggerated mandibles as weapons used in fighting contests for access to food and reproduction. The shapes and sizes of these mandibular weapons could influence the fighting behaviours and outcomes of the contests. This study examined the allometry, fighting behaviour and assessment strategy of a stag beetle, Cyclommatus mniszechi. Firstly, the allometric relationships between mandible and body sizes were identified to determine whether the males could be grouped into different morphs based on the allometries. Secondly, the behavioural sequences of male-male fights were characterized using sequential analyses of randomly and size-matched contests. Finally, the correlational predictions between contest duration (and aggressiveness) and RHP (resource holding potential) were examined to test alternative assessment strategies (self-, cumulative and mutual assessment). Allometric analyses show a non-linear positive allometry between mandible and body size in C. mnizechi males, and that they consist of dimorphic males defined quantitatively as the majors and minors by body sizes at the switch point of elytra length of 15.03 mm. Nine behavioural elements were identified from the contests, including ‘touch’, ‘defensive posture’, ‘body raising’ ‘attack’, ‘push’, ‘tussle’, ‘clamp1’(head), ‘clamp2’ (thorax or abdomen) and ‘retreat’. The major and minor males have different fighting behavioural sequences, where the minor males tend to stay within phases (more behavioural transitions within phases) of the contests and more likely to tussle than the major males. Mandible size is the main determinant of the outcomes of the contests and can be used as a reliable proxy for RHP in C. mnizechi. In randomly matched contests, strong positive relations between contest duration and winner’s and loser’s mandible sizes indicate that self-assessment determines strategic decisions in C. mniszechi. In size-matched contests, a positive relation between contest duration and mandible sizes further support the self-assessment strategy. However, males showed behavioural matching in contests, many physical contacts with rare injuries and unidirectional behavioural progressions in phases (from low towards high aggression with rare de-escalation). Therefore, the fighting contests of C. mniszechi may not settle entirely on the basis of pure self-assessment, and that cumulative or mutual assessment may also play an important role in contest decisions.
Andersson MB (1994) Sexual selection. Princeton University Press,
Arnott G, Elwood RW (2009) Assessment of fighting ability in animal contests. Anim Behav 77:991-1004 https://doi.org/10.1016/j.anbehav.2009.02.010
Bakeman R, Robinson BF, Quera V (1996) Testing sequential association: Estimating exact p values using sampled permutations. Psychol Methods 1:4-15 https://doi.org/10.1037/1082-989x.1.1.4
Bland JM, Altman DG (1995) Multiple significance tests: the Bonferroni method. BMJ 310:170 https://doi.org/10.1136/bmj.310.6973.170
Bonduriansky R (2007) Sexual selection and allometry: a critical reappraisal of the evidence and ideas. Evolution 61:838-849 https://doi.org/10.1111/j.1558-5646.2007.00081.x
Bonduriansky R, Day T (2003) The evolution of static allometry in sexually selected traits. Evolution 57:2450-2458 https://doi.org/10.1111/j.0014-3820.2003.tb01490.x
Briffa M, Elwood RW (2009) Difficulties remain in distinguishing between mutual and self-assessment in animal contests. Anim Behav 77:759-762 https://doi.org/10.1016/j.anbehav.2008.11.010
Chapin KJ, Peixoto PEC, Briffa M (2019) Further mismeasures of animal contests: a new framework for assessment strategies. Behav Ecol https://doi.org/10.1093/beheco/arz081
Clark JT (1977) Aspects of variation in the stag beetle Lucanus cervus (L.) (Coleoptera: Lucanidae). Syst Entomol 2:9-16 https://doi.org/10.1111/j.1365-3113.1977.tb00350.x
Csardi G, Nepusz T (2006) The igraph software package for complex network research. InterJournal, Complex Systems 1695:1-9
Darwin C (1871) The descent of man, and selection in relation to sex. London: Murray 415
Dennenmoser S, Christy JH (2013) The design of a beautiful weapon: compensation for opposing sexual selection on a trait with two functions. Evolution 67:1181-1188 https://doi.org/10.1111/evo.12018
Eberhard WG, Gutierrez EE (1991) Male Dimorphisms in Beetles and Earwigs and the Question of Developmental Constraints. Evolution 45:18-28 https://doi.org/10.1111/j.1558-5646.1991.tb05262.x
Eberhard WG, Rodríguez RL, Huber BA, Speck B, Miller H, Buzatto BA, Machado G (2018) Sexual Selection and Static Allometry: The Importance of Function. The Quarterly Review of Biology 93:207-250 https://doi.org/10.1086/699410
Elias DO, Kasumovic MM, Punzalan D, Andrade MC, Mason AC (2008) Assessment during aggressive contests between male jumping spiders. Anim Behav 76:901-910 https://doi.org/10.1016/j.anbehav.2008.01.032
Elwood RW, Arnott G (2012) Understanding how animals fight with Lloyd Morgan's canon. Anim Behav 84:1095-1102 https://doi.org/10.1016/j.anbehav.2012.08.035
Emlen DJ (1997) Alternative reproductive tactics and male-dimorphism in the horned beetle Onthophagus acuminatus (Coleoptera: Scarabaeidae). Behav Ecol Sociobiol 41:335-341 https://doi.org/10.1007/s002650050393
Emlen DJ (2001) Costs and the diversification of exaggerated animal structures. Science 291:1534-1536 https://doi.org/10.1126/science.1056607
Emlen DJ (2008a) The Evolution of Animal Weapons. Annu Rev Ecol Evol S 39:387-413 https://doi.org/10.1146/annurev.ecolsys.39.110707.173502
Emlen DJ (2008b) The Evolution of Animal Weapons. Annual Review of Ecology, Evolution, and Systematics 39:387-413 https://doi.org/10.1146/annurev.ecolsys.39.110707.173502
Emlen DJ, Nijhout HF (2000) The development and evolution of exaggerated morphologies in insects. Annu Rev Entomol 45:661-708 https://doi.org/10.1146/annurev.ento.45.1.661
Enquist M, Leimar O, Ljungberg T, Mallner Y, Segerdahl N (1990) A test of the sequential assessment game: fighting in the cichlid fish Nannacara anomala. Anim Behav 40:1-14 https://doi.org/10.1016/s0003-3472(05)80660-8
Friard O, Gamba M, Fitzjohn R (2016) BORIS: a free, versatile open-source event-logging software for video/audio coding and live observations. Methods Ecol Evol 7:1325-1330 https://doi.org/10.1111/2041-210x.12584
Goyens J, Dirckx J, Aerts P (2015a) Mechanoreceptor distribution in stag beetle jaws corresponds to the material stress in fights. Arthropod Struct Dev 44:201-208 https://doi.org/10.1016/j.asd.2015.03.003
Goyens J, Dirckx J, Aerts P (2015b) Stag Beetle Battle Behavior and its Associated Anatomical Adaptations. J Insect Behav 28:227-244 https://doi.org/10.1007/s10905-015-9495-3
Goyens J, Dirckx J, Aerts P (2016) Jaw morphology and fighting forces in stag beetles. J Exp Biol 219:2955-2961 https://doi.org/10.1242/jeb.141614
Goyens J, Dirckx J, Aerts P, Davidowitz G (2015c) Costly sexual dimorphism in Cyclommatus metallifer stag beetles. Func Ecol 29:35-43 https://doi.org/10.1111/1365-2435.12294
Goyens J, Dirckx J, Dierick M, Van Hoorebeke L, Aerts P (2014) Biomechanical determinants of bite force dimorphism in Cyclommatus metallifer stag beetles. J Exp Biol 217:1065-1071 https://doi.org/10.1242/jeb.091744
Green PA, Patek SN (2018) Mutual assessment during ritualized fighting in mantis shrimp (Stomatopoda). Proc Biol Sci 285 https://doi.org/10.1098/rspb.2017.2542
Hongo Y (2007) Evolution of male dimorphic allometry in a population of the Japanese horned beetle Trypoxylus dichotomus septentrionalis. Behav Ecol Sociobiol 62:245-253 https://doi.org/10.1007/s00265-007-0459-2
Hsu Y, Lee S-P, Chen M-H, Yang S-Y, Cheng K-C (2008) Switching assessment strategy during a contest: fighting in killifish Kryptolebias marmoratus. Anim Behav 75:1641-1649 https://doi.org/10.1016/j.anbehav.2007.10.017
Huxley JS 46. The relative size of antlers in deer. In: Proc Zool Soc Lond, 1931. vol 3. Wiley Online Library, pp 819-864. https://doi.org/10.1111/j.1096-3642.1931.tb01047.x
Inoue A, Hasegawa E (2012) Effect of morph types, body size and prior residence on food-site holding by males of the male-dimorphic stag beetle Prosopocoilus inclinatus (Coleoptera: Lucanidae). J Ethol 31:55-60 https://doi.org/10.1007/s10164-012-0350-0
Jennings DJ, Gammell MP, Carlin CoM, Hayden TJ (2004) Effect of body weight, antler length, resource value and experience on fight duration and intensity in fallow deer. Anim Behav 68:213-221 https://doi.org/10.1016/j.anbehav.2003.11.005
Johnson HE, Bleich VC, Krausman PR, Koprowski JL (2006) Effects of antler breakage on mating behavior in male tule elk (Cervus elaphus nannodes). Eur J Wildl Res 53:9-15 https://doi.org/10.1007/s10344-006-0060-4
Kawano K (2000) Genera and Allometry in the Stag Beetle Family Lucanidae, Coleoptera. Ann Entomol Soc Am 93:198-207 https://doi.org/10.1603/0013-8746(2000)093[0198:Gaaits]2.0.Co;2
Knell RJ (2009) On the analysis of non-linear allometries. Ecol Entomol 34:1-11 https://doi.org/10.1111/j.1365-2311.2008.01022.x
Knell RJ, Pomfret JC, Tomkins JL (2004) The limits of elaboration: curved allometries reveal the constraints on mandible size in stag beetles. Proc Biol Sci 271:523-528 https://doi.org/10.1098/rspb.2003.2641
Kojima W (2014) Mechanism of synchronous metamorphosis: larvae of a rhinoceros beetle alter the timing of pupation depending on maturity of their neighbours. Behav Ecol Sociobiol 69:415-424 https://doi.org/10.1007/s00265-014-1854-0
Kojima W, Ishikawa Y, Takanashi T (2014) Chemically mediated group formation in soil-dwelling larvae and pupae of the beetle Trypoxylus dichotomus. Naturwissenschaften 101:687-695 https://doi.org/10.1007/s00114-014-1199-6
Kojima W, Lin C-P (2017) It takes two to tango: functional roles, sexual selection and allometry of multiple male weapons in the flower beetle Dicronocephalus wallichii bourgoini. Biol J Linn Soc 121:514-529 https://doi.org/10.1093/biolinnean/blx018
Kotiaho JS (2001) The discrimination of alternative male morphologies. Behav Ecol 12:553-557 https://doi.org/10.1093/beheco/12.5.553
Kuan CY (2011) To win"big": determining factors and sequential analyses in male-male combats of a stag beetle, Cyclommatus mniszechi (Coleoptera: Lucanidae).49
Lagarde F, Corbin J, Goujon C, Poisbleau M (2005) Polymorphisme et performances au combat chez les mâles de Lucane cerf-volant (Lucanus cervus). Revue d'écologie 60:127-137
Lane SM, Briffa M (2017) The price of attack: rethinking damage costs in animal contests. Anim Behav 126:23-29 https://doi.org/10.1016/j.anbehav.2017.01.015
Li HY (2004) Taiwanese stag beetles., vol 1st. Kissnature Publisher, Taipei, Taiwan
Lin C-P, Huang J-P, Lee Y-H, Chen M-Y (2009) Phylogenetic position of a threatened stag beetle, Lucanus datunensis (Coleoptera: Lucanidae) in Taiwan and implications for conservation. Conserv Genet 12:337-341 https://doi.org/10.1007/s10592-009-9996-8
McCullough EL, Ledger KJ, O'Brien DM, Emlen DJ (2015) Variation in the allometry of exaggerated rhinoceros beetle horns. Anim Behav 109:133-140 https://doi.org/10.1016/j.anbehav.2015.08.013
Mills MR, Nemri RS, Carlson EA, Wilde W, Gotoh H, Lavine LC, Swanson BO (2016) Functional mechanics of beetle mandibles: Honest signaling in a sexually selected system. J Exp Zool A Ecol Genet Physiol 325:3-12 https://doi.org/10.1002/jez.1961
Mizunuma T, Nagai S (1994) The Lucanid beetles of the world. 1994. Mushisha Tokyo
Moczek AP, Emlen DJ (2000) Male horn dimorphism in the scarab beetle, Onthophagus taurus: do alternative reproductive tactics favour alternative phenotypes? Anim Behav 59:459-466 https://doi.org/10.1006/anbe.1999.1342
Nijhout HF, Wheeler DE (1996) Growth Models of Complex Allometries in Holometabolous Insects. Am Nat 148:40-56 https://doi.org/10.1086/285910
O'Brien DM et al. (2019) Muscle mass drives cost in sexually selected arthropod weapons. Proc Biol Sci 286:20191063 https://doi.org/10.1098/rspb.2019.1063
Okamoto K, Hongo Y (2013) Interspecific contests between males of two Japanese stag beetle species, Lucanus maculifemoratus and Prosopocoilus inclinatus: what overcomes a body size disadvantage? Behaviour 150:39-59 https://doi.org/10.1163/1568539x-00003036
Palaoro AV, Briffa M (2017) Weaponry and defenses in fighting animals: how allometry can alter predictions from contest theory. Behav Ecol 28:328-336 https://doi.org/10.1093/beheco/arw163
Payne RJH (1998) Gradually escalating fights and displays: the cumulative assessment model. Anim Behav 56:651-662 https://doi.org/10.1006/anbe.1998.0835
Payne RJH, Pagel M (1997) Why do animals repeat displays? Anim Behav 54:109-119 https://doi.org/10.1006/anbe.1996.0391
Pinto NS, Palaoro AV, Peixoto PEC (2019) All by myself? Meta-analysis of animal contests shows stronger support for self than for mutual assessment models. Biol Rev Camb Philos Soc 94:1430-1442 https://doi.org/10.1111/brv.12509
Prenter J, Elwood RW, Taylor PW (2006) Self-assessment by males during energetically costly contests over precopula females in amphipods. Animal Behaviour 72:861-868 https://doi.org/10.1016/j.anbehav.2006.01.023
Reichert MS, Quinn JL (2017) Cognition in Contests: Mechanisms, Ecology, and Evolution. Trends Ecol Evol 32:773-785 https://doi.org/10.1016/j.tree.2017.07.003
Rink M, Sinsch U (2007) Radio-telemetric monitoring of dispersing stag beetles: implications for conservation. J Zoo 272:235-243 https://doi.org/10.1111/j.1469-7998.2006.00282.x
Romiti F, Tini M, Redolfi De Zan L, Chiari S, Zauli A, Carpaneto GM (2015) Exaggerated allometric structures in relation to demographic and ecological parameters in Lucanus cervus (Coleoptera: Lucanidae). J Morphol 276:1193-1204 https://doi.org/10.1002/jmor.20411
Rowland JM, Emlen DJ (2009) Two thresholds, three male forms result in facultative male trimorphism in beetles. Science 323:773-776 https://doi.org/10.1126/science.1167345
Shapiro SS, Wilk MB (1965) An Analysis of Variance Test for Normality (Complete Samples). Biometrika 52:591-611 https://doi.org/10.2307/2333709
Shiokawa T, Iwahashi O (2000) Mandible dimorphism in males of a stag beetle, Prosopocoilus dissimilis okinawanus (Coleoptera: Lucanidae). Applied Entomology and Zoology 35:487-494 https://doi.org/10.1303/aez.2000.487
Simmons LW, Emlen DJ (2006) Evolutionary trade-off between weapons and testes. Proc Natl Acad Sci U S A 103:16346-16351 https://doi.org/10.1073/pnas.0603474103
Smith ABT (2006) A Review of the Family-group Names for the Superfamily Scarabaeoidea (Coleoptera) with Corrections to Nomenclature and a Current Classification. Coleopt Bull 60:144-204 https://doi.org/10.1649/0010-065x(2006)60[144:Arotfn]2.0.Co;2
Songvorawit N, Butcher BA, Chaisuekul C (2017) Different allometric intercepts in major Aegus chelifer chelifer stag beetle males from urban and forest habitats. J Asia Pac Enotmol 20:835-839 https://doi.org/10.1016/j.aspen.2017.05.011
Songvorawit N, Butcher BA, Chaisuekul C (2018) Resource holding potential and the outcome of aggressive Interactions between paired male Aegus chelifer chelifer (Coleoptera: Lucanidae) stag beetles. J Asia Pac Enotmol 31:347-360 https://doi.org/10.1007/s10905-018-9683-z
Sugiura S, Yamaura Y, Makihara H (2007) Sexual and male horn dimorphism in Copris ochus (Coleoptera: Scarabaeidae). Zoolog Sci 24:1082-1085 https://doi.org/10.2108/zsj.24.1082
Taylor PW, Elwood RW (2003) The mismeasure of animal contests. Anim Behav 65:1195-1202 https://doi.org/10.1006/anbe.2003.2169
Tomkins JL, Kotiaho JS, LeBas NR (2005) Matters of scale: positive allometry and the evolution of male dimorphisms. Am Nat 165:389-402 https://doi.org/10.1086/427732
Vieira MC, Peixoto PEC, Patek S (2013) Winners and losers: a meta‐analysis of functional determinants of fighting ability in arthropod contests. Func Ecol 27:305-313 https://doi.org/10.1111/1365-2435.12051
Voje KL (2016) Scaling of Morphological Characters across Trait Type, Sex, and Environment. Am Nat 187:89-98 https://doi.org/10.1086/684159