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研究生: 謝章壬
Hsieh, Cheng-Ren
論文名稱: 墾丁地區共域蜥蜴亞目的群聚與覓食生態
Assemblage Composition and Feeding Ecology of Sympatric Lacertilians in Kenting Area
指導教授: 李亞夫
Lee, Ya-Fu
學位類別: 碩士
Master
系所名稱: 生物科學與科技學院 - 生命科學系
Department of Life Sciences
論文出版年: 2014
畢業學年度: 102
語文別: 英文
論文頁數: 89
中文關鍵詞: 群聚生物多樣性覓食效率覓食行為覓食率覓食成功率壁虎蜥蜴共域性
外文關鍵詞: assemblage, biodiversity, foraging behavior, feeding efficiency, feeding rate, feeding success rate, geckoes, lizards, sympatric
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  • 有鱗目 (Squanmata) 的爬蟲動物廣泛分布世界各地,其中蜥蜴亞目 (Lacertilia) 包含蜥蜴和壁虎兩大類群。 在不同的棲地類型中,植披結構與環境因子 (如溫度) 的差異及其所具備的資源不盡相同,出現其中的生物群聚組成與物種間資源使用模式及覓食策略也可能有所差異。 本研究探討在墾丁地區不同棲地類型中環境因子如何影響蜥蜴亞目的群聚組成及分布,處在不同棲地類型中,蜥蜴的棲地選擇是否改變;使用不同覓食策略的物種在食性組成上的差異;獵物以及各物種對彼此覓食行為的影響。 我測試假說:關係密切的共域性相似物種在資源使用有分隔,並可能影響群聚組成。
    本研究在墾丁龜仔角試驗林,針對栽植林、破碎森林、連續森林、及草生地四類型棲地,設置了198個20 × 20公尺的樣格,從2012年1月至2013年5月,進行每月兩次為期4-5天的調查,每次調查以十分鐘內在樣格中以目視遇測法紀錄所見之蜥蜴。 我同時紀錄各類型地表所佔覆蓋比例及中層灌叢與高層樹冠層所占覆蓋百分比。 本研究共紀錄三科8種蜥蜴,包括舊大陸鬣蜥科 (Agamidae):斯文豪氏攀蜥 (Japalura swinhonis Günter, 1864);正蜥科 (Lacertidae):梭德氏草蜥 (Takydromus sauteri Van Denburgh, 1909);石龍子科 (Scincidae):麗紋石龍子 (Plestiodon elegans Boulenger, 1887)、長尾真稜蜥 (Eutropis longicaudata Hallowell, 1856)、多線真稜蜥 (Eutropis multifasciata Kuhl, 1820)、台灣滑蜥 (Scincella formosensis Van Denburgh, 1912)、股鱗蜓蜥 (Sphenomorphus incognitus Thompson, 1912)、與印度蜓蜥 (Sphenomorphus indicus Gray, 1853),及4種壁虎 (壁虎科 Gekkonidae):疣尾蝎虎 (Hemidactylus frenatus Schlegel, 1836)、無疣蝎虎 (Hemidactylus bowringii Gray, 1845)、鉛山壁虎 (Gekko hokouensis Pope, 1928)、及鱗趾虎 (Lepidodactylus lugubris Duméril éril and Bibron, 1836)。
    日行性蜥蜴的平均豐度及物種數在栽植林及破碎森林達到最高,在夜間,四種棲地類型間之壁虎平均豐度及物種數則沒有差異。 整體而言,栽植林均具備最高的蜥蜴的多樣性;另一方面,日間的破碎森林,以及夜間的栽植林則呈現較高均勻度。 除了在栽植林中股鱗蜓蜥有較高的豐度,在其他森林類型中斯文豪氏攀蜥皆有較高的豐度。 夜晚時,除了草生地的疣尾蝎虎豐度較高,其餘三種森林類型中皆是以鉛山壁虎的豐度最高。 逐步迴歸分析顯示,溫度是影響數量較多的三種蜥蜴:斯文豪氏攀蜥、長尾真稜蜥、及股鱗蜓蜥分布的共同重要因子。 不同物種有各自偏好的棲地類型,斯文豪氏攀蜥偏好有較高比例的樹的環境,長尾真稜蜥及股鱗蜓蜥雖然沒有明顯偏好的棲地類型,但皆會避免在灌叢及裸露地活動。
    在捕捉到的四種蜥蜴之食性分析中發現,印度蜓蜥及股鱗蜓蜥皆只發現4個昆蟲目,斯文豪氏攀蜥的食性包含10個目的節肢動物及蜥蜴和植物,而長尾真稜蜥的食性亦包含相同10個目的節肢動物及蜥蜴和蝸牛。 物種間型態與覓食策略的差異造成不同的食性組成。 覓食策略為坐等型的斯文豪氏攀蜥之食性組成中,以鞘翅目 (Coleoptera)、膜翅目 (Hymenoptera)、以及直翅目 (Orthoptera) 共佔66%的相對出現頻度及超過75%的體積百分比。 以主動覓食的三種石龍子科蜥蜴食性組成中,膜翅目、以及直翅目相對出現頻度較高,佔整體38%,並佔50%的體積百分比。
    本研究同時在夜晚針對墾丁地區共棲的三種最常見壁虎,疣尾蝎虎、無疣蝎虎、以及鉛山壁虎,於2012年6月至2013年9月間,在恆春熱帶植物園八個建築物的外牆定期進行行為觀察。 並以錄影的方式記錄壁虎的覓食行為, 探討壁虎間捕食相同或不同獵物時覓食行為的差異,及周圍出現個體的種類、數量、距離、與行為對於壁虎覓食之影響。 我測試假說 (1) 壁虎的覓食表現會受到獵物的種類及大小影響,及 (2) 壁虎的覓食行為會受鄰近個體的物種、數量、及距離影響。
    從晚上7點至凌晨4點,皆有三種壁虎的活動。 疣尾蝎虎數量隨時間逐漸降低,無疣蝎虎及鉛山壁虎則無明顯變化。 三種壁虎的覓食效率、覓食率、及覓食成功率沒有差異。 當三種壁虎單獨覓食相同獵物:鱗翅目 (Lepidoptera) 時,彼此不管在攻擊前的等待時間、接近獵物的速度、及攻擊時與獵物間的距離皆無明顯差異。 疣尾蝎虎共捕食了5個目的昆蟲及一種壁虎。 在捕食壁虎時,疣尾蝎虎接近獵物的等待時間高過捕食昆蟲,在捕食雙翅目 (Diptera) 時,攻擊時與獵物間的距離最短。 無疣蝎虎及鉛山壁虎皆只記錄到捕食兩個目的昆蟲,分別為膜翅目與鱗翅目;鱗翅目與鞘翅目。 無疣蝎虎在捕食鱗翅時的接近速度高於捕食膜翅目,而鉛山壁虎在捕食補不同獵物時,並未有覓食行為上的差異。 隨著獵物的體型增加,疣尾蝎虎不管是等待時間、接近速度、或是與獵物間的距離皆隨之上升。 無疣蝎虎與獵物間的距離及鉛山壁虎之等待時間也隨獵物體型增加而上升。 最近個體的物種及其與覓食者間的距離均不影響三種壁虎的行為。 疣尾蝎虎接近獵物的平均速度則隨周圍壁虎數量增加而下降。 三種壁虎的覓食率皆不受周圍個體數量影響,但疣尾蝎虎之覓食成功率隨著周圍個體數量增加而降低。
    結果顯示,在栽植林中,蜥蜴有較多機會做體溫調節,這也表現在栽植林中較高的物種豐富度及數量。 三種壁虎在單獨覓食時的行為表現相似,但各自在面對不同獵物時,行為表現受到獵物物種及大小影響。 鄰近個體的數量,造成覓食上的部分差異,且影響疣尾蝎虎的覓食成功率。有較高攻擊性的疣尾蝎虎,在食物資源的競爭中較佔優勢,但其覓食行為仍會受到其他物種的影響,甚至因此降低覓食成功率。

    Suborder Lacertilia of order Squanmata is comprised of lizards and geckos that are prevailing around the world. Different habitat types are comprised of specific environments and resources, thus result in different assemblage compositions. The study investigated (1) the effects of environmental factors on lizard assemblage compositions and distribution patterns among different habitat types, (2) habitat selection of different species in each habitat type, (3) the diet compositions with different foraging strategies of lizards, and (4) how the prey types and the interactions among species affect the foraging behaviors of geckos. Lizards are ectotherms, so that temperature-related factors, including ambient temperature, wind speed, relative humidity, and the amount of solar radiation, may influence their activity patterns directly or indirectly. I tested the hypothesis that closely-related sympatric taxa often reveal resource partitioning, which may influence assemblage compositions.
    The study site was situated in the Guililaou Exprtimental Forests, Kenting area. I set 198 grids, which were 20 m × 20 m in size in four habitat types, including plantations, fragmented woods of forest edge, intact woods with continuous canopy, and grasslands. I conducted 4-5 day field works twice a month from January 2012 to May 2013. When surveyed in grids, I used visual encounter method and recorded the lizards in 10 minutes. The grids were further classified into three vertical levels where the tree canopy coverage, shrub crown coverage, and the proportion of each ground coverage were estimated. I recorded 8 species of lizards and 4 species of geckoes. They included Swinhoe’s japaluras (Japalura swinhonis Günter, 1864, Agamidae), Sauter’s grass lizards (Takydromus sauteri Van Denburgh, 1909, Lacertidae),elegant skinks (Plestiodon elegans Boulenger, 1887), long tailed skinks (Eutropis longicaudata Hallowell, 1856), common sun skinks (Eutropis multifasciata Kuhl, 1820), formosan smooth skinks (Scincella formosensisVan Denburgh, 1912), brown forest skinks (Sphenomorphus incognitus Thompson, 1912), and Indian forest skinks (Sphenomorphus indicus Gray, 1853) of the family Scincidae, and common house geckos (Hemidactylus frenatus Schlegel, 1836), Bowring’s geckos (Hemidactylus bowringii Gray, 1845), Hekou’s geckos (Gekko hokouensis Pope, 1928), and common smooth-scaled geckos (Lepidodactylus lugubris Duméril and Bibron, 1836) of the family Gekkonidae.
    The mean abundance and richness of diurnal lizards were higher in both plantations and fragmented woods of forest edge, while geckos revealed no differences among these habitat types at night. The heterogeneity of lizard assemblages was the highest in plantations, whereas evenness peaked in fragmented woods of forest edge at daytime and plantations at night, respectively. Sphenomorphus incognitus dominated plantations, but J. swinhonis was the most abundant in other three habitat types. Among geckos, G. hokouensis was the prevailing species in all habitat types, whereas H. frenatus occurred in grasslands most frequently. Temperature was the essential factor that influenced the distributions of the three most dominant lizards, J. swinhonis, E. longicaudata, and S. incognitus. Japalura swinhonis preferred the habitat with higher tree density, on the other hand, E. longicaudata, and S. incognitus exhibited no habitat preferences and avoided shrub and exposed land.
    I totally captured four species. Both S. incognitus and S. indicus contained comprised four orders of arthropods in the fecal samples. The diet of both J. swinhonis and E. longicaudata comprised of 10 same orders of arthropods, along with some lizard remains. In addition, J. swinhonis were found ingesting plant items, and E. longicaudata consuming snails. Coleoptera, Hymenoptera, and Orthoptera accounted for 66% relative frequency and more than 75% volume of the diet composition of J. swinhonis. In the diet of family Scincidae, Hymenopterans and Orthopterans were the main food items, accounting to 38% of relative frequency and more than 50% of volume. The difference of morphology and foraging strategy between lizards cause the difference on the diet compositions.
    The study used three sympatric geckos, H. frenatus, H. bowringii, and G. hokouensis, to investigate the differences in foraging behaviors when capturing the same or different prey. I chose eight buildings in the Hengchun Tropical Botanical Garden (HTBG) to observe the behaviors of geckos. I recorded the foraging behaviors of geckos at night from June 2012 to September 2013. I tested the hypothesis that prey type and prey size will influence gecko foraging performances. I also tested the hypothesis that the foraging behaviors of geckos will be affected by species, number, and distance to neighboring geckos.
    The activity time of most geckos initiated from 19:00 to 04:00; however, the movement pattern of each species was different. The abundance of H. frenatus reached to the peak at about 19:00, but the abundance of H. bowringii and G. hokouensis were similar of each hour all night. The feeding rate, feeding success rate, and feeding efficiency were similar among these gecko species. When these three geckos captured lepidopterans, they showed similar foraging behaviors. Hemidactylus frenatus predated five insect orders and one gecko species. Once capturing geckos, the approaching latency was higher than capturing insects, while the approaching distance was the shortest in capturing dipterans. Hemidactylus bowringii hunted lepidopterans with a faster approaching speed than hunting Hymenopterans, whereas G. hokouensis showed similar foraging behaviors when capturing coleopterans and lepidopterans. The approaching latency, speed, and distance of H. frenatus all increased with increasing prey size; although only the approaching distance of H bowringii and the approaching latency of G. hokouensis will increase with increasing prey size. The kind of species of the nearest gecko, and the distance between two foraging geckos did not affect the foraging behaviors of each species. The mean approaching speed of H. frenatus decreased with the increasing number of neighboring geckos. The number of nearby individuals did not affect the feeding rate of these three geckos, but the success rate of H. frenatus decreased with the increasing number of neighboring individuals.
    The results indicated that lizards had more opportunities to do their thermoregulation in the plantations, which revealed the highest species richness and abundance. Moreover, when encountering different prey, the prey species and size affected the foraging behaviors. The number of nearby individuals will cause differences patterns in foraging behaviors, which influence the subsequent feeding success rate. Even H. frenatus had higher aggressive characteristics with more advantage to resource competition than other two species, the foraging behaviors of H. frenatus still affected by other gecko species. Therefore, the feeding success rate of H. frenatus decreased.

    Table of contents 摘要 i Abstract iv 致謝 viii Table of contents ix List of tables xi List of figures xii I. Introduction 1 Statement of the problem 1 Background 2 Animal assemblage compositions 2 Lizard activity patterns 5 Habitat selection 6 Lacertilians in Hengchun Peninsula 7 Food resource use 7 Relevant cues affecting foraging behavior 8 The effect of neighboring individuals 10 Interactions among foraging geckos 12 Hengchun Peninsula 13 Objectives 13 II. Assemblage composition, activity patterns and habitat use of sympatric lacertilians 15 Introduction 15 Materials and Methods 17 Study sites 17 Habitat sampling 18 Lizard sampling 20 Fecal sampling 20 Data analyses 22 Results 25 Lacertilia assemblages in the different habitat types 25 Environmental factors 35 Activity patterns of each species in each habitat type 38 Lizard diets 39 Discussion 43 III. Neighboring effects of conspecific and heterospecifics on foraging behaviors of geckos 51 Introduction 51 Materials and methods 53 Study sites 53 Sampling and observations 54 Data analysis 55 Results 56 Gecko census 56 Foraging behaviors of geckos 56 Effects of prey types and prey size on gecko foraging 58 Effects of neighboring individuals on gecko foraging behavior 61 Discussion 63 IV. Conclusions 69 V. Literature cited 71 Vita 89 List of tables Table Page 1-1. The relative frequency (RF) of occurrence, relative abundance (RA), relative importance (RI), and density of lizards in different habitat types 26 1-2. The relative frequency (RF) of occurrence, relative abundance (RA), relative importance (RI), and density of geckos in different habitat types. 28 1-3. Simpson’s diversity and Smith-Wilson evenness indices of Lacertilia assemblages in the different habitat types 29 1-4. Morisita’s index of similarity of the Lacertilia assemblages among the different habitat types. 30 1-5. The differences of environmental factors among each habitat type. 36 1-6. Stepwise regression analysis for environmental factors that explained the mean abundance of J. swinhonis, E. longicaudata, and S incognitus in all habitats 37 1-7. The estimated proportions of 8 ground coverages and lizard selection indices. 40 1-8. The relative frequency (RF) of occurrence, relative volume (RV), and relative importance (RI) of each prey type in the diets of families Agamidae and Scincidae. 43 2-1. The rate of feeding success (%) of geckos with different numbers of neighboring individuals 63 2-2. The rate of feeding success (%) of H. frenatus with different numbers of neighboring conspecifics 63 2-3. The rate of feeding success (%) of H. frenatus with different numbers of neighboring heterospecifics 63   List of figures Figure Page 1-1. Location maps showing Taiwan, Kenting within the Hengchun Peninsula, and the enlarged boundary of the study area 21 1-2. The rarefaction curve of Lacertilia assemblage in Kenting area 26 1-3. Changes in the relative abundance of lacertilian species in different habitat types. 29 1-4. The mean abundance of lizards at different seasons in each habitat type.. 30 1-5. The mean abundance of lizards at different periods of lizard in each habitat type.. 31 1-6. The mean abundance of J. swinhonis at different seasons in each habitat type.. 32 1-7. The mean abundance of E. longicaudata at different seasons in each habitat type. 33 1-8. The mean abundance of S. incognitus at different seasons in each habitat type 33 1-9. The species richness of lizards at different seasons in each habitat type. 34 1-10. The species richness of lizards at different periods of the day in each habitat type. 34 2-1. Activity patterns of three gecko species overnight. 57 2-2. Approaching latency of : G. hokouensis, : H. bowringii, : H. frenatus when encountering different prey.. 58 2-3. Approaching speed of : G. hokouensis, : H. bowringii, : H. frenatus when encounterung different prey.. 59 2-4. Approaching distance of : G. hokouensis, : H. bowringii, : H. frenatus when encountering different prey. 60 2-5. The relationship between the number of neighboring geckos and the mean approaching speed (cm/s) of (a) G. hokouensis, (b) H. bowringii, and (c) H. frenatus. 62

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