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研究生:陳溫柔
研究生(外文):Wen-jou Chen
論文名稱:臺灣地區澤蟹屬蟹類親緣關係暨西南部惡地區域厚圓澤蟹之適應策略研究
論文名稱(外文):The Phylogeny of Geothelphusa in Taiwan and the Adaptive Strategy of the crab Geothelphusa ancylophallus from the Badland in the Southwestern Taiwan
指導教授:徐芝敏徐芝敏引用關係鄭金華
指導教授(外文):Minna J. HsuJin-Hua Cheng
學位類別:博士
校院名稱:國立中山大學
系所名稱:生物科學系研究所
學門:生命科學學門
學類:生物學類
論文種類:學術論文
論文出版年:2007
畢業學年度:95
語文別:中文
論文頁數:117
中文關鍵詞:適應策略厚圓澤蟹臺灣地區澤蟹屬蟹類親緣關係惡地
外文關鍵詞:Geothelphusa ancylophallusadaptive strategybadlandTaiwanGeothelphusaPhylogeny
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本研究分兩部份,第一部份主要探討臺灣本島以及蘭嶼與綠島澤蟹屬Geothelphusa種間之親緣關係,第二部份則探討臺灣西南部惡地區域內優勢種厚圓澤蟹G. ancylophallu在泥岩環境的適應策略。
澤蟹屬種間之親緣關係研究,首先藉由SEM針對22種澤蟹的雄性第一生殖器(G1)的微細構造進行測量與分析比較。由11種G1形質特徵可將臺灣地區澤蟹分為東、西兩大群,平均距離係數為0.97(± 0.26, range 0.64 - 1.51, n = 22)。澤蟹種間在G1形質特徵上的主要差異,包括在末節的倒三角形狀棘的圈數、簡單型棘的數目、第一交接器全長與末節長的比值。
其次藉由mtDNA COI片段序列(658 bp)分析臺灣地區27種澤蟹。結果顯示臺灣本島澤蟹祖先種約在374萬年前開始分化。隨後由本島北部向東、西兩方向散佈(dispersal)分化,此結果與前述G1形質特徵的分析結果相符。推測澤蟹祖先種由臺灣本島西北方登陸往東邊分化的族群有一群往東北分化,形成宜蘭地區的澤蟹族群(d = 0.089)。另有一群往南分化為梨山地區(d = 0.078)、花蓮中橫的澤蟹物種。然後再繼續往南散佈分化為台東地區的物種(d = 0.078),最後分化為恆春半島、蘭嶼與綠島的澤蟹物種(d =0.043)。至於臺灣西部的族群,向南散佈至西南部時(d = 0.058),有一群分化為黃綠澤蟹與楠西澤蟹,續分化為藍灰澤蟹(d = 0.009)、霧台澤蟹(d = 0.019)與蔡氏澤蟹(0.009-0.022)。另有一群分化為厚圓澤蟹,續而往南分化為力里澤蟹、神山澤蟹、內埔澤蟹和屏東澤蟹。
同樣藉mtDNA COI片段序列(658bp)進一步分析高屏溪水域澤蟹的遺傳結構,結果顯示蔡氏澤蟹在高屏溪水系三大支流(楠梓仙溪、荖濃溪、隘寮溪)可區分為獨立的3群,族群間有顯著差異(d = 0.013 - 0.018)。其中隘寮溪群與霧台澤蟹關係相近(d = 0.012),支持率為73% - 82%,可歸為同種。另外荖濃溪群與藍灰澤蟹關係亦相近(d = 0.009),支持率為60% ,可否歸為同種尚需其他佐證。
厚圓澤蟹G. ancylophallu在泥岩惡地景觀區域內適應策略如下:1.) 生殖活動與雨量有顯著相關,集中在春、夏兩季,在6月份達最高鋒。2.) 母蟹平均育苗數為32.5隻(± 1.3, n = 9),第一期稚蟹平均殼寬為5.3 mm(± 0.13, n=15),屬於大型稚蟹,而且第一期稚蟹依附在母蟹腹夾內受保護約4天(n = 4)。3.) 厚圓澤蟹在洞旱季來臨前完成築洞,並持續使用到下次雨季。在旱季為抗旱、避敵之用,濕季為交配、孕育的場所。4.) 具防衛洞穴行為,洞主將測試蟹趕出洞外所付出平均時間為152.3秒(± 110.1)。洞主對來自地面震動響聲,約第2.3次(± 0.97),便有反應。5.) 由洞口覆蓋實驗:得知洞穴連續使用超過4個月者佔總有效洞穴67.0 %以上,洞穴使用最長時間為13個月。由線圈拉線測試:活動距離以夜間距離較長,平均約146.8 cm(± 73.75, n = 38),其中成功回到原洞者距離平均約174.9 cm(± 72.6, n = 8),故厚圓澤蟹對洞穴具有忠誠度(< 60﹪, 8/38)。
The thesis is divided into two parts, the phylogeny of the Geothelphusa spp. from Taiwan and the adaptive strategy of Geothelphusa ancylophallu from the badland in the southwestern Taiwan.
Through SEM studies, the male first gonopod (G1) microstructures of 22 species of Geothelphusa are compared. From the Numerical Analysis of 11 G1 measurements, it shows that Geothelphusa spp can be divided into the eastern and the western geographic groups in Taiwan. In addition, the main differences in G1 micro-morphology among Geothelphusa, including the rings of scales, the numbers of simple seta on the terminal segment, and the ratios of total length of G1 divided by the terminal segment length of G1.
Twenty-seven species of Geothelphusa from type locality in Taiwan were compared via the sequences of mtDNA COI (658 bp). The results indicate that the ancestors of Geothelphusa landed onto the northwestern Taiwan about 3.74 mya. After that, Geothelphusa dispersed from northern Taiwan into the eastern and the western geographic groups, this result was consisted with the previous conclusion obtained by G1 SEM studies.
The eastern geographic group was further dispersed to the northeast and evolved to the present-day Ilan Geothelphusa spp. Another subgroup distributed down south to the eastern mountains where are now Lishan area. The last subgroup reached more southern to the Hengtsuen peninsula, Lanyu, and Greenisland. The western geographic group was further dispersed down south and evolved to the present Geothelphusa olea and G. nanhsi which then evolved to G. caesia、G. wutai and G. tsayae. Another subgroup evolved to G. ancylophallus which then evolved to G. lili、G. shernshan、G. neipu and G. pingtung.
The genetic structures of Geothelphusa in Kaoping River area were studied further via the sequences of mtDNA COI (658bp). The results indicate that G. tsayae have three main subgroups according to the river they live on (Nantsishan River, Laolong River, Eyeliou River). Among them, the Eyeliou-River subgroup has the sequences divergence of 1.2% to the G. wutai, with supporting rate of 73%-82%, indicating that they may belong to the same species. In addition, Laolong-River subgroup has the sequences divergence of 0.9% to the G. caesia group, with supporting rate of 60%, indicating that they may be the same species as well but more evidences are needed to clarify the situation.
The adaptive strategy of the crabs Geothelphusa ancylophallu, distribute near the badland in the southwestern Taiwan, was studied. The strategies are as follows: 1) the breeding activity correlated to precipitation, mainly occurring in spring and summer, and the maximum in June. 2) The female crabs breed 32.5 offspring in average and the mean carapace width of the first stage crabs (C1) was 5.3mm. The C1 stayed with their mother for about 4 days. 3) G. ancylophallu finished building holes before the dry season and the holes could be used until the next wet season. The holes are used for drought-resistance and defending predators during the dry season and mating and breeding during the wet season. 4) G. ancylophallu could defense their holes. The residents spent 152.3 ± 110.1 seconds driving the invaders out of their holes and attacked them actively when near their holes. The residents’ alert to sound from the ground. 5) Through the experiment of hole covering, 67.0% of the valid holes were used more than 4 months, with the longest of 13 months. The moving distance of G. ancylophallu at night was 146.8 ± 73.8 cm (n = 38). The G. ancylophallu are fidelity to their holes, especially in dry season.
目 錄
誌謝..………………………………………………………………………………... i
摘要..………………………………………………………………………………... ii
Abstract...……………………………………………………………………………. iv
目錄..…………………………………………………………………………….…. vi
圖目錄..…………………………………………………………………………….. ix
表目錄..…………………………………………………………………………….. xi
第一部 臺灣地區澤蟹屬蟹類親緣關係之研究…...………………………….... 1
前言 ………………………………………………………………………………... 1
材料與方法 ..………………………………………………………………………. 5
一、材料來源 ..……………………………………………………………….. 5
二、採集與鑑定………………………………………………………………. 5
(一)採集方法 ..…………………………………………………. 5
(二)標本鑑定與保存……………………………………………. 6
三、掃瞄式電顯技術分析 ...…………………………………………………. 6
(一)標本處理 ..…………………………………………………. 6
(二)生殖肢SEM構造與測量 …………………………………. 7
(三)典型區別分析 ..……………………………………………. 9
(四)數值分類分析 ..……………………………………………. 9
四、粒線體DNA片段定序與分析 …………………………………………. 9
(一)Genomic DNA萃取 .………………………………………. 10
(二)聚合脢連鎖反應 ...………………………………………… 10
(三)洋菜膠電泳分析 ...………………………………………… 10
(四)PCR產物之定序反應……………………………………… 10
(五)DNA序列比對 .....………………………………………… 10
(六)分生親緣關係分析 ...……………………………………… 10
五、高屏溪水域澤蟹屬蟹類遺傳結構 ……………………………………… 11
六、臺灣地區澤蟹屬蟹類遺傳結構………………………………………….. 12
結果..………..………………………………………………………………………. 12
一、掃瞄式電顯技術分析 ..………………………………………………… 12
(一)生殖肢SEM構造與測量 .…………………………………. 12
澤蟹屬 Geothelphusa Stimpson, 1858 …………………………….. 13
黃灰澤蟹 G. albogilva Shy, Ng & Yu, 1994……………………….. 13
厚圓澤蟹G. ancylophallus Shy, Ng & Yu, 1994…………………… 14
雙色澤蟹G. bicolor Shy, Ng & Yu, 1994 ..………………………... 16
藍灰澤蟹G. caesia Shy, Ng & Yu, 1994 …………………………… 17
灰甲澤蟹G. cinerea Shy, Ng & Yu, 1994..…………………………. 18
扁足澤蟹G. dolichopodes Shy, Ng & Yu, 1994 ……………………. 19
顯齒澤蟹G. eucrinodonta Shy, Ng & Yu, 1994 ……………………. 20
寬甲澤蟹G. eurysoma Shy, Ng & Yu, 1994………………………... 21
�蛈熂A蟹G. ferruginea Shy, Ng & Yu, 1994………………………. 21
細足澤蟹G. gracilipes Shy, Ng & Yu, 1994……………………….. 22
宜蘭澤蟹 G. ilan Shy, Ng & Yu, 1994……………………………... 23
蘭嶼澤蟹 G. lanyu Shy, Ng & Yu, 1994…………………………… 24
力里澤蟹 G. lili Chen, Cheng & Shy, 2005..………………………. 25
綠島澤蟹 G. lutao Shy, Ng & Yu, 1994……………………………. 26
宮崎氏澤蟹 G. miyazakii (Miyake & Chiu, 1965)…………….. 27
高山澤蟹 G. monticola Shy, Ng & Yu, 1994………………………. 27
南澳澤蟹G. nanao Shy, Ng & Yu, 1994…………………………… 28
楠西澤蟹 G. nanhsi Shy, Ng & Yu, 1994………………………….. 29
內埔澤蟹 G. neipu Chen, Cheng & Shy, 1998…………………….. 30
黃綠澤蟹 G. olea Shy, Ng & Yu, 1994…………………………….. 31
屏東澤蟹G. pingtung Tan & Liu, 1998…………………………….. 32
神山澤蟹 G. shershan Chen, Cheng & Shy, 2005…………………. 33
達觀澤蟹 G. takuan Shy, Ng & Yu, 1994………………………….. 34
大里澤蟹 G. tali Shy, Ng & Yu, 1994……………………………… 35
太魯閣澤蟹 G. taroko Shy, Ng & Yu, 1994………………………... 36
大武澤蟹 G. tawu Shy, Ng & Yu, 1994……………………………. 36
蔡氏澤蟹 G. tsayae Shy, Ng & Yu, 1994…………………………... 37
霧台澤蟹 G. wutai Shy, Ng & Yu, 1994…………………………… 39
(二)典型區別分析………………………………………………. 40
(三)數值分類分析………………………………………………. 41
二、高屏溪水域澤蟹屬蟹類遺傳結構………………………………….….… 41
(一)澤蟹屬蟹類種類與分布……………………………………. 41
(二)粒線體片段DNA定序與分析……………………………. 42
(三)高屏溪水域與鄰近區域澤蟹物種演化路線………………. 43
(四)高屏溪水域優勢澤蟹分化時間估算………………………. 43
三、臺灣地區澤蟹屬蟹類遺傳結構………………………………………….. 44
(一)粒線體片段DNA定序與分析……………………………. 44
(二)建構親緣樹…………………………………………………. 44
(三)臺灣地區澤蟹屬蟹類物種演化路線………………………. 45
(四)臺灣地區澤蟹屬蟹類分化時間估算………………………. 45
(五)臺灣地區澤蟹屬蟹類趨同演化現象………………………. 46
討論..………..………………………………………………………………………. 47
結語..………..………………………………………………………………………. 54


第二部 臺灣西南部惡地區域厚圓澤蟹之適應策略研究……………………... 57
前言..………..………………………………………………………………………. 57
材料與方法..………………………………………………………………………... 60
一、厚圓澤蟹族群動態研究……………………………………………….… 60
(一)調查區……………………………………………….……… 60
(二)固定樣區……………………………………………..……… 61
(三)調查期間與步驟……………………………………………. 62
(四)資料分析……………………………………………………. 63
二、厚圓澤蟹棲地資源利用與防禦…………………………………..……… 64
(一)厚圓澤蟹洞穴結構與棲地資源利用……………….……… 64
(二)厚圓澤蟹棲地資源防禦…………………………………….. 65
結 果………………………..………………………………………..……………... 69
一、環境因子………………………………..………………………………… 69
(一)雨量與氣溫…...…………………..………………………… 69
(二)棲地土壤含水量與溼度……………………..…………… 69
二、個體形質相對成長探討………………………………………..………… 70
(一)成熟蟹直線線性關係..………………………..…………… 70
(二)成熟蟹非線性關係…………………………..……………… 71
(三)雌雄成熟蟹成長的差異………………………………… 71
(四)雄雌與稚蟹個體合併分析………………………………. 72
三、厚圓澤蟹族群動態與結構探討..………………………………………… 72
(一)有效洞口數推估族群結構與動態……………….………… 73
(二)捕獲個體推估族群結構與動態…………………………… 74
四、厚圓澤蟹棲地資源利用與防禦..………………………………………… 74
(一)洞穴結構與棲地資源利用………………………………… 74
(二)棲息地資源防禦……………….…………………………… 76
討論..……………………..…………………………………………………………. 78
結語..……………………..………………………………………………………… 83
參考文獻..……………………..……………………………………………………. 86
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