基於知識共享和自己的研究自己分享之理念，未來將會把我個人發表的相關研究，以中英對照的方式呈現在部落格上，並搭配簡短的說明，試圖讓非科學專業的讀者也能夠理解。這算是對我自己的一個交代，也算是對漫長的學習研究階段作個摘要。
[關於此研究]
此研究以資料文章（data paper）的形式發表於線上公開取閱（open access）的期刊Zookeys。呼應全球生物多樣性資訊機構（Global Biodiversity Information Facility，縮寫GBIF）提倡，將埋藏於研究室的珍貴調查資料發表，使得生物多樣性資料得以永存且可被引用。
[本文開始] 原始文獻連結在此
台灣的龜山島的軟體動物相 Molluscan fauna of Gueishan Island, Taiwan
作者：黃致維Chih-Wei Huang 1,2、熊大維Ta-Wei Hsiung 2、林思民Si-Min Lin 1、巫文隆 Wen-Lung Wu 2
1 國立臺灣師範大學生命科學系
Department of Life Science, National Taiwan Normal University, No. 88, Sec. 4, Tingzhou Rd., Wenshan Dist., 11677, Taipei, TAIWAN, R.O.C.
2 中央研究院生物多樣性研究中心
Biodiversity Research Center, Academia Sinica, No. 128 Academia Road Sec. 2, Nankang Dist., 11529, Taipei, TAIWAN, R.O.C.
通訊作者 Corresponding author：巫文隆 Wen-Lung Wu (malacolg@gate.sinica.edu.tw)
學術編輯 Academic editor：V. Chavan
收稿日期：2012年10月26日 | 接受日期：2013年1月14日 | 發表日期：2013年1月24日
Received 26 October 2012 | Accepted 14 January 2013 | Published 24 January 2013
摘要 Abstract
此資料彙整龜山島（台灣唯一的活火山島嶼）軟體動物調查的歷年研究文獻和2011-2012年間的田野調查，提供龜山島目前已知的軟體動物物種分布與名錄。文獻回顧包括發表於1934-2003年間的七篇研究，累積紀錄37科61屬112種軟體動物。根據我們的田野調查，共發現23科28屬34種軟體動物，其中14種是龜山島新記錄：大駝石鱉Liolophura japonica、雜斑蓮花青螺 Lottia luchuana、黑肋蜑螺Nerita costata、蘭富蜑螺Nerita rumphii、烏來芝麻蝸牛Diplommatina suganikeiensis、波紋玉黍螺Littoraria undulata、台灣山椒蝸牛Solenomphala taiwanensis、山椒蝸牛未知種Assiminea sp.、花松螺Siphonaria laciniosa、盤蜷Laevapex nipponica、八丈島罌粟蝸牛Carychium hachijoensis、台灣椎實蝸牛Succinea erythrophana、鬼牙細煙管蝸牛Zaptyx crassilamellata和細錐蝸牛Allopeas pyrgula。龜山島的軟體動物總計有45科71屬126種。以上資料已透過GBIF發布，並彙整至台灣貝類資料庫。
This dataset records the occurrence and inventory of molluscan fauna on Gueishan Island, the only active volcanic island in Taiwan, based on the literature survey and field investigation conducted between 2011 and 2012. The literature review involved seven studies published from 1934 to 2003, which collectively reported 112 species from 61 genera and 37 families of Mollusca on Gueishan Island. Through our field investigation, we identified 34 species from 28 genera and 23 families. Fourteen of these species were new records on Gueishan Island: Liolophura japonica, Lottia luchuana, Nerita costata, Nerita rumphii, Diplommatina suganikeiensis, Littoraria undulata, Solenomphala taiwanensis, Assiminea sp., Siphonaria laciniosa, Laevapex nipponica, Carychium hachijoensis, Succinea erythrophana, Zaptyx crassilamellata, and Allopeas pyrgula. In Total, there are 126 species from 71 genera and 45 families of Mollusca on Gueishan Island. These data have been published through GBIF [http://taibif.org.tw/ipt/resource.do?r=gueishan_island] and integrated into the Taiwan Malacofauna Database (http://shell.sinica.edu.tw/).
關鍵字 Keywords
軟體動物 Mollusca、腹足綱 Gastropoda、雙殼綱 Bivalvia、頭足綱 Cephalopoda、多板綱 Polyplacophora、台灣 Taiwan、龜山島 Gueishan Island
計畫詳情 Project details
計畫名稱：龜山島軟體動物相調查
Project title: Investigation of molluscan fauna of Gueishan Island, Taiwan.
人員：黃致維（鑑定、資料蒐集、資料管理、資料發布）、熊大維（鑑定、資料蒐集、資料管理）、李彥錚（鑑定）、林思民（計畫指導）、巫文隆（計畫指導、資料管理）
Personnel: Chih-Wei Huang (collection identifier, data collector, data manager, data publisher), Ta-Wei Hsiung (collection identifier, data collector, data manager), Yen-Chen Lee (collection identifier), Si-Min Lin (Project Director), Wen-Lung Wu (Project Director, data manager).
經費來源：中央研究院、行政院國家科技委員會、行政院農委會林務局
Funding: Academia Sinica; National Science Council, Executive Yuan, R.O.C.(Taiwan); Forest Bureau, Council of Agriculture, Executive Yuan, R.O.C.(Taiwan).
研究區域描述：龜山島位於距離台灣本島約10公里處。龜山島是由165萬年前的火山活動所形成，經歷多次火山噴發，最近一次噴發是在2萬年前（Juang et al. 2011），龜山島被認為是台灣附近唯一的活火山島。龜山島的陸域面積約2.85平方公里，最高海拔為398公尺。島上有兩座湖泊，淡鹹水的龜首湖（Head Lake）和淡水的龜尾湖（Tail Lake）（圖一）。大約在19世紀中期，島上開始有人類拓殖而形成小聚落，直到西元1977年，因軍事需要而將所有居民搬遷至台灣本島。直到西元2000年，龜山島由東北角暨宜蘭海岸國家風景區管理處管理，島上的動物相才開始有系統化的調查，並開放觀光遊客登島。
Study area descriptions/descriptor: Gueishan Island is located about 10 km from Taiwan. The island was formed via volcanic activity about 1.65 Ma ago and experienced multiple volcanic eruption events until 20 ka ago (Juang et al. 2011). It is considered the only active volcanic island near Taiwan. The land area of the island is about 2.85 km2, and the highest peak of the island is 398 meters above sea level. There are two lakes on the island, one of which consist of brackish water (Head Lake) and the other of freshwater (Tail Lake) (Figure 1). Humans colonized Gueishan Island in mid-19th century, by forming a small village. Later in 1977, all residents were moved back to Taiwan due to military requirements for the island. The fauna of this island were not investigated systematically until 2000, when the island came under the management of the Northeast and Yilan Coast National Scenic Area Administration, Tourism Bureau, MOTC and was open to tourists.
研究描述：棲息於島嶼上物種，因為族群數量小且資源有限，容易面臨滅絕。島嶼的物種數則是遷入物種和滅絕物種的動態平衡結果。火山島的生物相會因為火山噴發而全數滅絕後，自鄰近地區擴散拓殖而來，因而成為島嶼生物地理學的有趣研究案例。龜山島上的生物可能經歷過數次的火山噴發滅絕後，於冰河時期海平面下降時，再度自台灣本島擴散拓殖而來。人類活動可能在有意或無意間，提供軟體動物拓殖的機會。以往的龜山島軟體動物調查，並未針對陸棲軟體動物進行廣泛的調查。我們搜尋各種文獻報告資料庫，以彙整前人於龜山島地區的軟體動物調查結果，除此之外，也在2011-2012年間，進行龜山島的海生、淡水和陸生軟體動物調查，以期建立龜山島的軟體動物清單。我們同時考量地形與棲地類型，特別針對以往研究所忽略的陸生軟體動物，進行調查。根據文獻回顧與我們的田野調查結果，龜山島軟體動物共有45科71屬126種。此研究提供龜山島生物多樣性的基礎資料。
Design description: Island species are vulnerable to extinction due to their relatively small population size and limited access to resources. The number of species on an island represents a dynamic equilibrium between immigration and extinction. Volcanic islands provide particularly interesting cases of island biogeography, in that their biota is erased by volcanic activity and recolonized from neighboring regions. Species on Gueishan Island may have under gone several cycles of extinction after volcanic eruption, followed by recolonization from Taiwan when the sea-level dropped during glacial periods. Human activity may also have provided opportunities for colonization of mollusks, either intentionally or accidentally. Investigations of molluscan fauna have been previously conducted on Gueishan Island, but these did not involve a comprehensive examination of land snails. We performed a literature survey using diverse databases, in order to collect previously identified reports on molluscan fauna of Gueishan Island. In addition, we performed field sampling of mollusks in marine, freshwater and terrestrial environments during 2011 and 2012 to establish the inventory of molluscan fauna of Gueishan Island. We considered both the topography of the island and the habitats of mollusks during our field investigation. We focused on the terrestrial environment, as the majority of the earlier investigations examined non-terrestrial habitats. In total, our literature survey and field investigation identified 126 species from 71 genera and 45 families of Mollusca on Gueishan Island. This dataset provides basic information on the island’s biodiversity.
分類群涵蓋範圍 ：此資料包括45科71屬126種軟體動物（表一），棲息於龜山島的海洋、淡水或陸生環境。包括腹足綱（物種數佔調查結果的88.10%）、雙殼綱（8.73%）、頭足綱（1.59%）和多板綱（1.59%）。物種數最多的前五個科分別為寶螺科（20種，15.87%）、鐘螺科（13種，10.32%）、骨螺科（11種，8.73%）、蜑螺科（8種，6.35%）和玉黍螺科（5 種，3.97%）（圖二）。
Taxonomic coverage
General taxonomic coverage description: The coverage of this dataset includes 126 species from 71 genera and 45 families of Mollusks of marine, freshwater and terrestrial environments on Gueishan Island (Table 1). It includes Class Gastropoda (88.10%), Class Bivalvia (8.73%), Class Cephalopoda (1.59%), and Class Polyplacophora (1.59%). The top five representative families are Cypraeidae (20 species, 15.87%), Trochidae (13 species, 10.32%), Muricidae (11 species, 8.73%), Neritidae (8 species, 6.35%), and Littorinidae (5 species, 3.97%) (Figure 2).
分類階層 Taxonomic ranks
動物門 Phylum: Mollusca
綱 Class: Bivalvia, Cephalopoda, Gastropoda, Polyplacophora
目 Order: Arcoida, Caenogastropoda, Heterobranchia, Neoloricata, Neritimorpha, Octopoda, Ostreoida, Patellogastropoda, Pterioida, Unionoida, Veneroida, Vetigastropoda
科 Family: Achatinidae, Aplysiidae, Arcidae, Argonautidae, Assimineidae, Bradybaenidae, Bursidae, Camaenidae, Cardiidae, Chitonidae, Clausiliidae, Columbellidae, Conidae, Corbiculidae, Cypraeidae, Diplommatinidae, Ellobiidae, Fasciolariidae, Haliotidae, Littorinidae, Lottiidae, Muricidae, Nassariidae, Neritidae, Octopodidae, Ostreidae, Ovulidae, Patellidae, Pectinidae, Philomycidae, Phyllidiidae, Planaxidae, Planorbidae, Potamididae, Pteriidae, Ranellidae, Siphonariidae, Subulinidae, Succineidae, Thiaridae, Trochidae, Turbinellidae, Turbinidae, Unionidae, Veronicellidae
屬 Genus: Achatina, Acusta, Aegista, Allopeas, Aplysia, Argonauta, Assiminea, Astralium, Barbatia, Batillaria, Bradybaena, Bursa, Calliostoma, Calpurnus, Carychium, Cellana, Chicoreus, Chlamys, Chlorostoma, Collisella, Coniglobus, Conus, Corbicula, Crassostrea, Cristaria, Cymatium, Cypraea, Diplommatina, Dolabrifera, Drupa, Ergalatax, Haliotis, Laevapex, Liolophura, Littoraria, Lottia, Lunella, Mancinella, Meghimatium, Monodonta, Morula, Nassarius, Nerita, Nodilittorina, Notoacmea, Octopus, Ovula, Patella, Peristernia, Phyllidia, Pinctada, Planaxis, Pteria, Purpura, Pyrene, Saccostrea, Siphonaria, Solenomphala, Stomatella, Succinea, Tarebia, Tectus, Telasco, Tenguella, Thais, Thiara, Tridacna, Trochus, Vaginulus, Vasum, Zaptyx.
空間涵蓋範圍 Spatial coverage
General spatial coverage: The spatial coverage of the literature and our field investigation ranged from a latitude of 24°49'48"N to 24°51'0"N and a longitude of 121°55'48"E to 121°57'36"E. It includes the marine, intertidal, freshwater and terrestrial environment of Gueishan Island, Taiwan (Figure 1)
Coordinates: 24°49'48"N and 24°51'0"N Latitude; 121°55'48"E and 121°57'36"E Longitude
時間涵蓋範圍 Temporal coverage
1934–2012.
方法 Methods
採樣描述 Sampling description
文獻搜尋：針對以下資料庫，搜尋龜山島軟體動物調查的相關出版品，包括期刊、計畫報告、博碩士論文與書籍：（1）臺灣博碩士論文知識加值系統（此資料庫囊括1956年以來的博碩士論文，未查詢到本研究可用資訊）；（2）全國圖書書目資訊網（此資料庫整合國家圖書館與其他74座圖書館的國內出版品與部分的政府研究報告，搜尋到三筆相關資料(Wu 2002, National Museum of Marine Biology and Aquarium 2003, Hwang and Lee 2003)；（3）政府研究資訊系統（此資料庫收錄1993年以來的政府研究計畫，未查詢到本研究可用資訊）；（4）Google學術搜尋（此資料庫廣括各種來源，從期刊、書籍到網頁，查詢到兩筆期刊資料(Chen and Fu 2007, Lee and Chen 2010)）；（5）台灣貝類資料庫（該資料庫包含台灣軟體動物的分類資訊、物種分布和相關文獻，查詢到六筆相關資料(Lee and Wu 1998, Jung and Lai 1999, Wu 2002, National Museum of Marine Biology and Aquarium 2003, Hwang and Lee 2003, Chen and Fu 2007)）。另外，從Wu (2002)一書的參考文獻，找到三筆相關文獻(Hayasaka and Tan 1934, Kuroda 1938, Kuroda 1941)。總計搜尋到10筆可用的文獻資料，其中三筆文獻是描述龜山島附近海域漁船捕獲的軟體動物，並未紀錄詳細的採樣地理資訊，因而排除不納入此物種清單(Lee and Wu 1998, Chen and Fu 2007, Lee and Chen 2010)。其餘的七篇文獻，用於建置物種清單，包括採集點、採集者、學名等資訊，彙整記錄於Microsoft EXCEL 2010。上述提及的所有文獻，皆可在國家圖書館和國立臺灣圖書館調閱。
Literature survey: We searched for publications (including journals, project reports, theses and books) associated with the molluscan fauna of Gueishan Island from the following databases: (1) the National Digital Library of Theses and Dissertations in Taiwan (this contains details of theses and dissertations published since 1956, but did not contain publications relevant to this study); (2) the National Bibliographic Information Network (this catalog integrates information from National Central Library and 74 other libraries containing all publications with a Taiwan ISBN and selected government project reports; three publications (Wu 2002, National Museum of Marine Biology and Aquarium 2003, Hwang and Lee 2003) from this database met our requirement); (3) the Government Research Bulletin (this contains government project reports made since 1993, but did not contain reports relevant to this study); (4) Google Scholar (this contains a wide range of resources, from journals and books to webpages, and it provided two relevant journal articles (Chen and Fu 2007, Lee and Chen 2010)); (5) The Taiwan Malacofauna Database (this database contains taxonomy, distribution and references of all mollusks occurred in Taiwan, and provided six relevant publications (Lee and Wu 1998, Jung and Lai 1999, Wu 2002, National Museum of Marine Biology and Aquarium 2003, Hwang and Lee 2003, Chen and Fu 2007)). In addition, three relevant publications (Hayasaka and Tan 1934, Kuroda 1938, Kuroda 1941) were identified from citations in Wu (2002). In total, we identified ten relevant publications. Three of these publications (Lee and Wu 1998, Chen and Fu 2007, Lee and Chen 2010) were excluded because they described specimens acquired from fishing ports that had been captured by shrimp fishing or bottom trawling boats near Gueishan Island, without information of the precise sampling location. The seven remaining publications were used to establish the occurrence and inventory data. Sampling sites, names of collectors and the scientific name of each species were recorded using Microsoft EXCEL 2010. All of the publications mentioned above can be accessed in the National Central Library and the National Taiwan Library.
田野調查：本研究同時考量龜山島的地形與棲地類型進行調查。潮間帶、淡水和陸域環境，以肉眼搜尋軟體動物（圖一）。海水退潮時，檢視海邊的岩石、港口的人造物表面是否有附著軟體動物。針對淡水環境的龜尾湖，檢視周邊的水下或水邊的落葉和石頭等底質。陸域環境，包括龜尾湖步道、401高地步道和北岸步道，檢視樹葉枝幹、枯枝落葉堆、石頭底部、朽木等。我們蝸牛活動較活躍的雨後、清晨和夜間進行調查。每個物種至少採集一個空殼或活體，以作為存證標本。活體標本帶回實驗室後，冷凍於攝氏-80度，隨後轉置於95%酒精溶液，以利於長期保存。
Field Sampling: The topology of Gueishan Island and the types of mollusk habitat were considered for field investigation. Visual search was conducted for mollusks in intertidal, freshwater and terrestrial environments (Figure 1). The surface of rocks on the coastline and man-made concrete structures in port were searched for marine mollusks during low tide. Leaf litter and rocks under or near water around Tail Lake (the only freshwater habitat on island) were inspected for freshwater mollusks. We inspected from leaves, trunks, leaves litter, rocks and rotten woods for land snails along three trails: one trail around Tail Lake, another leads to the highest peak (401 Highland) on the island, and the other leads to the northern part of the island. We surveyed for land snails during their active periods: during and after rainfall, early morning, and night. At least one living individual or dead shells of each species was collected as voucher specimens. Living organisms were brought back to laboratory, fixed via freezing in a -80°C freezer, and subsequently transferred to 95% ethanol for long term preservation.
資料品質描述：以Garmin GPSmap 60CSx衛星定位儀測量，當誤差範圍小於10公尺時，記錄該採集點的經度、緯度和海拔，座標採用WGS84系統。採集回來的標本，分別由黃致維和熊大維鑑定。文獻回顧搜尋到的七篇文獻，缺乏清晰的物種照片或其他物種描述資訊。鑑定物種是依據以下台灣軟體動物相關文獻：Pace (1973)、Lai (1990, 1998)、Lee and Chen (2003)、Wu and Lee (2005)和Hsieh et al. (2006)。初步認定的龜山島新紀錄的物種，另由中央研究院生物多樣性研究中心的博士後研究員，李彥錚博士再次鑑定。十四種龜山島新紀錄種，皆有分布於臺灣，是龜山島以往調查沒有記錄到的物種。所有物種的學名，均透過台灣貝類資料庫和World Register of Marine Species資料庫比對確認。
Quality control description: Latitude, longitude and altitude of sampling sites were recorded using Garmin GPSmap 60CSx with uncertainty of less than 10 meters. Sampling sites were georeferenced (WGS84). All the specimens collected during the field investigation were identified independently by Huang and Hsiung. Seven earlier studies described the mollucan fauna of Gueishan Island, but these publications lack clear photos or other information for identifying specimens. Species identification was performed using the following guide books and publications about Taiwan malacofauna: Pace (1973), Lai (1990, 1998), Lee and Chen (2003), Wu and Lee (2005), and Hsieh et al. (2006). Newly recorded species were further confirmed by Dr. Yen-Chen Lee, a Mollusca specialist and postdoctoral researcher in the Biodiversity Research Center, Academia Sinica. Fourteen new recorded species were found to be native to Taiwan but previously unreported on Gueishan Island. The scientific names of all mollusks were checked against the Taiwan Malacofauna Database and World Register of Marine Species.
資料來源 Data resources
本研究資料儲存於GBIF
The data underpinning the analysis reported in this paper are deposited at GBIF, the Global Biodiversity Information Facility, http://taibif.org.tw/ipt/resource.do?r=gueishan_island
資料集 Datasets
資料集描述：此資料彙整七篇龜山島軟體動物相的文獻(Hayasaka and Tan 1934, Kuroda 1938, Kuroda 1941, Jung and Lai 1999, Wu 2002, National Museum of Marine Biology and Aquarium 2003, Hwang and Lee 2003)與田野調查的結果。資料包括調查日期、分類資訊、地理座標、海拔、棲地類型、採集者姓名、調查方式和文獻紀錄。根據發表於1934-2003年的文獻，紀錄37科61屬112種。我們在2011-2012年的田野調查，共記錄到23科28屬34種，其中14種為龜山島新紀錄種。彙整文獻紀錄和田野調查結果，龜山島軟體動物相共記錄有45科71屬126種，14種龜山島新紀錄種為棲息於海邊的大駝石鱉Liolophura japonica、雜斑蓮花青螺 Lottia luchuana、花松螺Siphonaria laciniosa、黑肋蜑螺Nerita costata、蘭富蜑螺Nerita rumphii、波紋玉黍螺Littoraria undulata；採集自淡水環境，龜尾湖的山椒蝸牛未知種Assiminea sp.、盤蜷Laevapex nipponica；採集自陸域環境的台灣山椒蝸牛Solenomphala taiwanensis、烏來芝麻蝸牛Diplommatina suganikeiensis、八丈島罌粟蝸牛Carychium hachijoensis、鬼牙細煙管蝸牛Zaptyx crassilamellata、細錐蝸牛Allopeas pyrgula和台灣椎實蝸牛Succinea erythrophana。此資料提供龜山島的生物多樣性與生物地理的基礎資料。此資料將由中央研究院生物多樣性研究中心的軟體動物學研究室維護。
Dataset description: This dataset incorporates seven publications (Hayasaka and Tan 1934, Kuroda 1938, Kuroda 1941, Jung and Lai 1999, Wu 2002, National Museum of Marine Biology and Aquarium 2003, Hwang and Lee 2003) associated with the molluscan fauna of Gueishan Island and field investigation results. The dataset includes sampling date, taxonomy information, GPS location, elevation, type of habitat, name of collector, method of collection, and literature record. Based on the literature published during the period between 1934 and 2003, 112 species from 61 genera and 37 families of Mollusca were recorded on Gueishan Island. Of the 34 species from 28 genera and 23 families identified during our 2011-2012 field investigation, fourteen species were new records on Gueishan Island. In total, our literature survey and field investigation documents 126 species from 71 genera and 45 families of Mollusca on Gueishan Island. The fourteen newly recorded species are: Liolophura japonica (Lischke, 1873), Lottia luchuana (Pilsbry, 1901), Siphonaria laciniosa (Linnaeus, 1758), Nerita costata Gmelin, 1791, Nerita rumphii Recluz, 1841, and Littoraria undulata (Gray, 1839), which were sampled from the marine environment; Assiminea sp. and Laevapex nipponica (Kuroda, 1947), which were discovered in a freshwater environment, Tail Lake; and Solenomphala taiwanensis (Habe, 1942), Diplommatina suganikeiensis (Pilsbry & Hirase, 1905), Carychium hachijoensis Pilsbry, 1902, Zaptyx crassilamellata Kuroda, 1941, Allopeas pyrgula (Schmacker & Boettger, 1891), and Succinea erythrophana Ancey, 1883 discovered in the terrestrial environment. This dataset provide basic information for the island’s biodiversity and biogeography. This dataset will be maintained by the Malacology Lab, Biodiversity Research Center, Academia Sinica.
Object name: Darwin Core Archive Molluscan fauna of Gueishan Island, Taiwan
Character encoding: UTF-8
Format name: Darwin Core Archive format
Format version: 1.0
Distribution: http://taibif.org.tw/ipt/archive.do?r=gueishan_island
Publication date of data: 2012-12-21
Language: English
Licenses of use: This work is licensed under a Creative Commons CCZero 1.0 License http://creativecommons.org/publicdomain/zero/1.0/legalcode
Metadata language: English
Date of metadata creation: 2012-09-21
Hierarchy level: Dataset
致謝 Acknowledgements
感謝宜蘭縣政府和東北角暨宜蘭海岸國家風景區管理處核准龜山島的調查。田野調查工作，要感謝頭城漁會，中央研究院生物多樣性研究中心軟體動物學研究室全體人員、國立臺灣師範大學生命科學系草魚研究室全體人員的協助。感謝Duncan Wright博士與兩位匿名審稿者，提供意見而使此文更加完整。
We would like to thank Yilan County Government and Northeast and Yilan Coast National Scenic Area Administration, Tourism Bureau, MOTC for approval of field investigation on Gueishan Island. Fieldwork was made possible by the assistance of Toucheng Fishermen’s Association, all members of the Malacology Laboratory (Biodiversity Research Center, Academia Sinica) and all members of the Laboratory of Grass Lizard and Fish (Department of Life Science, National Taiwan Normal University). We would like to thank Dr. Duncan Wright as well as two anonymous reviewers for helpful comments which greatly improved the manuscript.
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