摘要
文县疣螈(Tylototriton wenxianensis Fei, Ye et Yang, 1984)俗称山娃娃鱼,隶属有尾目(Caudata)、蝾螈科(Salamandridae)、疣螈属(Tylototriton),是我国特有的珍稀有尾两栖动物,因在甘肃文县发现而得名,在四川、重庆、贵州、湖南、安徽等地也有分布。目前,有关文县疣螈的研究报道甚少,仅限于分类、形态描述和生态习性方面,其他方面几乎处于空白。因此,研究文县疣螈的繁殖和种群分布不仅能填补该领域的空白,为其他研究提供基础资料,也能为该物种的保护工作起到一定的指导作用。
2006～2007年,在甘肃文县和四川青川县,采用直接计数法、样方法、标志重捕法和访问法,调查了文县疣螈的种群数量和分布。在繁殖场实地观察并记录了其雌雄特征、性比、繁殖行为等,并利用仪器测量了各种生境指标。室内饲养过程中,观察和记录了幼体的生长发育。数据以SPSS和Excel 2003软件统计分析。另外,还利用光学显微镜和透射电子显微镜观察和研究了精子和精巢的微观结构。
结果表明,文县疣螈在甘肃、四川的分布主要在文县和青川县,平武县也有分布,其中青川的种群数量最多。调查中,共发现文县疣螈繁殖场3处,其中2处发现成体,而在繁殖场之外仅发现零星个体。共发现成体83尾,幼体141尾,卵302粒。
成体雄螈和雌螈的形态量度无明显差异,利用数量性形态学特征鉴定该螈的性别不可行。因此,目前较为常用和可靠的方法仍是雌、雄泄殖腔部差异鉴别和解剖学方法。
该螈3月底出蛰,随着气温升高和降雨量增加,开始向繁殖场迁移。通常是雄性先迁入繁殖水塘周围的陆地上占区后,雌性才迁入,但雌性产卵后即迁出,而雄性迁出通常在繁殖末期。繁殖期性比明显偏雄,雌雄性比先逐渐升高,后逐渐下降,在5月中下旬达到最高。
求偶时雄螈占主动,并伴有竞争现象。求偶、交配和产卵均在繁殖水塘周围的陆地上进行。产卵环境颇为一致,一般在腐叶间或以腐叶覆盖的泥土上。体内受精,雌螈纳精后即开始选择产卵位点,然后筑巢,产卵多在筑巢后24h开始。从纳精到产下第一粒卵的间隔约为48 h(2d)。产卵时间多在傍晚至次日黎明,集中在20:00~次日4:00,昼时较少见到。每次产卵1粒,雌螈产卵期长达(26.8±1.3)d (n=6),产卵量在(42.8±11.9)粒(n=6)左右。窝卵数与成体大小成正相关,线性回归方程为:Y=-27.27+0.859x,[r=0.924, P<0.001]。
刚产出的卵近球形,周围由一层卵外胶次膜包裹,卵外胶次膜在(5.1±1.3)d(n=50)后消失。卵由内层的胚盘和外层的卵胶囊构成。初生卵单粒重0.203(0.098~0.233)g(n=52),卵径为(4.2±1.8)mm(n=52)。动物极浅棕色,植物极乳白色,中央具胚盘,由于地心引力作用胚盘总是向下。
精子除可见头部、颈部和尾部外,其他结构不明显。精巢的光镜观察可见曲细精管、精小叶、输出管等结构以及各级生精细胞和精子束。精巢间质为富含血管和淋巴管的疏松结缔组织,除成纤维细胞、血细胞外,还可见间质细胞。电镜下,各级生精细胞及支持细胞、间质细胞核较明显,滤泡细胞除核外,可见明显的线粒体、糙面内质网等细胞器。
文县疣螈的致危因素主要为生境退化和丧失、过度采集和环境污染等。其中,生境丧失是最直接的影响因素。生境丧失不仅剥夺了文县疣螈基本的生存条件,更使其繁殖活动受到严重影响。因此,应大力加强该物种生境的保护,在维护现有生境的基础上,开辟新的生境,如新建繁殖场和生境走廊等,保证其繁殖活动正常进行,尤其要避免对产卵场的干扰和破坏。另建议提高保护级别,鼓励国家和当地政府制定保护措施。同时,应加强宣传教育,尽量减少人为捕捉现象和栖息地的环境污染,并积极开展迁地保护和人工繁育。
Tylototriton wenxianensis (Fei, Ye et Yang, 1984)also called Shan wa-wa yu, on attachment to Tylototriton, Salamandridae, Caudata, is epidemic to China, first discovered in Wenxian of Gansu Province and also rarely distributed in Sichuan, Guizhou, Hunan, Anhui and Chongqing. Since the species is so rarely known and narrowly distributed, little studies were made on it, and they are limited in classification, morphology and ecology while almost no studies in other field. Therefore, the study on breeding and population distribution could not only fill the gap and provide basic document for other studies, but could also guide the protection of the species.
During 2006 to 2007, in Qingchuan of Sichuan province and Wenxian of Gansu province, we investigated into the population and distribution using direct counting method, quadrat sampling method, marking-recapture method and access method. We also observed and recorded the breeding habitat, sex ratio during the breeding season, male and female fetures, breeding behaviors, growth and development of, and captive feeding and measured kinds of habitat index with instruments. The data were analyzed with SPSS and MS Excel 2003. We also observed the microstructure and ultra structure of sperm and spermary through optical microscope and TEM, in contrast with the species of Salamandridae.
The result shows that the newts distributed in Gansu and Sichuan are mainly focused in Wenxian and Qingchuan, as well as Pingwu and Qingchuan has the largest population. The investigation has found 3 breeding sites (2 with adults) and no adults were found beyond the breeding sites, along with 83 adults, 141 juveniles and 302 eggs. There are no violent differences in morphological measurement of adult males and females, so it’s not feasible to distinguish between the sexes by quantitative morphology. The sex ratio was obviously desciscent to male in breeding seasons. The ratio of males to females rose at first and then fell. The population reached a peak during mid-May to late may as well as the ratio of females. The newt came out of hibernation in late March and set out to migrate to the breeding sites with the rising temperature and rainfall. The observation showed that at first the males migrated into the land around the pool and occupy the territory, and then the females got in. However, the females migrated out as soon as they finished breeding while the males at the end of breeding season. The breeding habitat looked almost the same. The female finished spawning in a short time without male involved. Spawning lasted 2-3 days smoothly in rainy seasons without interference. They moved to the spawning pools no sooner than they selected the proper sites. Spawning began one day later. The female spawned one a time lasting half a hour, and the whole course 5-6 hours. The course maintained a even pace slowly without a peak. The body size was positively related to the number of eggs per nest. Regression equation: Y=-27.27+0.859x, [r=0.924, P<0.001]. The eggs firstly laid were almost round wrapped by zonated secretion. The secretion was filmy when unfolded which was called meninx secundaria. The secretion disappeared in (5.1±1.3) d (n=50). The egg consisted of blastoderm inside and egg jelly outside. The firstly laid egg weighed 0.203(0.098~0.233) g (n=52) with the diameter of (4.2±1.8) mm (n=52). The animal pole was light brown and vegetate pole milky white. There’s a blastoderm in the center which kept hanging downwards attracted by the gravity from the earth.
The sperm consisted of head, neck and tail, which were different from those of reported salamanders. The ultra structure of spermary showed spermatogenous cells, spermatocytes, sustentacular cells, interstitial cell, follicular cells etc., and no spermoblasts and sperms were found.
The main factors causing the newt in severe danger are the loss and destruction of habitat, excessive seizure and pollution. The loss of habitat is the most direct factor which could not only deprive the newt of basic life condition but could also seriously affect the breeding behavior. Therefore, the habitat protection should be strengthened and open up such new habitat as new breeding sites and transit corridors when maintaining the current habitat. To guarantee regular breeding behavior and especially to avoid destroying the breeding sites. We suggest upgrading the protection level and encouraging the state and local government to draw up measures for protection. Meanwhile, artificial seizure and pollution in habitat should be reduced and allochthonous protection and artificial breeding should be carried out.
引文
蔡春抹,费梁.镇海棘螈Echinotriton chinhaiens(iChang)新模描述及其生态习性[J].两栖爬行动物学报, 1984, 4(2):109～114.
陈壁辉.安徽两栖爬行动物志[M].合肥:安徽科学技术出版社,1991:222～224.
邓其祥,余志伟.四川省细痣疣螈一新亚种—平武亚种[J].两栖爬行动物学报, 1984, 3 (2):75～77.
费梁,叶昌媛,杨戎生.疣螈属一新种和一新亚种(蝾螈目:蝾螈科)[J].动物学报, 1984, 30(1):85～91.
费梁,叶昌媛.蓝尾蝾螈繁殖生态的研究[J].生态学报, 1988, 8(3): 233～241.
费梁,叶昌媛,黄永昭.中国两栖动物检索[M].重庆:科学技术文献出版社重庆分社,1990.
费梁.中国两栖动物图鉴[M].郑州:河南科学技术出版社, 1999.
费梁,叶昌媛,黄永昭等.《中国两栖动物检索及图解》[M].四川科学技术出版社.2005: 41～42.
费梁,胡淑琴,叶昌媛等.中国动物志两栖纲上卷——总论、蚓螈目、有尾目[M].北京:科学出版社, 2006, 309～310.
耿家举,章菊明,蔡堡.东方蝾螈Cynops orientalis早期胚胎发育的初步观察[J].动物学报, 1960, 12(2):175～184.
国家林业部.国家重点保护野生动物名录[M].北京:国家林业部, 1989.
何晓瑞.我国特有种滇螈的绝灭及其原因分析[J].四川动物, 1998, 17(2): 58～59.
胡淑琴,赵尔宓,刘承钊.贵州两栖爬行动物调查及区系分析[J].动物学报, 1973,19(2):149.
胡思玉.贵州疣螈繁殖行为初步观察[J].毕节师范高等专科学校学报,1994,(4): 10～12.
江润祥.香港特别行政区两栖爬行动物多样性分析[J].四川动物, 2000,19(3): 112～115.
蒋志刚.动物行为原理与物种保护方法[M].北京:科学出版社, 2004, 299～306.
卡逊,刘惠宁,鲍嘉天.香港的两栖和爬行类[M].周家炜译.香港市政局出版, 1988, 1～132.
寇治通,邢一玲.蝾螈属一新种—呈贡蝾螈[J].两栖爬行动物学报, 1983, 2(4):51～54.
李义明.种群生存力分析:准确性和保护应用[J].生物多样性, 2003, 11(4): 340～350.
刘进辉,江辉,谭理琦,周玉林,李丽平.大鲵精巢的解剖及组织形态结构研究[J].经济动物学报, 2004, 8(3): 163～166 .
罗素元,谭兵兵,王乾兴,曾令媛.贵州疣螈精巢的显微和超微结构[J].遵义医学院学报. 1999, 22(3):20~22.
罗亚平,姜国诚,孙艳香,黄华苑.中国雄性大鲵生殖系统的解剖学和组织学研究[J].广西师范大学学报(自然科学版), 2003, 21(2): 83～87.
青川县志编纂委员会.青川县志[M].成都:成都科技大学出版社, 1992, 201～202.
曲韵芳,张继秀,郭汉身等.东方蝾螈排精和纳精的初步观察[J].动物学杂志, 1964,(2):77～78.
四川植物志编辑委员会.四川植物志第4卷[M].成都:四川人民出版社, 1985.
四川资源动物志编委会.四川资源动物志第一卷[M].成都:四川科学技术出版社, 1985.
覃丽梅,谢锋,郑中华,江建平.西藏山溪鲵精子的形态[J].动物学杂志, 2005, 40(2): 71～74.
田应洲,孙爱群,李松.贵州疣螈繁殖习性的观察[J].动物学杂志, 1997, 32(1):20～23.
田应洲,孙爱群,李松.贵州疣螈繁殖繁殖生态的研究[J]..四川动物, 1998, 17(2):60～64.
王宏元,张育辉.北方山溪鲵精巢显微结构的年周期变化[J].动物学研究, 2004, 25 (6):484～490.
王宏元,张育辉.北方山溪鲵精巢支持细胞的显微与超微结构观察[J].陕西师范大学学报(自然科学版), 2004, 32(1): 86～89.
王宏元,张育辉.北方山溪鲵精巢间质细胞的超微结构研究[J].西北农林科技大学学报(自然科学版), 2006, 35(2): 5～8.
王宏元,张育辉.北方山溪鲵精子发生不同阶段的显微与超微结构[J].动物学杂志, 2007,42(5):136~140.
王香亭.甘肃脊椎动物志[M].兰州:甘肃科学技术出版社, 1991, 1204～1205.
王延斌,陈永祥,胡思玉.贵州疣螈繁殖习性的观察[J].四川动物, 1995, 14(3):126～128.
谢锋,费梁,叶昌媛.镇海棘螈产卵位点及其对繁殖成功率的影响[J].动物学报, 2002, 40: 262～265.
杨道德,沈猷慧.东方蝾螈繁殖生态的研究[J].动物学研究,1993,14(3):215～220.
姚崇勇,张绳祖,龚大洁.甘肃有尾两栖动物的研究[A].见:中国黄山国际两栖爬行动物学学术会议论文集[C].北京:中国林业出版社,1993.38～47.
叶昌媛,费梁,胡淑琴.中国珍稀及经济两栖动物[M].成都:四川科学技术出版社, 1993, 1～412.
郑中华,谢锋,费梁,叶昌媛.镇海棘螈和大凉疣螈精子形态研究[J].两栖爬行动物学研究, 2000, 8: 228～232.
郑中华,江建平,谢锋,费梁,叶昌媛.龙洞山溪鲵精子形态研究[J].两栖爬行动物学研究, 2002, 9:4～9.
郑中华,谢锋,江建平,费梁.镇海棘螈的精子结构[J].动物学报, 2004,50(4): 622～629.
郑中华,江建平,谢锋.龙洞山溪鲵精子的超微结构[J].动物学报, 2005,51(4):703～709.
周海燕,杨楚彬,肖亚梅,刘文彬,刘筠.大鲵雄性生殖系统的组织学和超微结构研究[J].科学技术与工程, 2004, 4:187～192.
Arnold, SJ. The Evolution of Courtship Behaviour in Salamanders [D]. Ph. D. Thesis. University of Michigan. 1972.
Arnold, SJ. The Evolution of Courtship Behaviour in new world salamanders with some comments on old world salamanders [D]. In: Taylor, D. H., Ph. D. Thesis. University of Michigan. 1977.
Arntzen, JW. Seasonal variation in sex ratio and asynchronous presence at ponds of male and female Triturus newts [J]. Journal or Herpetology 2002, 36(1): 30～35.
Baker CL. Spermatozoa and spermatelcosis in Cryptobranchus and Necturus [J]. Tenn. Acad, Sci. 1963, 38: 1～11.
Barker KR, Baker CL. Urodele spermatelcosis: a comparative electron microcope study [A]. In: Baccetti B ed. Comparative Spermatology[C].New York: Academic Press, 1970, 81～84.
Cogalniceunu D. The relative importance of vision and olfation in mate recognition on male newts (Genus Triturus) [J]. Herpetologica 1994, 50: 344～349.
Cramp S, Simmons KEL. Handbook of the Birds Rope, the Middle East and North Africa, Vol.Ⅱ[M].Oxford: University Press.1980.
Dawley, EM. Identification of sex through odors by male red-spotted newts, Notophalmus viridescens [J]. Herpetologica 1984, 40: 101～105.
Douglas, ME. Migration and sexual selection in Ambystoma jeffersonianum [J]. Canadian Journal of Zoology 1979,57: 2303～2310.
Fawcett DW. A comparative view of sperm ultrastructure [J]. Biol.Reprod. Supl. 1970, 2: 90～127.
Fraile B ,Paniagua R ,Rodriguez M C ,et al., Annual changes in thenumber ,testosterone content and ultrastructure of glandular tissuecells of the testis in the marbled newt Triturus marmoratus [J]. 1989,167 :85～94.
Gabor CR, Krenz JD, Robert RG. Female choice, male interference, and sperm precedence in the red-spotted newt [J]. Behavioral Ecology2000, 11:115～124.
Garnier DH. A ndrogen and estrogen levels in the plasm a of Pleurodeles Waltl, M ichah, during the annual cycle [J]. Gen Comp Endocrinol, 1985,58 (3): 376～385.
Gong DJ, Mu M. Behavioural observation and description of the endangered knobby newt Tylototriton wenxianensis and its application in conservation [J]. AsiaticHerpetological Research 2008, 11:31～38.
Griffiths. Newts and salamanders of Europe [M].Poyser Natural History, London.1996, 322.
Halliday TK. The courtship of European newts: An evolutionary perspective [A]. In: Taylor DH, Guttman SI, eds. The reproductive biology of amphibians[C]. New York: Plenum Press, 1997,185～232.
Halliday, TK. The courtship of European newts: An evolutionary perspective [J]. Pp. 185～232. In D. H. Taylor and S. I. Guttman. The Reproductive Biology of Amphibians.Plenum Press, New York. 1997.
Halliday, TR. Sperm composition in amphibians [A]. Pp. 465～502. In T.R.Birkhead and A. P. Moller, Sperm Composition and Sex Selection[C]. Academic Press, London. 1998.
Harris, W. E. and J. R. Lucas. A state-based model of sperm allocation in a group-breeding salamander [J]. Behaviour Ecology 2002,13: 705～712.
Hasumi M , Iwasawa H, Nagahama Y. Seasonal changes in plasma a concentration of sex steroids in the salamander Hynobius nig rescens [J].Gen Comp Endocrinol, 1993,90: 51～57.
Houck, L. D. The effects of body size on male courtship success in a plethodontid salamander [J]. Animal Behaviour 1988, 36: 837～842.
Houck, L. D. and D. M. Sever. The role of the skin in reproduction and behaviour [A]. Pp. 351～388.
In H. Heatwole and G. Barthalamus (eds.), Amphibian Biology, Vol. 1, The Integument, Survey Beatty and Sons, Chipping Norton, New South Wales, Austrilia. 1994.
IUCN. 2006 IUCN Red List of Threatened Species. Cambridge, UK: IUCN, 2006.
Jackson GL, Hansen H, Kao C. Continuous exposure of Suffolk ewes to an equatorial photoperiod disripts expression of the annual breeding season [J]. Biology of Reproduction 1990, 42: 63～73.
Karsen, S. J., Lau, M.W.N. & Bogadek, A. Hong Kong Amphibians and Reptiles (2nd Edition) [M]. 1998.
Kawamura T, Sawada S. On the sexual isolation among different species and local races of Japanese newt [J]. Journal of Science-Hiroshima University Series B 1959, 118: 17～31.
Krenz JD, Sever D M. Mating and oviposition in paedomorphic Ambystoma talpoideam precedes the arrival of terrestrial males [J]. Herpetologica 1995,51: 387～393.
Kuramoto M. Scanning electron microscopic studies on the spermatozoa of some Japanesesalamanders (Hynobiidae: Crytobranchidae: Salamandridae) [J].Japanese Journal of Herpetology 1995, 16(2):49～58.
Kuramoto M, Tanaka S. Sperm morphology of Ehinotriton andersoni (Caudata: Salamandridae) [J]. Amphibia-Reptilia1997, 8 (3):308～313.
Lazard L. Steroidogenesis in axolotl testis. Histo chemistry of two major enzymes related to cell type, sperm atogenesis, and substrate [J]. Gen Comp Endocrinol, 1979,39 (3): 381～387.
Lee YH, Kwon AS, Ultra structure of spermatozoa in Urodele and primitive Anura (Amphibia) with phylogenetic considerations [J]. Korean J. SystematicZool. 1996, 3:252～264.
Mabry M, Verrell P A. All are one and one is all: sexual uniformity among widely separated population of the North American seal salamander, Desmognathus ochrophaeus [J]. Bio J Linn Soc, 2003, 78:1～10.
Malacarne G. Female attractiveness and male choice in the crested newts, Triturus cristatus carnifex (Laur.) [J]. Monitore Zoology Italia 1984, 18: 33～39.
Malacarne G, Vellano C. Behavioral evidence of a courtship pheromone in the crested newt, Triturus cristatus carnifex Laurenti [J].Copeia 1987: 245～247.
Massey, A. Sexual interactions in res-spotted newts’population [J]. Animal Behavior 1988, 36:205～210. Mead L S, Verrell P L. Evolution of courtship behavior patterns and reproductive isolation in the Desmognathus ochrophaeus complex [J]. Ethology 2002, 108: 403～427.
Nicander L, 1970. Comparative studies on the fine structure of vertebrate spermatozoa [A]. In: BaccettiBed. Comparative Spermatology [C]. NewYork: Academic Press, 4～12.
Nussbaum. The evolution of parental care in salamanders [J]. Miscellaneous Publications, Museum of Zoology, University of Michigan 1985, 169:1～50.
Peterson MG. Nest, but not egg, fidelity in a territorial salamander [J]. Ethology 2000,106: 781～794.
Petranka JW. Geographic variation in the mode of reproduction and larval characteristics of small-mouthed salamander (Ambystoma texanum) in the east-central United States [J]. Herpetologica 1982,38: 475～478.
Proper C. R. Courtship in the rough-skinned newt Taricha granulose. [J] Animal behavior, 1991., 41:547～554.
Pudney J, Canick J A, Mak P, et al. The differentiation of Leydig cells, steroid genesis, and the spermatogenetic wave in the testis of Necturus maculosus [J]. Gen CompEndocrinol , 1983, 50 :43～66.
Rafinski J, Osikowski A. Sperm mixing in the Alpine newts (Triturus alpestris) [J]. Canadian Journal of Zoology 2002, 80: 1293～1298.
Rollmann, SM., L. D. Houck and R. C. Feldhoff. Proteinacious pheromone affecting female receptivity in a terriestrial salamander [J]. Science 1999, 285: 1907～1909.
Salthe, SN. Courtship patterns and the phylogeny of Urodeles [J]. Copeia 1967: 100～117.
Salthe, SN.and Mecham JS.. Reproductive and courtship patterns [A]. In: Loffs, B.The physiology of the amphibians[C], Vol 2. New York: Academic Press: 1974, 309-521.
Sawada S. Studies on the local races of Japanese newt, Triturus phrhogaster Boie.Ⅱ.Sexual isolation mechanism [J]. Journal of Science-Hiroshima University Series B Division 1 1963,21:167～180.
Selmi MG, Brizzi R, Bigliardi E. Sperm morphology of salamandrids (Amphibia: Urodela): mplications forphylogenyan fertilization biology [J]. Tissue Cell 1997, 29(6):651～664.
Sequeira F, Ferrand N, Crespo E. Reproductive cycle of the golden striped salamander Chioglossa lusitanica (Caudata: Salamandridae) in NW Portugal [J]. Amphibia-Reptilia , 2003, 24:1～12.
Serbiolova, M. and H. Ota. Notes on the life-history and reproductive behaviour of Cynops ensicauda popei (Amphibia: Salamandridae) [J]. Herpetological Journal ,1995, 5: 310～315.
Sever DM, Klopfer NM. 1993. Spermathecal cytology of Ambystoma opacum(Amphibia: Ambystomatidae) and the phylogeny of sperm storageorgans in female salamanders [J].Morphology 217:115～127.
Sparreboom M, Xie F, Fei L. Reproductive behaviour of the Chinhai Salamander (Echinotriton chinhaiensis) (Caudata: Salamandridae) [J]. Amphibia-Reptilia, 2001, 22: 309～320.
Sparreboom M. Courtship in the newts of the genus Triturus and Paramesotriton [A]. In Rocek Z(ed.).1986. Studies in Herpetology[C], 529～534.
Sparreboom M. Een onbekende Salamander uit China: Paramesotriton caudopunctatus [J].Lacerta, Leiderdorp, 1981, 39(8):102～108.
Sparreboom M. Haultung und Nachzucht Tylototriton verrucosus [J]. Elaphe, 1999,7(2): 20～24.
Sparreboom M. On the sexual behavior of Paramesotriton caudopunctatus (Liu and Hu) (Amphibia: Caudata: Salamandridae) [J]. Amphibia-Reptilia, 1983, 4:25～33.
Sparreboom, M. and H. Ore. Notes on the life-history and reproductive behaviour of Cynops ensicauda Popi(Amphibia: Salamandridae) [J]. Amphibia-Reptilia 1995, 16: 15.
Twitty, VC. Of scientists and salamanders [M]. Freeman, San Francisco. 1966, 101.
Uribe MC, Rios GG and Brandon RA. Spermatogenesis in the urodele Ambystoma dumerilli [J]. Morphology. 1994,222: 287～299.
Utsunomiya Y, Utsunomiya T and Kawachi. Some ecological observations of Tylototriton andersoni, a terrestrial salamander occurring in the Tokunooshima Island. Proc. Japan Acad., 1978, 54, ser. B.(7):341～346.
Veith, M., Steinfartz S.and R. Zardoya. 1998. A molecular phylogeny of“true”salamander (family salamandridae) and the evolution of terrestriality of reproductive modes[J]. Jouanal of Zoology Syetem Evolution Research 36: 7～16.
Verrell P, Mabry M. The courtship behaviour of Plethodontid salamander [A]. In: Bruce D. The Biology of Plethodontid Salamander [M]. New York: Kluwer Academic/Plenum Publisher. 2000.
Verrell PA. Male mate choice for large fecund females in the red–spotted newt, Notophthalmus viridescens: how is size assessed [J]? Herpetologica 1985, 41:382～388.
Verrell PA. Male discrimination of large more fecund females in the smooth newt (Triturus vulgaris) [J]. Journal of Herpetology1986, 20:416～422.
Verrell PA. Male mate choice for fecund female in a plethodontid salamander [J]. Animal Behavior 1989, 38:1086～1088.
Verrell PA. Insemination temporarily inhabits sexual responsiveness in female salamanders (Desmongnathus ochphaeus) [J]. Behaviour 1991,119:51～64.
Verrell PA. Population and species divergence of chemical cues that influence male recognition of females in Desmognathine salamanders [J]. Ethology 2003, 109: 577～586.
Verrell, PL. and Krenz JD. Competition for mates in the mole salamander, Ambystoma talpoideum: tactics that may maximize male mating success [J]. Behaviour 1998, 135: 121～138.
Werner G, On the development and structure of the neckin urodele sperm [A]. In: Baccetti Bed. Comparative Spermatology [C] New York:Academic Press, 1970.8～15.
Whiteman TG. Territorial defense in gall aphids [J]. Nature 1999,279: 324～325.
Wortham JW, Brandon AJr, Martan J, Comparative morphology of some plethodontid salamander spermatozoa [J]. Copeia 1977, (4):666～680.
Wortham JW, Murphy JA, Martgan J, Brandon RA,Scanning electronmicroscopy of some salamander spermatoza [J].Copeia1982, (1):52～60.
Xie F, Fei L, Ye CY. Oviposition sites and their effects on the reproductive success in the Chinhai Salsmander (Echinotrition chinhaiensis) [J]. Acta Zoologica Sinica 2002, 40: 262～265.
Zuiderwijk A, Sparrebaum M. Sexual strategies in the newts Triturus cristatus and Triturus marmoratus [J].Bijdr Dierk, 1990, 60:51～64.
Zuiderwijk A, Sparrebaum M. Territorial behavior in crested newt Triturus cristatus and marbled newt T. marmoratus (Amphibia: urodela) [J]. Bijdr Dierk, 1986, 56:205～213.