大凉螈繁殖生态
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摘要
大凉螈是我国特有的珍稀有尾两栖动物,其种群数量目前呈现明显下降趋势,然而涉及该物种保护的繁殖生态学研究仍十分匮乏。通过融合围栏陷阱及标志重捕的样方调查法,对大凉螈石棉栗子坪种群繁殖个体和变态登陆幼体的迁徙、繁殖群体种群大小、繁殖场内雌雄有效性比变化等进行了研究。运用Jolly-Seber法估测了繁殖种群大小,运用单因素方差分析或Kruskal-Wallis秩和检验比较了不同时期进入繁殖场的雄性大凉螈头体长及体重,运用t检验或者Wilcoxon秩和检验比较了雌雄性间形态上的差异,运用t检验、t′检验或Wilcoxon秩和检验比较了野外抱对雄性与非抱对雄性间的体征差别,运用Pearson相关分析探讨了雌性产卵量与其身体形态的关系,同时观察了卵的孵化情况。研究结果表明:大凉螈的繁殖季为每年的4月下旬到7月下旬,幼体最早于8月上旬变态登陆。估测调查地繁殖场内雄性大凉螈繁殖种群大小约为391尾,雄螈较雌螈更早进入繁殖场且在场内停留时间更长,体重较轻的雄螈较晚迁入繁殖场。有效性比明显偏雄(雌/雄:0.03—0.10)。雌雄间具明显性二型性,雌性个体的头体长、体重及肥满度均大于雄性,而雄性的尾高和尾长占全长的比例则大于雌性。对比自然抱对雄性和非抱对雄性个体发现,抱对个体在头体长、体重和尾高等体征方面显著大于非抱对个体,暗示这些形态特征可能在雄性竞争配偶的过程中起到关键作用。雌螈在室内条件下平均产卵数为176枚,产卵历时2—4 d,产卵量与雌性肥满度正相关,卵的平均孵化期为15.7 d,孵出幼体平均全长为9.74 mm。
The Taliang crocodile newt(Liangshantriton taliangensis) is a rare urodelean species that is endemic to China. Its population has significantly decreased, but studies of the reproductive ecology of the species, which contribute to its conservation, are limited. By introducing the quadrat method, which involved pitfall trapping and marked-recapture, we evaluated the migration of courting adults and metamorphosed larvae, population size of mature individuals, and operational sex ratio change during the breeding season at the Liziping National Nature Reserve in Shimian County. We used the Jolly-Seber model to estimate the population size of breeding males. One-way analysis of variance and Kruskal-Wallis test were used to examine the differences in the snout-vent length and body mass of male newts that entered the breeding pond at different periods. Sexual dimorphism between male and female was tested by Student′s t-test and Wilcoxon rank-sum test. Additionally, t-test, t′-test, or Wilcoxon rank-sum test was used to compare morphological parameters between amplectant and unamplectant male L. taliangensis, which we observed in the field. Pearson correlation coefficient was calculated to determine the relationship between egg-laying number and morphological traits in the female. We also examined incubation of the spawn. The results showed that the breeding season of L. taliangensis was from late April to late July and the first batch of hatched larvae finished metamorphosis and landed in early August. The population size of breeding males at the studied site was approximately 391. Male newts entered the breeding pond earlier than the female and stayed longer, while males with lighter body masses accessed the pond at a later time. The operational sex ratio skewed to male from 0.03 to 0.10 during the entire breeding season. Sexual dimorphism was overt in this species, with female newts possessing significantly longer snout-vent lengths, heavier body masses, and bigger fatness indices than males, whereas males had higher tail fins and higher tail proportions. We also found that amplectant males had greater snout-vent lengths and body masses and higher tail fins than unamplectant males, suggesting that these morphological characters play important roles in intrasexual competition for mates. Additionally, we counted the egg-laying number of 28 individual females in captivity: the average was 176, the spawning period lasted for 2—4 days for an individual female, and the egg-laying number was positively correlated with the female′s fatness. The incubating process occurred in 15.7 days on average and newly hatched larva were approximately 9.74 mm long.
The Taliang crocodile newt(Liangshantriton taliangensis) is a rare urodelean species that is endemic to China. Its population has significantly decreased, but studies of the reproductive ecology of the species, which contribute to its conservation, are limited. By introducing the quadrat method, which involved pitfall trapping and marked-recapture, we evaluated the migration of courting adults and metamorphosed larvae, population size of mature individuals, and operational sex ratio change during the breeding season at the Liziping National Nature Reserve in Shimian County. We used the Jolly-Seber model to estimate the population size of breeding males. One-way analysis of variance and Kruskal-Wallis test were used to examine the differences in the snout-vent length and body mass of male newts that entered the breeding pond at different periods. Sexual dimorphism between male and female was tested by Student′s t-test and Wilcoxon rank-sum test. Additionally, t-test, t′-test, or Wilcoxon rank-sum test was used to compare morphological parameters between amplectant and unamplectant male L. taliangensis, which we observed in the field. Pearson correlation coefficient was calculated to determine the relationship between egg-laying number and morphological traits in the female. We also examined incubation of the spawn. The results showed that the breeding season of L. taliangensis was from late April to late July and the first batch of hatched larvae finished metamorphosis and landed in early August. The population size of breeding males at the studied site was approximately 391. Male newts entered the breeding pond earlier than the female and stayed longer, while males with lighter body masses accessed the pond at a later time. The operational sex ratio skewed to male from 0.03 to 0.10 during the entire breeding season. Sexual dimorphism was overt in this species, with female newts possessing significantly longer snout-vent lengths, heavier body masses, and bigger fatness indices than males, whereas males had higher tail fins and higher tail proportions. We also found that amplectant males had greater snout-vent lengths and body masses and higher tail fins than unamplectant males, suggesting that these morphological characters play important roles in intrasexual competition for mates. Additionally, we counted the egg-laying number of 28 individual females in captivity: the average was 176, the spawning period lasted for 2—4 days for an individual female, and the egg-laying number was positively correlated with the female′s fatness. The incubating process occurred in 15.7 days on average and newly hatched larva were approximately 9.74 mm long.
引文
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[4] 蒋志刚,江建平,王跃招,张鹗,张雁云,李立立,谢锋,蔡波,曹亮,郑光美,董路,张正旺,丁平,罗振华,丁长青,马志军,汤宋华,曹文宣,李春旺,胡慧建,马勇,吴毅,王应祥,周开亚,刘少英,陈跃英,李家堂,冯祚建,王燕,王斌,李成,宋雪琳,蔡蕾,臧春鑫,曾岩,孟智斌,方红霞,平晓鸽.中国脊椎动物红色名录.生物多样性,2016,24(5):500- 551.
[5] 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//两栖爬行动物学研究(第8辑)——亚洲两栖爬行动物学第四届国际学术会议专辑.贵州:贵州科技出版社,2000:228- 232.
[6] 刘晓雯,刘克武,杨守忠,刘绍龙,赵欣平,闵丽娥,江琰,喻东.大凉疣螈碱性磷酸酶的分离纯化及部分性质.中国生物化学与分子生物学报,2001,17(1):135- 138.
[7] 曹玫,杨玉华,张义正.大凉疣螈脑源性神经营养因子(BDNF)基因的克隆与序列分析.应用与环境生物学报,2002,8(3):314- 316.
[8] Weisrock D W,Papenfuss T J,Macey J R,Litvinchuk S N,Polymeni R,Ugurtas I H,Zhao E M,Jowkar H,Larson A.A molecular assessment of phylogenetic relationships and lineage accumulation rates within the family Salamandridae (Amphibia,Caudata).Molecular Phylogenetics and Evolution,2006,41(2):368- 383.
[9] 张明旺,饶定齐,余国华,杨君兴.从线粒体Cyt b基因探讨红瘰疣螈物种地位的有效性.动物学研究,2007,28(4):430- 436.
[10] Pyron R A,Wiens J J.A large-scale phylogeny of Amphibia including over 2800 species,and a revised classification of extant frogs,salamanders,and caecilians.Molecular Phylogenetics and Evolution,2011,61(2):543- 583.
[11] 刘畅.镇海棘螈和大凉疣螈年龄结构研究[D].成都:中国科学院研究生院,2009.
[12] 赵云,刘绍龙,张寒珍.大凉疣螈的骨骼系统.四川动物,2009,28(4):561- 564.
[13] 王宁宁,刘绍龙,曾麟.大凉疣螈皮肤的组织学观察.四川动物,2011,30(1):66- 68.
[14] 李蓓,邹立扣,罗燕.大凉疣螈栖息地现状调查及其保护.四川动物,2011,30(6):964- 966.
[15] 李蓓,邹立扣,王朋,龙梅,郭放,马炳存,罗燕,刘晓琳.大凉疣螈形态学观察及消化、呼吸系统解剖.四川农业大学学报,2012,30(1):98- 102.
[16] 李蓓,吴邦元,彭西,邹立扣.成体大凉疣螈肺脏的组织形态学观察.西南师范大学学报:自然科学版,2012,37(6):92- 96.
[17] 李蓓,史一珺,龙梅,罗燕,彭西,邹立扣.成体大凉疣螈肝脏的组织形态学观察.四川动物,2013,32(3):398- 401.
[18] Jiang Y,Li Z Y,Liu J B,Li Y,Ni Q Y,Yao Y F,Xu H L,Li Y,Zhang M W.The complete mitochondrial genome sequence of Tylototriton taliangensis (Amphibia:Caudata).Mitochondrial DNA Part A,2016,27(4):2639- 2640.
[19] 叶昌媛,费梁,胡淑琴.中国珍稀及经济两栖动物.成都:四川科学技术出版社,1993.
[20] 李成,李娟,梁春平,赵联军,王大军,江建平.王朗自然保护区地栖脊椎动物群落结构和生境类型的关系.生态学报,2017,37(12):4247- 4257.
[21] 费梁,胡淑琴,叶昌媛,黄永昭.中国动物志两栖纲(上卷)总论蚓螈目有尾目.北京:科学出版社,2006.
[22] 戴强,戴建洪,李成,刘志君,王跃招.关于肥满度指数的讨论.应用与环境生物学报,2006,12(5):715- 718.
[23] Heyer W R,Donnelly M A,McDiarmid R W,Hayek L A C,Foster M S.Measuring and Monitoring Biological Diversity:Standard Methods for Amphibians.Washington,London:Smithsonian Institution Press,1994.
[24] 牟迈.文县疣螈的繁殖生态学与物种保护研究[D].兰州:西北师范大学,2008.
[25] 龚大洁,牟迈,李晓鸿,滕继荣,张可荣.文县疣螈繁殖初探.动物学杂志,2008,43(4):48- 55.
[26] 向孙军.文县疣螈繁殖习性和胚胎发育的研究[D].长沙:湖南师范大学,2010.
[27] Kuzmin S L,Dasgupta R,Smirina é M.Ecology of the Himalayan newt (Tylototriton verrucosus) in Darjeeling Himalayas,India.Russian Journal of Herpetology,1994,1(1):69- 76.
[28] 张晓翠.云南新平哀牢山红瘰疣螈的繁殖迁徙[D].昆明:云南师范大学,2014.
[29] 王延斌,陈永祥,胡思玉.贵州疣螈繁殖习性的观察.四川动物,1995,14(3):126- 128.
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