COMMUNITY STRUCTURE AND INFECTION OF GASTROINTESTINAL HELMINTHS IN GLYPTOSTERNUM MACULATUM FROM TIBET AUTONOMOUS REGION, CHINA
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摘要: 为了解西藏特有鱼类黑斑原鮡(Glyptosternum maculatum)消化道寄生蠕虫的群落结构和感染情况, 于2019年5—8月对383尾黑斑原鮡进行了调查。在黑斑原鮡消化道中共发现7种寄生蠕虫, 分别为深槽绦虫未定种(Bothriocephalus sp.)、原头绦虫未定种(Proteocephalus sp.)、异肉吸虫未定种(Allocreadium sp.)、新棘吻虫未定种(Neoechinorhynchus sp.)、裸鲤棘头虫(Echinorhynchus gymnocyprii)、Contracaecum eudyptulae和杆咽线虫未定种(Rhabdochona sp.), 并对各个物种的形态特征进行了描述。将黑斑原鮡按整个群体、不同性别和不同全长群体进行划分, 分别对其消化道寄生蠕虫的群落多样性和优势虫种进行分析, 并对各寄生蠕虫物种的感染情况进行统计。结果表明: 在黑斑原鮡群体中, 消化道寄生蠕虫群落的Shannon-Wiener指数为1.53, Berger-Parker指数为0.37, 优势物种为C. eudyptulae, 其感染数量、感染率、感染强度和平均丰度均为最高; 在黑斑原鮡不同性别群体中, Shannon-Wiener指数为0.26—1.57, Berger-Parker指数为0.34—0.93, 优势物种为C. eudyptulae, 雄性群体中新棘吻虫未定种也为优势虫种, 两者的感染率和平均丰度均较高; 在黑斑原鮡不同全长群体中, Shannon-Wiener指数为0.22—1.59, Berger-Parker指数为0.34—0.94, 优势物种为C. eudyptulae或新棘吻虫未定种, 感染率和平均丰度基本以二者为最高。研究进一步明确了西藏鱼类寄生虫的种类组成和寄生特点, 为研究体内寄生蠕虫的环境适应性及与宿主的协同进化提供基础资料。Abstract: Glyptosternum maculatum is the only fish belong to Glyptosternum in China, and it is the fish that only lives in the Yarlung Zangbo River in Tibet Autonomous Region. In order to understand the community structure of gastrointestinal helminths and infection of G. maculatum, the endemic fish of Tibet Autonomous Region, 383 individuals of the species were dissected from May 2019 to August 2019. Seven species of gastrointestinal helminths were found of G. maculatum, including Bathybothrium sp., Proteocephalus sp., Allocreadium sp., Neoechinorhynchus sp., Echinorhynchus gymnocyprii, Contracaecum eudyptulae and Rhabdochona sp.. Most of them were widespread parasites. The population of G. maculatum were divided into whole group, different sex groups and different total-length groups. The gastrointestinal helminths’ community diversity, dominant species and infection of G. maculatum with different groups were respectively analyzed. The results were as follows: in the whole group of G. maculatum, the Shannon-Wiener index and the Berger-Parker index of the gastrointestinal helminths were 1.53 and 0.37 respectively, and the dominant species was C. eudyptulae, which had the highest infection quantity, infection rate, infection intensity and average abundance. In the different sex group, the Shannon-Wiener index was 0.2—1.57, the Berger-Parker index was 0.34—0.93, the dominant species was Contracaecum sp. in all sex groups, and the Neoechinorhynchus sp. was the dominant species in the female group. The infection rate and mean abundance of C. eudyptulae and Neoechinorhynchus sp. were high. In the different total-length group, the Shannon-Wiener index was 0.22—1.59, the Berger-Parker index was 0.34—0.94, the dominant species was C. eudyptulae or Neoechinorhynchus sp. The infection rate and mean abundances with both helminths were high. It can be concluded that the dominant species in the gastrointestinal helminths of native fishes in Tibet Autonomous Region were mainly nematodes, and most of them were heteroparasites. Fish were only their intermediate hosts, and birds play an important role in their transmission. This study was to understand the community structure and infection of gastrointestinal helminths of G. maculatums, to further clarify the species composition and parasitic characteristics of the parasites in Tibetan fishes, and to provide basic data for studying the environmental adaptability of endohelminths and the coevolution with the host.
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图 1 深槽绦虫未定种
A. 头节顶面观; B. 头节侧面观; C. 成熟节片; D. 妊娠节片; 1. 卵黄腺; 2. 子宫; 3. 卵巢; 4. 阴茎囊; 箭头指向生殖孔
Figure 1. Bathybothrium sp.
A. apical view of scolex; B. lateral view of scolex; C. mature proglottid; D. gravid proglottid; 1. vitelline follieles; 2. uterine; 3. ovary; 4. cirrus sac; The arrow points to the genital pore
图 4 新棘吻虫未定种
A.雌虫: 1. 吻; 2. 吻腺; 3. 体壁大核; B. 雄虫: 1. 吻; 2. 吻腺; 3. 体壁大核; 4. 睾丸; 5. 黏液腺贮囊; 6. 黏液腺; 7. 贮精囊; 8. 交接囊
Figure 4. Neoechinorhynchus sp.
A. female: 1. proboscis; 2. lemniscus; 3. somatic giant nucleus; B. male: 1. proboscis; 2. lemniscus; 3. somatic giant nucleus; 4. testes; 5. cement reservoir; 6. cement gland; 7. seminal vesicle; 8. copulatory bursa
表 1 黑斑原消化道寄生蠕虫的物种组成
Table 1 Species composition of gastrointestinal helminths of G. maculatum in Tibet Autonomous Region
寄生蠕虫种类Gastrointestinal helminth 纲Class 目Order 科Family 属Genus 种Species 绦虫Cestode 绦虫纲Cestoda 头槽目Bothriocephalidea 三枝钩科Triaenophoridae 深槽绦虫属Bathybothrium 深槽绦虫未定种Bathybothrium sp. 原头目Proteocephalidea 原头科Proteocephalidae 原头绦虫属Proteocephalus 原头绦虫未定种Proteocephalus sp. 复殖吸虫Digenea 吸虫纲Trematoda 复殖目Digenea 异肉科Allocreadiidae 异肉吸虫属Allocreadium 异肉吸虫未定种Allocreadium sp. 棘头虫Acanthocephalan 始新棘头虫纲Eoacanthocephala 新棘头虫目Neoacanthocephala 新棘吻科Neoechinorhynchidae 新棘吻虫属Neoechinorhynchus 新棘吻虫未定种Neoechinorhynchus sp. 古棘头虫纲Palaeacanthocephala 棘吻目Echinorhynchidea 棘吻科Echinorhynchidae 棘吻虫属Echinorhynchus 裸鲤棘头虫Echinorhynchus gymnocyprii 线虫Nematode 线虫纲Nematoda 蛔目Ascaridida 异尖科Anisakidae 对盲囊线虫属Contracaecum Contracaecum eudyptulae 旋尾目Spirurida 杆咽科Rhabdochonida 杆咽线虫属Rhabdochona 杆咽线虫未定种Rhabdochona sp. 表 2 黑斑原群体消化道寄生蠕虫感染情况
Table 2 Infection of gastrointestinal helminths of G. maculatum in Tibet Autonomous Region
寄生蠕虫种类
Parasites helminths寄生蠕虫数量
Numbers of helminths被感染的宿主数量 (尾)
Numbers of infected host感染率
Prevalence (%)感染强度
Intensity平均丰度
Mean abundance深槽绦虫未定种Bothriocephalus sp. 27 25 6.53 1—3 0.07±0.29 原头绦虫未定种Proteocephalus sp. 47 30 7.83 1—6 0.12±0.52 异肉吸虫未定种Allocreadium sp. 44 19 4.96 1—10 0.11±0.75 新棘吻虫未定种Neoechinorhynchus sp. 141 69 18.02 1—13 0.37±1.16 裸鲤棘头虫E. gymnocyprii 7 2 0.52 1—6 0.02±0.31 Contracaecum eudyptulae 166 92 24.02 1—19 0.43±1.47 杆咽线虫未定种Rhabdochona sp. 12 9 2.35 1—3 0.03±0.23 表 3 不同性别黑斑原消化道寄生蠕虫感染情况
Table 3 Infection of gastrointestinal helminths of G. maculatum with different sexes in Tibet Autonomous Region
性别Sex 寄生蠕虫种类
Parasites helminths寄生蠕虫数量Numbers of helminths 被感染的宿主数量 (尾)Numbers of infected host 感染率
Prevalence (%)感染强度
Intensity平均丰度
Mean abundance雌性Female 深槽绦虫未定种
Bothriocephalus sp.16 14 9.66 1—3 0.11±0.37 原头绦虫未定种
Proteocephalus sp.11 7 4.83 1—4 0.08±0.41 异肉吸虫未定种Allocreadium sp. 4 3 2.07 1—2 0.03±0.20 新棘吻虫未定种Neoechinorhynchus sp. 44 20 13.79 1—19 0.30±1.64 裸鲤棘头虫E. gymnocyprii 5 1 0.69 5 0.03±0.42 Contracaecum eudyptulae 44 34 23.45 1—4 0.30±0.64 杆咽线虫未定种Rhabdochona sp. 6 3 2.07 1—3 0.04±0.31 雄性Male 深槽绦虫未定种
Bothriocephalus sp.11 11 5.45 1 0.05±0.23 原头绦虫未定种
Proteocephalus sp.36 23 11.39 1—6 0.18±0.63 异肉吸虫未定种Allocreadium sp. 40 16 7.92 1—10 0.20±1.28 新棘吻虫未定种Neoechinorhynchus sp. 102 48 23.76 1—10 0.50±1.23 裸鲤棘头虫E. gymnocyprii 1 1 0.50 1 0.00±0.07 Contracaecum eudyptulae 109 48 23.76 1—19 0.54±1.93 杆咽线虫未定种Rhabdochona sp. 6 6 2.97 1 0.03±0.17 性别不明Unknow sex 异肉吸虫未定种Allocreadium sp. 13 10 27.78 1—2 0.36±0.64 新棘吻虫未定种Neoechinorhynchus sp. 1 1 2.78 1 0.03±0.17 表 4 不同全长黑斑原消化道寄生蠕虫感染情况
Table 4 Infection of gastrointestinal helminths of G. maculatum with different total length in Tibet Autonomous Region
全长Total length (cm) 寄生蠕虫种类Parasites helminths 寄生蠕虫数量Numbers of helminths 被感染的
宿主数量(尾)Numbers of infected host宿主总数 (尾)Numbers of host 感染率Prevalence (%) 感染强度Intensity 平均丰度Mean abundance 10≤L<15 新棘吻虫未定种Neoechinorhynchus sp. 1 1 47 2.13 1 0.02±0.15 C. eudyptulae 16 12 25.53 1—3 0.34±0.67 15≤L<20 深槽绦虫未定种Bothriocephalus sp. 1 1 142 0.70 1 0.01±0.08 原头绦虫未定种Proteocephalus sp. 4 3 2.11 1—2 0.03±0.20 异肉吸虫未定种
Allocreadium sp.2 1 0.70 2 0.01±0.17 新棘吻虫未定种Neoechinorhynchus sp. 8 4 2.82 1—4 0.06±0.39 C. eudyptulae 29 23 16.20 1—4 0.20±0.55 20≤L<25 深槽绦虫未定种Bothriocephalus sp. 15 13 96 13.54 1—3 0.16±0.44 原头绦虫未定种Proteocephalus sp. 9 5 5.21 1—4 0.09±0.48 异肉吸虫未定种
Allocreadium sp.1 1 1.04 1 0.01±0.10 新棘吻虫未定种Neoechinorhynchus sp. 40 21 21.88 1—13 0.42±1.44 裸鲤棘头虫E. gymnocyprii 7 2 2.08 1—6 0.07±0.62 C. eudyptulae 36 25 26.04 1—4 0.38±0.74 杆咽线虫未定种
Rhabdochona sp.8 5 5.21 1—3 0.08±0.40 25≤L<30 深槽绦虫未定种Bothriocephalus sp. 10 10 79 12.66 1 0.13±0.33 原头绦虫未定种Proteocephalus sp. 24 17 21.52 1—3 0.30±0.67 异肉吸虫未定种
Allocreadium sp.16 11 13.92 1—4 0.20±0.63 新棘吻虫未定种Neoechinorhynchus sp. 51 30 37.97 1—5 0.65±1.05 C. eudyptulae 62 24 30.38 1—19 0.80±2.59 杆咽线虫未定种
Rhabdochona sp.4 4 5.06 1 0.05±0.22 30≤L<35 深槽绦虫未定种Bothriocephalus sp. 1 1 19 5.26 1 0.05±0.23 原头绦虫未定种Proteocephalus sp. 10 5 26.32 1—6 0.53±1.39 异肉吸虫未定种
Allocreadium sp.25 6 31.58 1—10 1.32±2.85 新棘吻虫未定种Neoechinorhynchus sp. 41 13 68.42 1—10 2.16±2.59 C. eudyptulae 23 8 42.11 1—13 1.21±2.97 表 5 黑斑原消化道寄生蠕虫群落的多样性及优势物种
Table 5 Diversity and dominant species of gastrointestinal helminths’ community of G. maculatum in Tibet Autonomous Region
分类
ClassificationShannon-Wiener指数
Shannon-Wiener indexBerger-Parker指数
Berger-Parker index优势物种
Dominant species整个群体
Whole group1.53 0.37 C. eudyptulae 性别Sex 雌性Female 1.57 0.34 C. eudyptulae和新棘吻虫未定种
C. eudyptulae and Neoechinorhynchus sp.雄性Male 1.47 0.36 C. eudyptulae 性别
不明
Unknow sex0.26 0.93 C. eudyptulae 全长Total length 10≤L<15 0.22 0.94 C. eudyptulae 15≤L<20 1.03 0.66 C. eudyptulae 20≤L<25 1.59 0.34 新棘吻虫未定种Neoechinorhynchus sp. 25≤L<30 1.49 0.37 C. eudyptulae 30≤L<35 1.33 0.41 新棘吻虫未定种Neoechinorhynchus sp. 表 6 西藏3个水域土著鱼类消化道寄生蠕虫的群落结构比较
Table 6 Comparison of community structure of native fishes’ gastrointestinal helminths in three waters of Tibet Autonomous Region
水域Water area 宿主Host 消化道寄生蠕虫物种数Total number of gastrointestinal species Shannon-Wiener指数 Shannon-Wiener index 优势物种Dominant species 文献来源Literature sources 拉萨河Lhasa River 拉萨裸裂尻鱼Schizopygopsis younghusbandi 6 1.54 希蚋杆咽线虫
Rhabdochona hellichi[30] 双须叶须鱼Ptychobarbus dipogon 4 — 新棘吻虫未定Neoechinorhynchus sp. 高原裸鲤Gymmocypris waddellii 5 — 对盲囊线虫未定种Contracaecum sp. 尖裸鲤Oxygymnocypris stewartii 5 — 对盲囊线虫未定种Contracaecum sp. 巨须裂腹鱼Schizothorax macropogon 4 1.24 新棘吻虫未定种, 希蚋杆咽线虫Neoechinorhynchus sp., Rhabdochona hellichi 异齿裂腹鱼Schizothorax oconnori 0 0 — 细尾高原鳅Triplophysa stenura 4 — 胃瘤线虫未定种
Eustrongylides sp.哲古措Chuguco Lake 高原裸鲤
C. waddellii5 0.70—1.27 线虫nematode [32] 雅鲁藏布江日喀则江段Shigatse section of the Yarlung Zangbo River 黑斑原
G. maculatum7 1.53 C. eudyptulae 本文this paper -
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