
Biodiv Sci ›› 2026, Vol. 34 ›› Issue (2): 25250. DOI: 10.17520/biods.2025250 cstr: 32101.14.biods.2025250
• Original Papers: Animal Diversity • Previous Articles Next Articles
Enmei Yang1, Zuwei Yang2, Xi Lu1, Anru Zuo2, Yueti Li2, Yubao Duan1,*(
)
Received:2025-07-01
Accepted:2025-11-12
Online:2026-02-20
Published:2026-03-23
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E-mail: About author:First author contact:# Co-first authors
Supported by:Enmei Yang, Zuwei Yang, Xi Lu, Anru Zuo, Yueti Li, Yubao Duan. Behavioral ethogram and establishment of PAE coding system for three hornbills during the breeding season in Yingjiang, Yunnan[J]. Biodiv Sci, 2026, 34(2): 25250.
| 编码 Code | 姿势 Posture | 描述 Definition | 双角犀鸟 Great hornbill | 花冠皱盔犀鸟 Wreathed hornbill | 冠斑犀鸟 Oriental pied hornbill |
|---|---|---|---|---|---|
| P1 | 站 Standing | 双腿或单腿直立以承受身体重量 Stand upright on both legs or one leg to support body weight | √ | √ | √ |
| P2 | 蹲 Squatting | 双腿弯曲, 略贴于腹部, 背部略低, 身体重心略向后移 Bend both legs, drawing them slightly toward the abdomen, lower the back slightly, and shift the body's center of gravity backward | √ | √ | √ |
| P3 | 走 Walking | 双脚左右交替移动, 使身体向前或向侧面发生位移 The body moves forward or sideways by alternating steps of the left and right feet | √ | √ | |
| P4 | 跳跃 Jumping | 双腿蓄力下蹬以使身体腾空 Bend and forcefully extend both legs to propel the body into the air | √ | √ | √ |
| P5 | 飞 Flying | 双翅扇动或伸直, 在空中运动 Flapping or extending wings to move through the air | √ | √ | √ |
| P6 | 悬挂 Hanging | 双腿弯曲, 使重心向上抓住巢口, 身体挂在巢口 Bend the legs to shift the center of gravity upward, grip the nest entrance, and suspend the body from the nest edge | √ | √ | √ |
| P7 | 踩背 Treading | 交配时雄性踩在雌性背上 During copulation, the male mounts the female from behind | √ | √ | √ |
Table 1 Posture codes and definitions of three hornbill species
| 编码 Code | 姿势 Posture | 描述 Definition | 双角犀鸟 Great hornbill | 花冠皱盔犀鸟 Wreathed hornbill | 冠斑犀鸟 Oriental pied hornbill |
|---|---|---|---|---|---|
| P1 | 站 Standing | 双腿或单腿直立以承受身体重量 Stand upright on both legs or one leg to support body weight | √ | √ | √ |
| P2 | 蹲 Squatting | 双腿弯曲, 略贴于腹部, 背部略低, 身体重心略向后移 Bend both legs, drawing them slightly toward the abdomen, lower the back slightly, and shift the body's center of gravity backward | √ | √ | √ |
| P3 | 走 Walking | 双脚左右交替移动, 使身体向前或向侧面发生位移 The body moves forward or sideways by alternating steps of the left and right feet | √ | √ | |
| P4 | 跳跃 Jumping | 双腿蓄力下蹬以使身体腾空 Bend and forcefully extend both legs to propel the body into the air | √ | √ | √ |
| P5 | 飞 Flying | 双翅扇动或伸直, 在空中运动 Flapping or extending wings to move through the air | √ | √ | √ |
| P6 | 悬挂 Hanging | 双腿弯曲, 使重心向上抓住巢口, 身体挂在巢口 Bend the legs to shift the center of gravity upward, grip the nest entrance, and suspend the body from the nest edge | √ | √ | √ |
| P7 | 踩背 Treading | 交配时雄性踩在雌性背上 During copulation, the male mounts the female from behind | √ | √ | √ |
| 编码 Code | 部位 Part | 动作 Act | 双角犀鸟 Great hornbill | 花冠皱盔犀鸟 Wreathed hornbill | 冠斑犀鸟 Oriental pied hornbill |
|---|---|---|---|---|---|
| 1 | 头颈部 Head and neck | 抬头 Head up | √ | √ | √ |
| 2 | 低头 Head down | √ | √ | √ | |
| 3 | 后仰 Tilt head backward | √ | √ | √ | |
| 4 | 立颈 Erect neck | √ | √ | √ | |
| 5 | 伸颈 Extend neck | √ | √ | √ | |
| 6 | 缩颈 Retract neck | √ | √ | √ | |
| 7 | 左转 Turn left | √ | √ | √ | |
| 8 | 右转 Turn right | √ | √ | √ | |
| 9 | 后转 Turn backward | √ | √ | √ | |
| 10 | 喙 Beak | 张 Open beak | √ | √ | √ |
| 11 | 合 Close beak | √ | √ | √ | |
| 12 | 鸣 Vocalization | √ | √ | √ | |
| 13 | 啄 Peck | √ | √ | ||
| 14 | 甩 Shake | √ | √ | ||
| 15 | 衔 Hold in beak | √ | √ | √ | |
| 16 | 吞 Swallow | √ | √ | √ | |
| 17 | 眼部 Eyes | 睁眼 Open eyes | √ | √ | √ |
| 18 | 注视 Stare | √ | √ | √ | |
| 19 | 闭眼 Close eyes | √ | √ | ||
| 20 | 躯干 Torso | 挺背 Arch back | √ | √ | √ |
| 21 | 屈背 Hunch back | √ | √ | √ | |
| 22 | 起身 Rise up | √ | √ | √ | |
| 23 | 下蹲 Crouch | √ | √ | √ | |
| 24 | 翅 Wings | 展翅 Spread wings | √ | √ | √ |
| 25 | 抖翅 Shake wings | √ | √ | √ | |
| 26 | 扇翅 Flap wings | √ | √ | √ | |
| 27 | 拢翅 Fold wings | √ | √ | √ | |
| 28 | 腿部 Legs | 直立 Stand upright | √ | √ | √ |
| 29 | 弯曲 Bend legs | √ | √ | √ | |
| 30 | 挠 Scratch | √ | √ | √ | |
| 31 | 抓 Grasp | √ | √ | √ | |
| 32 | 尾部 Tail | 上翘 Raise tail | √ | √ | |
| 33 | 下压 Lower tail | √ | √ | √ | |
| 34 | 排便 Defecate | √ | √ |
Table 2 Act codes of three hornbill species
| 编码 Code | 部位 Part | 动作 Act | 双角犀鸟 Great hornbill | 花冠皱盔犀鸟 Wreathed hornbill | 冠斑犀鸟 Oriental pied hornbill |
|---|---|---|---|---|---|
| 1 | 头颈部 Head and neck | 抬头 Head up | √ | √ | √ |
| 2 | 低头 Head down | √ | √ | √ | |
| 3 | 后仰 Tilt head backward | √ | √ | √ | |
| 4 | 立颈 Erect neck | √ | √ | √ | |
| 5 | 伸颈 Extend neck | √ | √ | √ | |
| 6 | 缩颈 Retract neck | √ | √ | √ | |
| 7 | 左转 Turn left | √ | √ | √ | |
| 8 | 右转 Turn right | √ | √ | √ | |
| 9 | 后转 Turn backward | √ | √ | √ | |
| 10 | 喙 Beak | 张 Open beak | √ | √ | √ |
| 11 | 合 Close beak | √ | √ | √ | |
| 12 | 鸣 Vocalization | √ | √ | √ | |
| 13 | 啄 Peck | √ | √ | ||
| 14 | 甩 Shake | √ | √ | ||
| 15 | 衔 Hold in beak | √ | √ | √ | |
| 16 | 吞 Swallow | √ | √ | √ | |
| 17 | 眼部 Eyes | 睁眼 Open eyes | √ | √ | √ |
| 18 | 注视 Stare | √ | √ | √ | |
| 19 | 闭眼 Close eyes | √ | √ | ||
| 20 | 躯干 Torso | 挺背 Arch back | √ | √ | √ |
| 21 | 屈背 Hunch back | √ | √ | √ | |
| 22 | 起身 Rise up | √ | √ | √ | |
| 23 | 下蹲 Crouch | √ | √ | √ | |
| 24 | 翅 Wings | 展翅 Spread wings | √ | √ | √ |
| 25 | 抖翅 Shake wings | √ | √ | √ | |
| 26 | 扇翅 Flap wings | √ | √ | √ | |
| 27 | 拢翅 Fold wings | √ | √ | √ | |
| 28 | 腿部 Legs | 直立 Stand upright | √ | √ | √ |
| 29 | 弯曲 Bend legs | √ | √ | √ | |
| 30 | 挠 Scratch | √ | √ | √ | |
| 31 | 抓 Grasp | √ | √ | √ | |
| 32 | 尾部 Tail | 上翘 Raise tail | √ | √ | |
| 33 | 下压 Lower tail | √ | √ | √ | |
| 34 | 排便 Defecate | √ | √ |
| 编码 Code | 环境 Environment | 双角犀鸟 Great hornbill | 花冠皱盔 犀鸟 Wreathed hornbill | 冠斑犀鸟 Oriental pied hornbill | |
|---|---|---|---|---|---|
| E1 | 巢树 Nest trees | 巢口 Nest entrance | √ | √ | √ |
| E2 | 树枝 Branches | √ | √ | √ | |
| E3 | 非营巢树 Non-nesting trees | 1-25 m | √ | √ | √ |
| E4 | 25-50 m | √ | √ | √ | |
| E5 | > 50 m | √ | √ | √ | |
| E6 | 空中 Air | 空中 Air | √ | √ | √ |
Table 3 Environment codes and classification of three hornbill species
| 编码 Code | 环境 Environment | 双角犀鸟 Great hornbill | 花冠皱盔 犀鸟 Wreathed hornbill | 冠斑犀鸟 Oriental pied hornbill | |
|---|---|---|---|---|---|
| E1 | 巢树 Nest trees | 巢口 Nest entrance | √ | √ | √ |
| E2 | 树枝 Branches | √ | √ | √ | |
| E3 | 非营巢树 Non-nesting trees | 1-25 m | √ | √ | √ |
| E4 | 25-50 m | √ | √ | √ | |
| E5 | > 50 m | √ | √ | √ | |
| E6 | 空中 Air | 空中 Air | √ | √ | √ |
| 行为多样性指数 Behavioral diversity index | 双角犀鸟 Great hornbill | 花冠皱盔犀鸟 Wreathed hornbill | 冠斑犀鸟 Oriental pied hornbill |
|---|---|---|---|
| 绝对行为多样性指数 Absolute diversity index (A) | 4.9534 | 4.9798 | 4.4142 |
| 最大行为多样性指数 Maximum diversity index (Amax) | 5.2480 | 5.2854 | 4.7004 |
| 相对行为多样性指数 Relative diversity index (r) | 0.9439 | 0.9422 | 0.9391 |
| 校正行为多样性指数 Regulate diversity index (r-variable) | 1.6511 | 1.6599 | 1.7077 |
Table 4 Behavioral diversity index of three hornbill species
| 行为多样性指数 Behavioral diversity index | 双角犀鸟 Great hornbill | 花冠皱盔犀鸟 Wreathed hornbill | 冠斑犀鸟 Oriental pied hornbill |
|---|---|---|---|
| 绝对行为多样性指数 Absolute diversity index (A) | 4.9534 | 4.9798 | 4.4142 |
| 最大行为多样性指数 Maximum diversity index (Amax) | 5.2480 | 5.2854 | 4.7004 |
| 相对行为多样性指数 Relative diversity index (r) | 0.9439 | 0.9422 | 0.9391 |
| 校正行为多样性指数 Regulate diversity index (r-variable) | 1.6511 | 1.6599 | 1.7077 |
| 行为多样性指数 Behavioral diversity index | 双角犀鸟 Great hornbill | 花冠皱盔犀鸟 Wreathed hornbill | 冠斑犀鸟 Oriental pied hornbill | |||
|---|---|---|---|---|---|---|
| 雄 Male | 雌 Female | 雄 Male | 雌 Female | 雄 Male | 雌 Female | |
| 绝对行为多样性指数 Absolute diversity index (A) | 2.2304 | 2.2197 | 2.0640 | 2.0640 | 2.2264 | 1.3455 |
| 最大行为多样性指数 Maximum diversity index (Amax) | 3.5850 | 3.7004 | 3.3219 | 3.3219 | 3.1699 | 2.5850 |
| 相对行为多样性指数 Relative diversity index (r) | 0.6221 | 0.5998 | 0.6213 | 0.6213 | 0.7023 | 0.5205 |
| 校正行为多样性指数 Regulate diversity index (r-variable) | 1.4072 | 1.4004 | 1.3023 | 2.0640 | 1.1132 | 1.3455 |
Table 5 Behavioral diversity index of three hornbill species by sex
| 行为多样性指数 Behavioral diversity index | 双角犀鸟 Great hornbill | 花冠皱盔犀鸟 Wreathed hornbill | 冠斑犀鸟 Oriental pied hornbill | |||
|---|---|---|---|---|---|---|
| 雄 Male | 雌 Female | 雄 Male | 雌 Female | 雄 Male | 雌 Female | |
| 绝对行为多样性指数 Absolute diversity index (A) | 2.2304 | 2.2197 | 2.0640 | 2.0640 | 2.2264 | 1.3455 |
| 最大行为多样性指数 Maximum diversity index (Amax) | 3.5850 | 3.7004 | 3.3219 | 3.3219 | 3.1699 | 2.5850 |
| 相对行为多样性指数 Relative diversity index (r) | 0.6221 | 0.5998 | 0.6213 | 0.6213 | 0.7023 | 0.5205 |
| 校正行为多样性指数 Regulate diversity index (r-variable) | 1.4072 | 1.4004 | 1.3023 | 2.0640 | 1.1132 | 1.3455 |
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