我国东南部地区夏季两种雀形目鸟类的代谢产热特征及其体温调节

采用封闭式流体压力呼吸计,分别在5-35℃和5-40℃的环境温度范围内测定了白头鹎(Pycnonotussinensis)和丝光椋鸟(Sturnussericeus)的代谢率、热传导和体温等指标,探讨其代谢产热特征。结果显示:在环境温度(Ta)为5-35℃时,白头鹎的体温基本维持恒定,平均温度为40.3±0.1℃,热中性区为26.6-32.8℃,基础代谢率为73.10±4.11mlO2/h,是体重预期值的79%;Ta在5-26℃范围内,代谢率(MR)与Ta呈负相关,回归方程为:MR[mlO2/h]=265.37-7.24Ta(℃);Ta在5-30℃范围内,热传导值最低且基本保持恒定,平均为0.24±0.01mlO2/g·h·℃,是体重预期值的126%。丝光椋鸟的热中性区为27.6-34.5℃,平均体温为40.5±0.1℃(5-40℃),基础代谢率为160.64±9.20mlO2/h,是体重预期值的90%;最低热传导为0.16±0.05mlO2/g·h·℃,是体重预期值的129%。在5-25℃范围内,MR与Ta的回归方程为:MR[mlO2/h]=377.96-7.88Ta(℃)。白头鹎和丝光椋鸟的基本生物学特征为:较高的体温,热传导和上临界温度,较宽的热中性区和较低的代谢率,符合南方小型鸟类的代谢特征.     

Effects of temperature acclimation on body mass and energy budget in the Chinese bulbul Pycnonotus sinensis

Chinese bulbuls(Pycnonotus sinensis) are small passerine birds that inhabit areas of central, southern and eastern China. Previous observations suggest that free–living individuals of this species may change their food intake in response to seasonal changes in ambient temperature. In the present study, we randomly assigned Chinese bulbuls to either a 30 °C or 10 °C group, and measured their body mass(BM), body temperature, gross energy intake(GEI), digestible energy intake(DEI), and the length and mass of their digestive tracts over 28 days of acclimation at these temperatures. As predicted, birds in the 30 °C group had lower body mass, GEI and DEI relative to those in the 10 °C group. The length and mass of the digestive tract was also lower in the 30 °C group and trends in these parameters were positively correlated with BM, GEI and DEI. These results suggest that Chinese bulbuls reduced their absolute energy demands at relatively high temperatures by decreasing their body mass, GEI and DEI, and digestive tract size.     

环境温度对白头鹎代谢产热和蒸发失水的影响

基础代谢率(BMR)是恒温动物维持正常生理机能的最小产热速率,是动物在清醒时维持身体各项基本功能所需的最小能量值。"能量需求"假设认为,鸟类短期BMR的调整可以通过改变内部器官的大小来适应能量需求。以白头鹎(Pycnonotus sinensis)为研究对象,分别在10℃和30℃的不同环境中适应4周后,测定了其BMR、蒸发失水(EWL)和内部器官(心、肝、肾、胃、小肠和整体消化道)的重量,同时分析了白头鹎内部器官的变化及与BMR的关系。驯化4周后,白头鹎暖温组(30℃)的体重有明显降低的趋势;经协方差校正体重后,低温组(10℃)单位体重BMR与整体BMR明显高于暖温组,同时两组间EWL表现出显著差异。协方差分析表明,低温组白头鹎的肝脏、肾脏、小肠及总消化道重量显著高于暖温组。低温引起白头鹎能量需求增加,内部器官发生相应改变,要求摄入更多的氧气以维持代谢平衡,导致散失更多的蒸发水。     

白头鹎对季节性驯化和温度适应的生理反应

在自然环境中,有机体某一特性的可塑性变化在适应和功能方面是非常重要的。以成年白头鹎(Pycnonotus sinensis)为研究对象,分别在野外驯化条件和实验室温度适应条件下测定静止代谢率(RMR)、蒸发失水率(EWL)和代谢活性器官的重量,同时测定肝脏及肌肉的线粒体呼吸和细胞色素C氧化酶活力(COX)。野外季节性驯化在冬季和夏季时测定;实验室温度适应分为2组,分别在10℃和30℃下适应4周后进行测定。结果显示,与夏季驯化和暖适应组相比,冬季驯化和冷适应组RMR和EWL较高、代谢活性器官较重,线粒体呼吸速率和COX活性显著增加;器官重量和细胞产热能力的适应性变化可能是导致了RMR适应性调节。结果表明,白头鹎可以表现出对季节性驯化和温度适应的生理反应,利用这种能力应付野外环境温度的波动。生理能量特性的可塑性是鸟类能量代谢的共同特征。     

Metabolism and thermoregulation in Maximowiczi's voles (Microtus maximowiczii) and Djungarian hamsters (Phodopus campbelli)

1 Metabolic rates (Vo₂), body temperature (T_b), and thermal conductance (C) were first determined in newly captured Maximowiczi's voles (Microtus maximowiczii) and Djungarian hamsters (Phodopus campbelli) from the Inner Mongolian grasslands at a temperature range from 5 to 35 °C.

2 The thermal neutral zone (TNZ) was between 25 and 32.5 °C for Maximowiczi's voles and between 25 and 30 °C for Djungarian hamsters. Mean T_b was 37.0±0.1 °C for voles and 36.2±0.1 °C for hamsters. Minimum thermal conductance was 0.172±0.004 ml O₂/g h °C for voles and 0.148±0.003 ml O₂/g h °C for hamsters.

3 The mean resting metabolic rate within TNZ was 2.21±0.05 ml O₂/g h in voles and 2.01±0.07 ml O₂/g h in hamsters. Nonshivering thermogenesis was 5.36±0.30 ml O₂/g h for voles and 6.30±0.18 ml O₂/g h for hamsters.

4 All these thermal physiological properties are adaptive for each species and are shaped by both macroenvironmental and microenvironmental conditions, food habits, phylogeny and other factors.

Seasonal effects on thermoregulatory responses of the Rock Kestrel, Falco rupicolis

Thermoregulatory responses are known to differ seasonally in endotherms and this is often dependent on the environment and region they are resident. Holarctic animals are exposed to severe winters and substantial seasonal variation in ambient temperature. In contrast, those in the Afrotropics have less severe winters, but greater variation in temperature, rainfall and net primary production. These environmental factors place different selection pressures on physiological responses in endotherms. In this study, metabolic rate (VO₂) and body temperature (T_b) were measured in captive bred Rock Kestrels (Falco rupicolus) from the Afrotropics after a period of summer and winter acclimatisation. Resting metabolic rate was significantly lower after the winter acclimatisation period than after the summer acclimatisation period, and there was a shift in the thermoneutral zone from 20–33 °C in summer to 15–30 °C in winter. However, no significant difference in basal metabolic rate between summer and winter was found. The results show that Rock Kestrels reduce energy expenditure at low ambient temperatures in winter as expected in an Afrotropical species.     

白头鹎的鸣唱结构及其鸣唱微地理变异

对武汉市区白头鹎(Pycnonotus sinensis)的鸣唱类型和音节类型进行了统计、分析,并按照采样地点划分为组(微地理种群),对组间、组内不同白头鹎个体间的鸣唱差异进行了探讨.分析来自市区5个样点的26只雄性白头鹎的667个鸣唱,共发现18种基本鸣唱型、53种音节类型.每只雄性白头鹎具有1-3种基本鸣唱型,每只个体能唱6.7(4-1 4)种音节类型.平均每个鸣唱由5.0(3-11)个音节、4.6(3-8)种音节类型组成.白头鹎的鸣唱顺序模式为平稳过渡型,并能通过以下三种方式在基本鸣唱型的基础上产生鸣唱变异:1)省略、添加或替换鸣唱中的个别音节;2)对鸣唱中的某个部分进行重复或重复不同次数;3)将不同的鸣唱型进行拼接组合.每只个体每种基本鸣唱型至少具有2.0(1-5)个变 异类型,这种变异在个体间和个体内普遍存在.个体间能共享1-2种鸣唱型.所划分的5个组内普遍存在鸣唱型和音节类型的共享,而组间则无鸣唱型共享,有音节类型共享.采用Jaccard相似性系数衡量白头鹎个体间音节的共享情况,发现个体间的音节共享程度在组内 明显大于组间,因此认为越是邻近分布的白头鹎个体具有越相似的鸣唱     

丝光椋鸟的代谢产热特征及体温调节的日周期变化

为探讨丝光椋鸟(Sturnus sericeuts)代谢产热特征及体温调节的日周期变化,本研究采用开放式氧气分析仪及数字式温度计,测定了丝光椋鸟24 h体温、体重(Mb)和基础代谢率(BMR)的连续变化.结果显示,丝光椋鸟体温、体重及代谢率的日周期变化存在显著差异.丝光椋鸟夜间体温明显低于白昼,其中体温在凌晨5时降至(40.4±0.1)℃;体重昼夜变化明显,且在20:00～次日6:00时雌雄丝光椋鸟体重的下降与时间分别存在明显的线性关系,雌性为Mb=83.46(±0.12)-0.41(±0.02)t(R2=0.992,P＜0.01);雄性为Mb =76.74(±0.15)-0.39(±0.02)t(R2=0.986,P＜0.01)(式中的t代表时间).丝光椋鸟的BMR在凌晨4时降至最低,为(1.96±0.06) ml/(g·h).结果表明,丝光椋鸟通过内源性的调节,即夜间降低体温、体重及代谢率等途径,调节生理能量平衡,从而适应昼夜环境变化.     

Seasonal variation in the metabolism-temperature relation of House Sparrows (Passer domesticus) in KwaZulu-Natal,South Africa

Many birds exhibit considerable phenotypic flexibility in metabolism to maintain thermoregulation or to conserve energy. This flexibility usually includes seasonal variation in metabolic rate. Seasonal changes in physiology and behavior of birds are considered to be a part of their adaptive strategy for survival and reproductive success. House Sparrows (Passer domesticus) are small passerines from Europe that have been successfully introduced to many parts of the world, and thus may be expected to exhibit high phenotypic flexibility in metabolic rate. Mass specific Resting Metabolic Rate (RMR) and Basal Metabolic Rate (BMR) were significantly higher in winter compared with summer, although there was no significant difference between body mass in summer and winter. A similar, narrow thermal neutral zone (25–28 °C) was observed in both seasons. Winter elevation of metabolic rate in House Sparrows was presumably related to metabolic or morphological adjustments to meet the extra energy demands of cold winters. Overall, House Sparrows showed seasonal metabolic acclimatization similar to other temperate wintering passerines. The improved cold tolerance was associated with a significant increase in VO₂ in winter relative to summer. In addition, some summer birds died at 5 °C, whereas winter birds did not, further showing seasonal variation in cold tolerance. The increase in BMR of 120% in winter, compared to summer, is by far the highest recorded seasonal change so far in birds.     

Reproductive function of Selenoprotein M in Chinese mitten crabs (Eriocheir sinesis)

Selenoproteins are present in all major forms of life, including eukaryotes, bacteria and archaea. In eukaryotic animals, selenoproteins often function as antioxidants, but rare or absent in other phyla, such as plants and fungi (except for the green alga Chlamydomonas). Selenoprotein M (SelM) is a selenocysteine containing protein with redox activity, which is involved in the antioxidant response. However, information remains limited about SelM physiology and function in marine invertebrates, particularly in crustaceans. Hence, we investigated the reproductive functionality of SelM in the Chinese mitten crab (Eriocheir sinensis), which is a commercially important yet disease vulnerable aquaculture species. The full-length SelM cDNA (928bp) strand was cloned by using PCR, based on an initial expressed sequence tag (EST) that was isolated from a hepatopancreatic cDNA library. The SelM cDNA contained a 390bp open reading frame (ORF) that encoded a putative 129 amino acid (aa) protein. SelM mRNA expression in E. sinensis was (a) tissue-specific, with the highest expression observed in the hepatopancreas, testis, ovaries and intestines. Based on this information, we then detected the different stages of tissue expression for SelM in the testis, ovary, and male crab hepatopancreas and hemolymph, and the enzyme activity of SelM in the testis. Overall, SelM was isolated successfully from the Chinese mitten crab, and its involvement in the regulation of reproduction during the period of rapid development in E. sinensis was confirmed.     

埋植雌二醇对养殖中华鲟雌性幼鱼血清生理指标的影响

用硅橡胶作载体,将剂量30 mg/kg的雌二醇(estradiol)埋植入人工养殖11龄、平均体重46.00 kg的雌性中华鲟(Acipenser sinensis)的背部肌肉内。埋植前及埋植后的第30、120、200天从尾部取血,测定血清中总蛋白、白蛋白、球蛋白、甘油三酯、总胆固醇、血糖和卵黄蛋白原的含量。结果显示,与对照组相比,除血糖含量无显著变化外,实验组鱼血清中总蛋白、白蛋白、球蛋白、甘油三酯、总胆固醇和卵黄蛋白原的含量显著升高。此结果为进一步探索外源物质促进中华鲟卵巢发育的可能途径奠定前期基础。     

笼养黑颈长尾雉和白颈长尾雉代谢产热特征及体温调节(英文)

采用封闭式流体压力呼吸仪,在5～35℃的环境温度范围内测定了黑颈长尾雉(Syrmaticushumiae)和白颈长尾雉(Syrmaticusellioti)的代谢率(MR)、热传导(C)和体温(Tb)等指标,探讨了其代谢产热特征。结果显示:黑颈长尾雉和白颈长尾雉的热中性区(TNZ)分别为24.5～31.6℃和23.0～29.2℃。在5～35℃的温度范围内,黑颈长尾雉和白颈长尾雉能保持稳定的体温,分别为(40.47±0.64)和(40.36±0.10)℃;在热中性区内,黑颈长尾雉和白颈长尾雉的平均基础代谢率(BMR)分别为(1.36±0.84)和(2.03±0.12)mLO2/(g.h),分别是体重预期值的77%和86%。在下临界温度以下,黑颈长尾雉和白颈长尾雉的最小热传导分别是(0.12±0.01)和(0.17±0.01)mLO2/(g.h.℃),分别是体重预期值的119%和124%。这两种鸟的生理生态学特征是:黑颈长尾雉和白颈长尾雉都具有较低的代谢率,较高的体温和热传导,能较好地适应南方湿热的气候特征。     

Metabolic thermogenesis and evaporative water loss in the Hwamei Garrulax canorus

Basal metabolic rate (BMR) is thought to be a major hub in the network of physiological mechanisms connecting life history traits. Evaporative water loss (EWL) is a physiological indicator that is widely used to measure water relations in inter- or intraspecific studies of birds in different environments. In this study, we examined the physiological responses of summer-acclimatized Hwamei Garrulax canorus to temperature by measuring their body temperature (T_b), metabolic rate (MR) and EWL at ambient temperatures (T_a) between 5 and 40 °C. Overall, we found that mean body temperature was 42.4 °C and average minimum thermal conductance (C) was 0.15 ml O₂ g⁻¹ h⁻¹ °C⁻¹ measured between 5 and 20 °C. The thermal neutral zone (TNZ) was 31.8–35.3 °C and BMR was 181.83 ml O₂ h⁻¹. Below the lower critical temperature, MR increased linearly with decreasing T_a according to the relationship: MR (ml O₂ h⁻¹)=266.59–2.66 T_a. At T_as above the upper critical temperature, MR increased with T_a according to the relationship: MR (ml O₂ h⁻¹)=−271.26+12.85 T_a. EWL increased with T_a according to the relationship: EWL (mg H₂O h⁻¹)=−19.16+12.64 T_a and exceeded metabolic water production at T_a>14.0 °C. The high T_b and thermal conductance, low BMR, narrow TNZ, and high evaporative water production/metabolic water production (EWP/MWP) ratio in the Hwamei are consistent with the idea that this species is adapted to warm, mesic climates, where metabolic thermogenesis and water conservation are not strong selective pressures.     

光周期对白头鹎体重、器官重量和能量代谢的影响

光周期是四季环境变化的最直接表现因素之一,并影响动物的生理变化特征。为探讨光周期驯化对白头鹎(Pycnonotussinensis)体重、器官重量及能量代谢的影响,以室温28℃、不同光周期(16L∶8D,LD组和8L∶16D,SD组)对两组白头鹎进行为期4周的光周期驯化,测定其体重、各器官鲜重和干重、基础代谢率(BMR)和食物摄入能、排泄能及同化能并计算同化率。结果发现,SD组个体体重、内部器官(肝、小肠)重量、BMR及同化率相应显著高于LD组个体;短光照刺激白头鹎显著降低摄入能、排泄能及同化能。这些结果表明:光周期对白头鹎的体重、器官重量、BMR及能量收支有着一定影响,并且短光照较长光照更能引起白头鹎体重、器官重量及能量代谢的明显变化,同时验证了中心限制假说,即白头鹎BMR与中心器官代谢(肝、小肠等)具有相关性,中心器官是改变白头鹎BMR的主要原因之一。     

Energy metabolism and thermoregulation in pygmy lorises (Nycticebus pygmaeus) from Yunnan Daweishan Nature Reserve

The pygmy loris (Nycticebus pygmaeus) is a small prosimian living in Vietnam, Laos, eastern Cambodia and the south part of China. In China it is only found in Pingbian, Hekou, Jinping, Luchun of Yunnan. As N. pygmaeus is seriously threatened by hunting, trade and habitat destruction, it is listed in Appendix II of CITES, and in 2006 the IUCN classified it as “vulnerable”. In order to understand the characteristics of energy metabolism and thermoregulation of N. pygmaeus, the resting metabolic rate (RMR) and body temperature (T_b) at different ambient temperature (T_a) of pygmy lorises, as well as body mass, energy intake, digestable energy intake, digestability and the thermal conductance were measured in captivity. The results obtained mainly are as follows: (1) Pygmy loris feed dry food averaged 12.90 ± 1.02 g/d. They could gain 214.87 ± 16.65 kJ/d from food intake, and earned 200.15 ± 16.36 kJ digestable energy intake per day with 90.13 ± 1.34% of the digestability. (2) The T_b at room temperatures was a little low (35.23 ± 0.16 °C) and varied with T_a from 25 °C to 35 °C. There was a positive relationship between T_b and T_a, which was described as: T_b = 27.22 + 0.34T_a (r = 0.880). (3) The resting metabolic rate (RMR) of the pygmy loris was 0.3844 ± 0.0162 mlO₂/g/h, which was 51.91 ± 1.90% of the previous predicted rate by Kleiber (1961) [21]. (4) The average thermal conductance of the pygmy loris (N. pygmaeus) was 0.0449 ± 0.0031 mlO₂/g/h/°C. These characteristics of energy metabolism and thermoregulation of N. pygmaeus in Yunnan Daweishan Nature Reserve might be considered as the adaptive characteristics to their environment in tropical semi-evergreen forests and secondary forests.     

中国石龙子母体孕期调温诱导幼体表型: 母体操纵假说的实验检测

卵胎生是由卵生繁殖模式通过逐渐增加卵滞留和胚胎在母体子宫内发育的时间进化而来的繁殖模式。有鳞类爬行动物(蜥蜴和蛇)有着较高的繁殖模式多样性,因而是研究卵胎生繁殖模式进化及其适应意义的理想动物模型。至今对于卵胎生进化的选择压力尚无定论,目前有3种关于卵胎生进化的假说受到学者的关注,其中母体操纵假说最受关注但尚未得到充分的检测。研究继1995年母体操纵假说提出之后,以栖息于温带气候环境下的卵生中国石龙子(Eumeces chinensis)为模型动物检测该假说。37条中国石龙子怀卵母体采自浙江丽水市郊。将怀卵母体分置于3个热处理中,其中12条母体提供每日14 h光照时间,13条母体提供10 h光照,其余12条母体没有任何调温机会(体温随室内环境温度而改变)。结果显示:怀卵母体选择体温向下漂移。3种处理下的雌体繁殖特征没有显著差异。长、短调温组下母体产卵时间要早于非调温组母体,但新生卵的胚胎历期没有显著差异。用5种热处理孵化卵,孵化温度分别为:1=室内波动温度孵化;2=27℃;3=24—30℃;4=22—32℃(3和4孵化处理中,孵化箱内的温度每隔1d改变1次,即卵分别在22和24℃孵化24 h,随后在32和30℃孵化24 h,每2d循环1次直至孵出);5=在实验室后院内模拟石龙子野外巢址孵化。结果显示:孵出幼体的体长、腹长和头部大小(头长和头宽)在3个母体热处理间存在显著差异,其他形态学特征不存在母体热处理间的显著差异;孵化温度以及孵化温度和母体热处理的交互作用对所有的幼体形态学特征均无显著影响。孵化温度以及孵化温度和母体热处理的交互作用对幼体疾跑速和最大持续运动距离无显著影响;但不同的母体热处理显著影响幼体疾跑速和最大持续运动距离。研究结果不仅为"热变异对在一定孵化温度范围内表型无显著变化的物种的幼体表型没有重要的修饰作用"这一假说提供了有力证据,并且支持母体操纵假说的两个主要预测:雌体在孕期通过体温漂变行为提供体内胚胎发育的最适热环境,而由母体调温行为诱导的后代表型的变异将增强后代的适合度。     

Thermoregulatory responses in seasonally acclimatized captive Southern White-faced Scops-owls

Seasonal thermoregulatory responses that are associated with cold tolerance have been reported for many species that inhabit regions where winters are severe (e.g. Holarctic), but relatively few studies have focused on species from regions where the climate is more unpredictable (e.g. Southern Africa). In this study, metabolic rate (VO₂) and body temperature (T_b) was measured during summer and winter in captive Southern White-faced Scops-owl (Ptilopsis granti), to test for thermoregulatory responses representing energy conservation in winter. During winter the Southern White-faced Scops-owls increased resting metabolic rate (RMR) by 45% to regulate a set point T_b—a result similar to what had been shown in small passerines from the Holarctic region. Increased RMR and increased conductance at cold T_a's are suggestive of improved cold tolerance. Basal metabolic rate (BMR) was 0.60 mL O₂ g⁻¹ h⁻¹ and showed no seasonal flexibility. Thus, contrary to expectation, the Southern White-faced Scops-owls showed seasonal thermoregulatory responses that are unlikely to represent energy conservation which was expected for a medium-sized bird inhabiting unpredictable climates in Southern Africa.     

白头鹎的代谢率与器官重量在季节驯化中的可塑性变化

动物能量代谢的生理生态特征与物种的分布和丰富度密切相关,基础代谢率(BMR)是恒温动物维持正常生理机能的最小产热速率,是动物在清醒时维持身体各项基本功能所需的最小能量值,是内温动物能量预算的重要组成部分.本研究测定了白头鹎(Pycnonotus sinensis)的BMR、内部器官(肝、心、肌胃、小肠、肾和整体消化道)和肌肉的重量,分析了白头鹎内部器官和肌肉重量的季节性变化及与BMR的关系.方差分析表明,白头鹎的BMR存在明显的季节性变化,冬季较高,夏季最低.其内部器官及肌肉重量的变化同样有明显的季节性.相关分析表明,白头鹎的BMR与肝、心、消化道等内部器官和肌肉重量存在明显的相关性.     

Energy metabolism and thermoregulation in pygmy lorises (Nycticebus pygmaeus) from Yunnan Daweishan Nature Reserve

The pygmy loris (Nycticebus pygmaeus) is a small prosimian living in Vietnam, Laos, eastern Cambodia and the south part of China. In China it is only found in Pingbian, Hekou, Jinping, Luchun of Yunnan. As N. pygmaeus is seriously threatened by hunting, trade and habitat destruction, it is listed in Appendix II of CITES, and in 2006 the IUCN classified it as “vulnerable”. In order to understand the characteristics of energy metabolism and thermoregulation of N. pygmaeus, the resting metabolic rate (RMR) and body temperature (T_b) at different ambient temperature (T_a) of pygmy lorises, as well as body mass, energy intake, digestable energy intake, digestability and the thermal conductance were measured in captivity. The results obtained mainly are as follows: (1) Pygmy loris feed dry food averaged 12.90 ± 1.02 g/d. They could gain 214.87 ± 16.65 kJ/d from food intake, and earned 200.15 ± 16.36 kJ digestable energy intake per day with 90.13 ± 1.34% of the digestability. (2) The T_b at room temperatures was a little low (35.23 ± 0.16 °C) and varied with T_a from 25 °C to 35 °C. There was a positive relationship between T_b and T_a, which was described as: T_b = 27.22 + 0.34T_a (r = 0.880). (3) The resting metabolic rate (RMR) of the pygmy loris was 0.3844 ± 0.0162 mlO₂/g/h, which was 51.91 ± 1.90% of the previous predicted rate by Kleiber (1961) [21]. (4) The average thermal conductance of the pygmy loris (N. pygmaeus) was 0.0449 ± 0.0031 mlO₂/g/h/°C. These characteristics of energy metabolism and thermoregulation of N. pygmaeus in Yunnan Daweishan Nature Reserve might be considered as the adaptive characteristics to their environment in tropical semi-evergreen forests and secondary forests.