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30 January 2025, Volume 45 Issue 1

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Catalogue of mammals in China (2024)

WEI Fuwen, YANG Qisen, WU Yi, JIANG Xuelong, LIU Shaoying, HU Yibo, GE Deyan, LI Baoguo, YANG Guang, LI Ming, ZHOU Jiang, LI Song, LI Sheng, YU Wenhua, CHEN Bingyao, ZHANG Zejun, ZHOU Caiquan, WU Shibao, ZHANG Li, CHEN Zhongzheng, CHEN Shunde, DENG Huaiqing, JIANG Tinglei, ZHANG Libiao, SHI Hongyan, LU Xueli, LI Quan, LIU Zhu, CUI Yaqian, LI Yuchun, HE Kai

2025, 45(1):  1-16.  DOI: 10.16829/j.slxb.151039

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Following the publication of the ‘Catalogue of mammals in China (2021)’and the release of Taxonomy and Distribution of Mammals in China, the mammalian catalogue of China has garnered significant attention from researchers worldwide. This surge in attention is primarily attributed to significant national research projects, such as the Second Qinghai-Xizang Plateau Scientific Expedition and Research Program, which have facilitated further exploration of China’s biodiversity and enhanced our understanding of the mammalian fauna. Between 2022 and 2024, a series of new mammalian species have been established, augmenting the 2022 catalogue by 42 species. In order to present a comprehensive overview of the valid mammalian taxa in China, we hereby present an updated edition of the ‘Catalogue of mammals in China (2024)’. The documentation conducted until the end of December 2024 revealed a total of 58 families, en-compassing 263 genera and comprising 736 valid mammalian species in China.

ORIGINAL PAPERS

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Spatial distribution and conservation implications of the type localities of mammals in China

GE Deyan, WEN Zhixin, LI Rui, LIU Qitao, YANG Qisen

2025, 45(1):  17-27.  DOI: 10.16829/j.slxb.150984

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The type locality of a species refers to the geographical location where the specimen was firstly collected and described. In biological taxonomy, phylogenetic reconstruction, and evolutionary studies, samples from the type locality are considered more reliable taxonomically and can better represent the current evolutionary state of the species compared to other geographic populations. Moreover, these samples are also important in research across various fields of biological science and in the preservation of genomic resources. Based on the analysis of the spatial distribution density and changes in the areas where type localities of valid species of extant mammals in China were discovered in different historical periods, this article collected the coordinate information on type localities for all of these valid species. The results show that approximately 42% of the valid species of extant mammals in China have their original type localities within the country. The Himalaya mountains, Hengduan Mountains, and Taiwan are regions where type localities of extant mammals in China are highly concentrated, with over 54% of the species (over 170 species) distributed in these areas. Analysis of the density of type localities of mammals discovered in different historical periods reveals that most species described in the early history were not within China. This is related to the development of taxonomy in Europe after the introduction of Linnaeus’s binomial nomenclature, as well as to the broader distribution of these species. The Himalayan Mountains, the Hengduan Mountains, and Taiwan have been areas where new species have been more concentratedly discovered since 1800, which is related to the presence of a large number of endemic and narrowly distributed species in these areas. Focusing on the regions where topotypes of extant mammals are concentrated and accelerating the collection and preservation of type specimens and high-quality genetic resources are of great significance for the protection and research of biodiversity in China, as well as for the training of the next generation of taxonomists.

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Population sizes and spatial distribution of Macaca mulatta brevicaudus in Hainan, China

CHEN Guangle, XING Lu, DU Wanru, REN Baoping

2025, 45(1):  28-35.  DOI: 10.16829/j.slxb.150889

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Rhesus macaque (Macaca mulatta) is a second-class protected species in China, and its subspecies the Hainan macaque (M. m. brevicaudus) is only found in Hainan Island, at the lowest latitude in China. Information on population size and spatial distribution of this subspecies dated from 30 years ago. It is urgent to obtain population size and distribution as the baseline information for the sake of conservation of Hainan macaques. From June 2021 to December 2023, a systematic survey of this subspecies was conducted via literature, questionnaire, and field investigation. We found 204 groups (5 100 individuals) of Hainan macaques inhabiting Hainan Island. These groups occupied less than 1% of the island area, mainly in the southern mountainous region. The average group size of the Hainan macaque is 25 ±11 individuals, which is the smallest among the six rhesus macaque subspecies in China. Eight isolated subpopulations of those 204 groups were identified based on their spatial distances and living status. Over the past 30 years, the distribution range of Hainan macaques has markedly shrunk, and the number of counties where the species has been recorded reduced from 18 to 10. About 96. 1% of the macaque groups are now restricted to nature reserves. We suggest those isolated groups should be paid more attention for the sake of conservation, especially the group in Wenchang. Strong human disturbance, habitat degradation, and habitat loss are the main threats to the survival of Hainan macaques.

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Mammals of Liupanshan National Nature Reserve in Ningxia

CHENG Xiaofu, MA Gang, HE Yijun, YIN Xiaohui, LI Cheng, LI Xinrang, MA Bocheng, GUAN Hongxin, ZHOU Hongwei, SONG Wenyu

2025, 45(1):  36-48.  DOI: 10.16829/j.slxb.150892

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During 2022 to 2023, we conducted a comprehensive investigation using various mammal survey methods to enhance knowledge and management of mammal diversity and distribution in the Ningxia Liupanshan National Nature Reserve. With 118 infrared cameras deployed over 45 430 camera working days, we recorded 20 mammal species belonging to 19 genera, 12 families, and five orders. Thirty small mammal survey plots were established, with a total of 7 425 trap-nights leading to the capture of 280 non-volant small mammals from 16 species belonging to 13 genera, seven families, and three orders. Using mist net and hand-netting methods, we registered a new record of a bat species within the reserve. Medium and large mammals such as Panthera pardus exhibited healthy populations with ample food resources and wide ranges. Through field surveys and literature compilation, a list of mammal species in this area was established totaling 64 species from 50 genera, 21 families, and six orders. Compared to previous lists, the present list has an increase of 17 mammal species, with 12 of which recorded from recent literature or image vouchers, and 5 from field survey of this study, including Erinaceus amurensis, Chimarrogale leander, Ochotona syrinx, Anourosorex squamipes, and Plecotus austriacus. The reserve houses approximately 75. 3% of all mammal species found in Ningxia, including a significant proportion (20. 3%) of carnivores and a large population of Panthera pardus. Moreover, this area serves as a pivotal zone where the Palaearctic and Oriental realms intersect, highlighting its conservation significance. The findings provide fundamental data for monitoring and conservation management efforts on mammalian resources within the Liupanshan National Nature Reserve.

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Population density, activity rhythm, and habitat utilization of hog badger Arctonyx albogularis in the Zhejiang Qingliangfeng National Nature Reserve

WANG Yang, LI Liang, ZHENG Nanzhong, YU Lin, TONG Genping, FU Wenyan, SONG Xiao, ZHAO Shanshan, XU Aichun

2025, 45(1):  49-59.  DOI: 10.16829/j.slxb.150854

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As higher-level consumers in the ecosystem, medium-sized carnivores plays an important role in the function-ality, structure, and stability of the ecosystem. The hog badger (Arctonyx albogularis) is a medium-sized carnivore belonging to the Mustelidae family and occupies a high position in the food chain of forest ecosystems in East China. However, there is limited research on its population density, daily activity rhythm, and habitat utilization characteristics. Our study represents a three-year investigation in the Zhejiang Qingliangfeng National Nature Reserve using infrared cameras. A total of 110 infrared cameras were deployed in the Qianqingtang and Longtangshan areas, accumulating 118 800 camera-days. Population density of hog badgers was estimated using the random encounter model (REM) and the daily activity rhythm of hog badgers and how it is affected by sympatric species was explored using the non-parametric kernel density index method. A single-species occupancy model was constructed to assess the habitat utilization characteristics of hog badgers. The results showed that the density of hog badgers in the Qingliangfeng National Nature Reserve was 23. 26 ±2. 97 individuals per square kilometer, exhibiting seasonal and interannual variations. The daily activity rhythm of hog badgers indicated nocturnal and crepuscular behavior, differing significantly from the activity rhythms of Masked palm civets (Paguma larvata) and Chinese ferret-badger (Melogale moschata), but overlapping significantly with that of Siberian weasel (Mustela sibirica). The occupancy model revealed that during the breeding season, hog badgers preferred to inhabit broad-leaved forests, coniferous forests, and mixed forests, with habitat utilization positively correlated with temperature. During the non-breeding season, hog badgers showed a preference for slopes with slight grades, locations closer to human settlements, and regions with higher temperatures. This study broadens the general understanding of the ecological features of wild hog badgers and the role of medium-sized carnivores in ecosystems, providing a scientific basis for their protection and management.

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Temporal and spatial distribution patterns of Chinese ferret-badger (Melogale moschata) and masked palm civet (Paguma larvata) in the same region of Lushan National Nature Reserve

SUN Jie, ZHANG Mi, LUO Wei, GU Hangmin, ZHANG Weiwei

2025, 45(1):  60-69.  DOI: 10.16829/j.slxb.150895

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Species with sympatric distributions typically undergo ecological niche differentiation to minimize competition and facilitate coexistence. This research explores the spatio-temporal distribution patterns of the Chinese ferret-bad-ger (Melogale moschata) and masked palm civet (Paguma larvata), their interactions with environmental factors, and the mechanisms behind their coexistence. We analyzed the daily activity rhythms and the degree of temporal niche overlap between these species within the Lushan National Nature Reserve using daily activity intensity index, kernel density estimation, and overlap index methods. The MaxEnt model was used to predict suitable habitats for both species and ecological niche overlap was assessed using ENMTools 1. 4. 4. We also compare their spatial niche preferences and investigate how environmental factors influence their coexistence strategies. The results reveal that both the Chinese ferret-badger and masked palm civet are predominantly nocturnal. They exhibit a high degree of overlap in their daily activity rhythms (Δ = 0.93, P > 0. 05), indicating that temporal niche differentiation does not significantly reduce competition between them. Suitable habitats for both species are primarily located along the forest edges at the mountain bases within the reserve, where there is substantial spatial overlap, although some differences exist. Both species show similar preferences for environmental factors, with altitude being the most significant, yet they differ in their preferences for vegetation types and slopes. These spatial differences in ecological niches may help mitigate competition for food resources and territories. This study underscores that the spatio-temporal distribution patterns of the Chinese ferret-badger and masked palm civet in the Lushan National Nature Reserve exhibit minimal niche differentiation. It is recommended to fully consider the environmental tolerance of both species, actively enhance habitat protection, and control human distur-bances to effectively conserve these and other sympatric species.

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Population structure and reproductive parameters of rhesus macaque in Wulingyuan National Scenic Spot

SHEN Chengye, YU Yang, LEI Shuyao, XIANG Zuofu

2025, 45(1):  70-77.  DOI: 10.16829/j.slxb.150853

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Among the six extant subspecies of macaques in China, the Fujian subspecies (Macaca mulatta littoralis) is one of the subspecies that has received relativeiy less attention. In this study, we investigated the population size, mating and birth seasons of rhesus macaque in the Wulingyuan area of Hunan Province through field tracking surveys based on individual identification. The results are as follows: (1) There are 18 monkey groups, with an average group size of 47. 78 ±22. 51, with ranging from 16 to 79 individuals. The overall sex ratio (male/female) is 1∶1. 57. The sex ratio for adult individuals is 1∶2. 47, for sub-adult individuals is 1∶3. 25, for adolescent individuals is 1∶1. 08, and for infant monkeys is 1∶1. 05. (2) Copulation events with ejaculation occur between October and February of the following year, with the highest mating frequency observed in November. (3) The distribution of birth events is from March to June, with a peak from April to May. The average birth date is 28th April, and the median birth date is 1st May with a standard deviation of 19. 11 days. These results indicate that the rhesus macaque in study area is seasonality and conforms to the pulse model; (4) The birth rate is 66. 27%, the mortality rate is 1. 80%, and the growth rate is 18. 09%. The results can provide basic data for research on the ecology, behaviour and conservation biology of macaques in Wulingyuan National Scenic Spot.

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The impact of anticoagulant rodenticides on the genetic diversity and anticoagulant resistance in the Norway rat population

ABUDUKEYOUMU Kader, SUN Ting, WANG Dawei, LIU Xiaohui, WANG Xiuhui, SONG Ying

2025, 45(1):  78-85.  DOI: 10.16829/j.slxb.150891

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To understand the genetic diversity and the level of anticoagulant resistance in Norway rats (Rattus norvegicus) in Kashgar region, Xinjiang after continuous use of anticoagulant rodenticides, we collected Norway rat samples after the application of the anticoagulant rodenticides in the year of 2019, 2020, 2022, and 2023. We first analyzed the different mitochondrial Cyt b haplotypes and their frequencies in different temporal populations, as well as the phylogenetic relationship among different haplotypes. Then, we analyzed the Vkorc1 variations and their frequencies in the temporal populations and assessed the anticoagulant level of the populations. The results revealed a total of six different Cyt b haplotypes in 100 rats, and they clustered into two groups in the haplotype network. The highest and lowest haplotype diversity was found in the population of 2019 and 2023, respectively, indicating an obvious bottleneck effect. Some low-frequency haplotypes disappeared in the populations of 2020, 2022, and 2023. Fst analysis suggested significant genetic differentiation between the population of 2022 and other temporal populations. Analysis of the Vkorc1 gene revealed one silent mutation Ile82Ile and no resistance-associated Vkorc1 missense mutations were found, indicating a low level of resistance in the local populations. All these findings suggest that the rat control in the entire village has led to a decrease in genetic diversity and significantly affected the genetic structure of the population. However, the rat populations in the studied area have not developed anticoagulant resistance yet, and the first-generation anticoagulant rodenticides are still effective. This study provides a reference for further research on the population structure of Norway rats and also provides theoretical guidance for the use of anticoagulant rodenticides in local Norway rat populations.

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Relationship between the level of blood hormones and the shedding and regeneration of male Père David’s deer (Elaphurus davidianus) antlers

MENG Qinghui, XIA Zhiqiang, SHAN Yunfang, LI Junfang, HE Yonghui, CUI Yanhong, HU Huagang, XIAO Xuan, BAI Jiade, ZHONG Zhenyu, MENG Xiuxiang, ZHANG Fuxian, BAI Chao, ZHANG Chenglin

2025, 45(1):  86-95.  DOI: 10.16829/j.slxb.150791

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The antlers of male Père David’s deer (Elaphurus davidianus) were shed during the winter solstice, and hormones play a crucial role in the regeneration of antler cartilage. This study collected blood samples from male Père David’s deer (Elaphurus davidianus) to measure hormone levels, comparing pre-winter solstice (November-December) and post-winter solstice (January-March) periods, as well as analyzing the fluctuations during the antler growth period (Winter Solstice-Start of Insects Awakening, December to March of the following year). The aim is to provide further evidence for the relationship between antler cartilage regeneration and hormone levels. Results showed that blood estradiol (E₂), testosterone (T), growth hormone (GH), insulin-like growth factor-1 (IGF-1), and melatonin (MT) levels exhibited significant annual rhythmic changes (F_11,486= 3. 98, n = 507, P = 0.004). Blood parathyroid hormone (PTH) levels showed synchronous monthly changes with antler ossification (F_11,486= 1. 72, n = 507, P = 0.081). After the winter solstice, blood E₂, GH, T, and PTH levels in deer antlers began to gradually increase, while IGF-1 levels increased later. Throughout the antler growth period, blood IGF-1, MT, GH, T, and E2 levels were significantly positively correlated with the regeneration speed of deer antlers, while blood PTH levels were significantly positively correlated with antler weight gain. During the Major Snow solar term (early December), the content of MT, GH, and IGF-1 in the blood began to increase, and the content of E₂ and T was at the second peak of the year, which was extremely significantly different from the content of E₂ and T during the breeding season of male deer (F7,152 = 4. 08, n = 160, P = 0.003). During the winter solstice, the slow growth phase Ⅲ and the rapid growth phase Ⅳ of the antlers, the content of MT, GH, and IGF-1 in the blood of deer antlers increased sharply (F_3,80 = 4. 69, n = 84, P = 0.002); the content of E₂ first decreased and then increased, with significant differences (F_3,80 = 3. 14, n = 84, P = 0.045), while the content of T decreased from (0. 696 9 ±0. 09) ng/mL (n = 84) to (0. 664 7 ±0. 06) ng/mL (n = 84), with no significant difference (F_3,80 = 5. 79, P = 0.178). After the Rainwater solar term (late February), during the stagnation and ossification phase V of antler growth, the content of GH in the blood decreased significantly, with significant differences (F_3,80 = 3. 25, n = 84, P = 0.037), while the content of E₂ and T continued to increase. PTH levels continued to rise before shedding and throughout the antler growth period, increasing from (11. 78 ±2. 54) pg/mL (n = 22) during the Major Snow solar term to (69. 23 ±7. 89) pg/mL (n = 22) during the Vernal Equinox solar term, with extremely significant differences (F_1,20 = 4. 44, P = 0.001). In conclusion, E₂, MT, GH, IGF-1, T, and PTH are all involved in the entire process of winter antler regeneration in deer antlers. Low physiological concentrations of melatonin and testosterone during the winter solstice are directly related to antler shedding. This study can provide a basis for improving the breeding conditions of male deer in winter and contribute to research on the regulation of cartilage proliferation by hormones and the diverse mechanisms of antler regeneration.

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Distribution characteristics of mineral elements in habitat soil-food-feces continuum of giant pandas

SHU Yan, HUANG Yuling, LIU Yuan, ZHU Dapeng, LIU Kun, TAN Jinchao, WANG Fengmei, FANG Tingting, YUAN Shibin, WANG Le

2025, 45(1):  96-107.  DOI: 10.16829/j.slxb.150852

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To investigate the distribution characteristics of mineral elements in the habitat soil-food-faeces continuum of wild giant pandas, we collected samples of soil, dietary bamboos, and feces from the core area of Foping National Nature Reserve in Shaanxi Province in the year of 2022. All samples were analyzed for the absolute and relative contents of nine nutritionally essential minerals: Fe, Mn, Zn, Cu, K, Ca, Mg, P, and Na. The results showed that the absolute content of mineral elements in soil-food-feces was not entirely consistent. The elements with high absolute content in the soil were not always rich in the food bamboos and feces, and minerals that were of less absolute abundance in the soil were not always low in the food and feces. Minerals in soil and bamboo leaves showed a close positive or negative correlation, whereas elements in the feces showed significant positive correlations without detected negative relevance. After the whole process of ingestion, digestion, and absorption of bamboo leaves by giant pandas, the relative contents of most minerals changed in the feces. The feces showed lower relative concentration of P and Na than in the food when feeding on Bashania fargesii; whereas there was no significant difference in the content of Ca, Mg, K, Mn, Cu, and Zn. Furthermore, Fe was significantly higher in feces than in food. When feeding on Fargesia qinlingensis, there was no significant difference in the relative content of Mn, the relative content of Fe in feces was significantly higher than in food, and the relative contents of the other seven elements in feces were significantly lower than in food. Therefore, the contents of mineral elements in soil-food-feces were not completely consistent. When giant pandas fed on different bamboo leaves of different species, their utilization rates of different mineral elements varied. Among the 9 essential elements, Fe had the lowest absorption rate. Our results illustrated the distribution characteristics of mineral elements in the habitat soilfood-feces continuum and the differences in the utilization of 9 essential mineral elements by giant pandas, which help us further understand the nutritional strategies of wild giant pandas and provide references for the mineral nutrition regulation on giant pandas in captive settings.

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Evaluation of aboveground biomass of edible bamboo for giant panda in the Yingjing area of Giant Panda National Park

QIN Weirui, LIU Jingyi, SONG Xinqiang, FU Mingxia, PAN Han, CHENG Yong, ZHOU Caiquan, YANG Biao

2025, 45(1):  108-118.  DOI: 10.16829/j.slxb.150841

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Evaluating food resources and assessing habitat quality for giant pandas (Ailuropoda melanoleuca) requires estimating the aboveground biomass of edible bamboo. In the Yingjing area of the Giant Panda National Park (GPNP), we conducted an investigation into the aboveground biomass of different bamboo species across various forest types. This involved employing sampling and harvesting methods to preliminarily estimate the aboveground biomass of bamboo suitable for giant pandas. The findings revealed that aboveground biomass of edible bamboo for giant pandas did not exhibit significant differences across different forest types. However, there was a considerable disparity in the aboveground biomass among various edible bamboo species. Notably, the majority of the aboveground biomass followed the pattern of branches < leaves < stems. In the Yingjing area of the GPNP, the distribution area of giant panda edible bamboo, including Chimonobambusa szechuanensis, Yushania brevipaniculata, Bashania faberi, and Qiongzhuea multigemmia, was as follows: 20 770. 44 hm², 31 549. 34 hm², 30 714. 25 hm², and 23 340. 65 hm², respectively. The total aboveground biomass for each bamboo species was 640 543. 67 t, 177 086. 42 t, 344 601. 63 t, and 152 888. 26 t. Additionally, the energy provided by each square meter of stems and leaves was 16 246. 17 kJ, 3 157. 89 kJ, 7 786. 47 kJ, and 2 814. 41 kJ, respectively. From the perspective of the aboveground biomass of edible bamboo, it can be concluded that, within a specific space and time range, there is no shortage of food source bamboo supply in the study area. The result of this study can offer valuable insights for devising scientifically informed protection and management measures for giant pandas and their habitat.

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Feeding on human-planted eucalyptus by the white-headed langur (Trachypithecus leucocephalus) in the wild and its nutritional analysis

LIU Yanchen, GUO Qiuyan, LU Meijing, WU Shijun, NONG Dengpan, HUANG Chengming, FAN Penglai, ZHOU Qihai

2025, 45(1):  119-127.  DOI: 10.16829/j.slxb.150883

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Few mammals feed on eucalyptus and wild white-headed langurs (Trachypithecus leucocephalus) have been found eating parts of this poisonous trees. From August 2021 to July 2022, data on the feeding on cultivated eucalyptus (Eucalyptus urophylla×E. grandis) were collected systematically from one free-ranging group of white-headed langurs in the Chongzuo White-headed Langur National Nature Reserve, Guangxi. We explored possible reasons why the whiteheaded langur feeds on eucalyptus by nutritional analysis. The results showed that white-headed langurs fed on barks, mature leaves, and young leaves of eucalyptus, accounting for 70. 4%, 28. 1% and 1. 4% of the total foraging time of eu-calyptus respectively. The mean durations of feeding bouts on barks and mature leaves were significant longer than that for young leaves. The frequencies of feeding bouts on bark and mature leaves were significantly higher in the rainy season than those in the dry season. The frequencies and mean duration of feeding bouts on barks, mature leaves, and young leaves of eucalyptus did not differ among ages of individuals. Compared to the young leaves of eucalyptus and the other major food plant species consumed by white-headed langurs, eucalyptus barks contain higher content of calcium, which indicates that the acquisition of mineral elements may be one of reasons to select eucalyptus as food.

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Changes in body weight of hand-reared chimpanzee infants (Pan troglodytes verus)

YANG Xinyi, FAN Baodan, LI Ruizi, SHI Jian, HAN Jingjing, LI Jing, ZHANG Yuanyuan, ZHANG Shu, CAO Yue, YAO Lin, ZHOU Na, LIU Xuefeng, LIU Dingzhen

2025, 45(1):  128-136.  DOI: 10.16829/j.slxb.150874

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Post-parturition infant rejection is common in captive chimpanzee. When this occurs, an immediate hand-rearing is required. Monitoring and observing the infant’s development and health status are essential for successful handrearing. Body weight serves as an important index of an individual’s growth and development and has been widely used in several mammalian species. However, there is currently no literature available on the changes in daily body weight gain in newborn chimpanzee infants. This study aimed to address this gap by monitoring and recording the changes in body weight during the first 12 months of life of six mother-rejected captive-born chimpanzee infants (2 females and 4 males) between 1999 and 2015 at Beijing Zoo and Dalian Forest Zoo. A linear regression model of weight-age was established based on the body weight data. A generalized linear model was then used to analyze the relationship between daily body weight gain and relative daily body weight gain and age by day, as well as to compare the two variables between two genders. The results showed that the body weight of chimpanzee infants increased linearly with age by day (y = 0.015 x + 2.139, R² = 0.830, P = 0.000 1), with females being significantly heavier than males of the same age by day (F = 10. 175, P < 0. 001). Both daily body weight gain and relative body weight gain decreased significantly with age by day (r = -0. 667, P = 0.035 and r = -0. 224, P = 0.042, respectively). The daily body weight gain and relative daily body weight gain in female infants were remarkably larger than those of males until they reached the ages of 235 and 184 days, respectively. Afterward, these two variables in female infants became smaller than those in male infants. Our results demonstrate sexual dimorphism in body weight, daily body weight gain and relative daily body weight gain of chimpanzees aged 0-12 months. These findings may provide a theoretical basis for increasing the survival rate of infants and enlarging the population size of chimpanzees in captivity.