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Downloaded from Brill.Com10/02/2021 03:41:10AM Via Free Access 66 Xiong Et Al Contributions to Zoology, 78 (2) 65-75 (2009) Food limitation and low-density populations of sympatric hamster species in North China Wenhua Xiong1, 2, Hong Liang1, 2, Xiaohui Yu1, 2, Lin Cong3, Zhibin Zhang1, 4 1State Key Laboratory of Integrated Management of Pest Insects and Rodents in Agriculture, Institute of Zoology, Chinese Academy of Sciences, Datun Road, Beijing 100101, China 2Graduate School of the Chinese Academy of Sciences, Beijing 100049, China 3Institute of Plant Protection, Heilongjiang Academy of Agricultural Sciences, China 4Email: [email protected] Key words: body condition, Cricetulus barabensis, density-dependent food restriction, population dynamics, Tscherskia triton Abstract Statistical analyses ..................................................................................... 68 Results .................................................................................................................... 68 Although food has been proposed as a possible factor on population Annual population abundances ............................................................ 68 regulation of small mammals, evidence of food restriction in field Laboratory food restriction experiment ............................................ 68 conditions is still lacking. Food restriction is generally thought to Muscle protein concentration ............................................................... 68 occur in high-density years of oscillating populations of small mam- Cleaned carcass index .............................................................................. 70 mals. By using two body condition indices (muscle protein concen- Comparisons between laboratory and field studies ..................... 70 tration and cleaned carcass index), we investigated variations of Discussion ............................................................................................................. 72 body condition of the greater long-tailed hamster (Tscherskia triton) Yearly variation of body condition ..................................................... 72 and the striped hamster (Cricetulus barabensis) in the Raoyang Seasonal variation of body conditions .............................................. 72 County of the North China Plain during a population decline- and Laboratory food restriction effect ....................................................... 73 low-phase from 1998 to 2003. Our laboratory experiments showed Implications of food restriction ............................................................ 73 that food restriction reduced muscle protein concentration and Acknowledgements .......................................................................................... 73 cleaned carcass index of greater long-tailed hamster. We found there References ............................................................................................................ 73 were significant differences in body condition of hamsters over the six years. Both the muscle protein concentration and cleaned carcass index of T. triton were significantly and positively correlated with its Introduction yearly trap successes, which reflected yearly population densities. Muscle protein concentration of C. barabensis was positively cor- Food availability has been emphasized as a major lim- related with the total yield of the wheat. This observation supported iting factor that influences population variations of the hypothesis on the in-phase relation between food resources or most vertebrates (Cameron and Eshelman, 1996; Huitu availability and rodent abundances. During the four seasons, muscle et al., 2007). Food per animal may determine body protein concentration of two species was highest in autumn, and lowest in summer. These two body condition indices of wild T. triton condition related to food restriction. Some previous in summer and in low phases were similar to 60% food restricted laboratory studies have shown that food restriction has animals raised in the laboratory, but significantly lower than non- remarkable effects on the physiological condition, re- food-restricted animals. Our results indicated that malnutrition oc- production and immune level of immunity of rats curred in summer when food was not abundant, and in low density (Sakamoto et al., 2000). For example, Liang and Zhang years with low grain production (an index reflecting food abundance (2006) demonstrated that food restriction during the of rodents) in our study region. Therefore, we concluded that food pregnancy period of the greater long-tailed hamster limitation may play an important role in seasonal and inter-annual negatively affected the body growth of F1 offspring, population fluctuation of these two hamster species. and delayed the physical development and neurode- velopment of both F1 and F2 offspring. However, evi- Contents dence of food impact in fluctuating wild populations is still lacking, and thus the underlying mechanisms are Introduction .......................................................................................................... 65 not fully understood (Lindell, 1997; Norrdahl and Materials and methods .................................................................................... 66 Study sites and trapping procedures .................................................. 66 Korpimäki, 2002; Ramsey et al., 2002). Measurement of body condition .......................................................... 67 Many population densities of small mammals expe- Laboratory food restriction on the greater long-tailed hamster 67 rience fluctuations in demography (Boonstra, 1994; Downloaded from Brill.com10/02/2021 03:41:10AM via free access 66 Xiong et al. – Food limitation and low-density populations of sympatric hamster species in North China Krebs et al., 1995; Norrdahl and Korpimäki, 1995; pests across the Northern China Plain where wheat, Krebs, 1996; Tkadle and Zejda, 1998). There are three corn and peanut are the main crops. Most crops are possible phase patterns between food and population harvested in autumn (from August to October), except density: in-phase, out-of-phase and phase with a time- wheat, which is harvested in spring (from March to lag (Krebs and Myers, 1974). The time-lag reflects the May). Crop seeds comprise over 90% of the diets of ability of animal population tracking changing food re- these two rodent species in autumn, and a large number source (Mihok and Boonstra, 1992; Boonstra et al., of crop seeds is stored in their burrows for over-win- 1998a; Norrdahl and Korpimäki, 2002). If the time lag tering (Wang et al., 1992, 1996). Population abun- is close to zero or very small, food availability should dance of both rodent species oscillates greatly among be higher in the increase phase or high-density phase of seasons and years and populations of both species animal populations, and lower in the decline phase or change slowly over the years. Population density of T. low-density phase. The out-of-phase relation is gener- triton generally reaches a peak in autumn, while that of ally accepted by ecologists because food availability C. barabensis generally peaks in spring, and both spe- would decrease with an increase of population density cies exhibit an approximate 10-year cycle in popula- and it would become lowest when population reaches tion size (Zhang et al., 1998). Although the behaviour, peak phase. Food shortage is usually thought to occur in reproduction, genetics and ecology of the two hamsters high-density years of oscillating populations of small have been well-documented (Wang et al., 1996; Zhang mammals (Boonstra, 1994). Andersson and Jonasson et al., 1998; Xie and Zhang, 2006), the mechanism un- (1986) found a negative correlation between the density derlying their population dynamics remains unclear. of rodents and frequency of food plants during two four- In this study, we investigated the seasonal and yearly year cycles on alpine heath. The out-of-phase relation variation of muscle protein content and cleaned car- may be more likely for more oscillating animals and cass index of the two hamster species during a decline their food resources. For example, many tree species and low phase of their populations from 1998 to 2003 produce massive crops of large seeds in low-density in the croplands in the Raoyang County of the North years of small rodents (Willson and Traveset, 2000; China Plain using field trapped animals. We wanted to Jansen et al., 2004; Xiao et al., 2005; Li and Zhang, test the season- or density-dependent food restriction 2007), which often results in “predator satiation” and hypothesis. We also compared muscle protein content benefits forest regeneration (Janzen, 1970). However, in and cleaned carcass index of wild animals with lab- theory, if animals could track fluctuations of food re- raised animals. We predicted that 1) food restriction sources more quickly, an in-phase relation is also pos- would occur in low-density years of hamsters if the sible. The in-phase relation is likely possible when in-phase relation hypothesis holds; 2) food restriction populations of animals and their food resources change would occur in summer, as high temperature plus slowly. poor food resources may reduce body condition of Body weight and body composition (such as muscle hamsters. protein content and muscle fat content) are generally affected by food and nutrition, and they are feasible to evaluate (Johns et al., 1984; Munoz et al., 1994; Lynch Materials and methods and Barbano, 1999). For
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