Dietary Habits of Wintering Pale Thrush Turdus Pallidus in the Urban Green Environment of Subtropical Okinawa-Jima Island, Japan
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Ornithol Sci 18: 205 – 213 (2019) SHORT COMMUNICATION Dietary habits of wintering Pale Thrush Turdus pallidus in the urban green environment of subtropical Okinawa-jima Island, Japan Shun KOBAYASHI1,#,*, Wakako YOSHIDA1, Hiroaki UI2, Yoko OKAWARA2 and Masako IZAWA1 1 Faculty of Science, University of the Ryukyus, 1 Senbaru, Nishihara, Okinawa 903–0213, Japan 2 Graduate School of Engineering and Science, University of the Ryukyus, 1 Senbaru, Nishihara, Okinawa 903–0213, Japan ORNITHOLOGICAL Abstract Knowledge of the feeding behavior of omnivorous migrant birds is impor- tant to understand their migration ecology. This study revealed the dietary habits of SCIENCE the Pale Thrush Turdus pallidus overwintering on Okinawa-jima Island, Japan. Food © The Ornithological Society items consumed were determined from the contents of the gizzards of 32 birds; at of Japan 2019 least 24 animal and eight plant species were identified. The contents of 84.4% of the studied gizzards were dominated by a single species or item. The Pale Thrush for- ages on various animals and plants, but in its winter habitat it feeds extensively and exclusively on select food items within a short period. Key words Food habit, Ryukyu Archipelago, Turdus pallidus, Urban, Winter bird Approximately 15% of bird species are migratory Tokyo, Japan, the Pale Thrush fed only on fruits, (Kirby et al. 2008), migrating between breeding and although the sample size of observed birds was small. non-breeding ranges in order to maximize energy These observations suggest that the diet of this spe- efficiency (Somveille et al. 2018). However, changes cies varies depending on its winter habitat. in food resources between years can cause annual In order to understand dietary habits, not only food fluctuations in migratory ranges (Newton 2008). Fur- species but also the amount of each species con- thermore, feeding habits can differ among individuals sumed is important. However, previous studies did in some omnivorous birds and individual specializa- not determine the frequency of occurrence or volume tion has been observed in some species (reviewed by of each food item and gizzard content sample sizes Bolnik et al. 2003 and Araújo et al. 2011). Therefore, were too small for analysis. Furthermore, in order to a detailed study of the feeding behavior of omnivo- understand the plasticity of the dietary habits of the rous migrant birds is of interest. species within its broad range, knowledge of its diet The Pale Thrush Turdus pallidus (Passeriformes: in the southern subtropical region where it winters Turdidae) has a body length of 24 cm (Takano 2015). is also desirable. Therefore, in the present study we It is widely distributed across East Asia. It breeds in used gizzard content analysis to clarify the feeding summer in northeast China and Russia and winters habits of the Pale Thrush in a subtropical urban green in southern and western Japan, Korea, and southern environment. China (Takano 2015). Previous studies of droppings have shown that this species is omnivorous, with MATERIALS AND METHODS its winter diet including animals such as arthropods and fruits from various plants growing in temperate 1) Study site Kyushu, Japan (Seki 1998). In contrast, Hamao et al. Samples were collected at the University of the (2009) observed that in the urban parks of temperate Ryukyus (Senbaru Campus), which is located in central Okinawa-jima Island (26°14–15′N, 127°45– (Received 26 August 2018; Accepted 17 February 2019) 46′E), the Ryukyu Archipelago, Japan. The campus # Corresponding author, E-mail: [email protected] * Present address: Faculty of Science, University of the Ryukyus, 1 has an area of 100 ha; it is surrounded by human Senbaru, Nishihara, Okinawa 903–0213, Japan residences, contains some scattered buildings, grass- 205 S. KOBAYASHI et al. land, and trees, and in addition to a 2 ha pond, has DISCUSSION 20 ha of secondary forest, and a 16 ha agricultural experimental field. Pale Thrushes inhabit this area The Pale Thrush was found to feed on a range of each winter from November to April. animal and plant species in winter in the subtropical urban green area. Previous studies have indicated 2) Gizzard contents analysis that Pale Thrushes mainly feed on plants in the tem- We collected dead Pale Thrush specimens from perate urban area (Hamao et al. 2009), or a combi- 2000 to 2017. The main cause of mortality was colli- nation of animal and plant food items at some sites sion with windows. Birds (N=38) were dissected and (Uchida 1913; Chiba et al. 1972; Seki 1998). As the the gizzards extracted and stored in 70% ethanol for abundance of arthropods decreases during winter analysis. The gizzard contents were washed through in urban areas (Galeotti et al. 1991), the probable a 1 mm sieve and the items remaining in the residue reason why thrushes were not observed feeding on were identified wherever possible to species level; animal matter in the temperate urban area was that when we could not do so, we used the label “item.” the abundance of arthropods was low. Our results, The wet weight of each species or item was mea- together with those of previous studies, indicate that sured to an accuracy of 1 mg using a microbalance the Pale Thrush is flexible in its feeding habits and (PG503-S; Mettler Toledo, Switzerland). changes according to food resource availability in their winter habitat. 3) Data analysis Although Pale Thrushes are omnivorous and utilize The frequency of occurrence (FO) of each spe- various food resources, our study showed that single cies or item was calculated, and a binomial test was food species or items tend to dominate the gizzard. In used to compare wet weight among species or items. general, birds have a short digestion time (Ziswiler & Statistical analysis was performed in R ver. 3.4.0 (R Farner 1972; McKechnie 2010). Our results indicate Core Team 2017). that the Pale Thrush intensively consumes specific foods over a short period. Uchida (1913) also showed that gizzard contents were dominated by one to four RESULTS items in this species (N=5). In addition, among Six of the 38 gizzard samples (15.8%) were empty various other omnivorous Turdidae gizzards were and were excluded from the analyses. dominated by one item in 51.4% (N=177) of Dusky From the gizzard samples (N=32), 24 animal Thrushes T. naumanni eunomus wintering in Tokyo, and eight plant species were identified; many items in 56.5% (N=62) of Brown-headed Thrushes T. remained unidentified (Table 1). The mean number chrysolaus resident in Tokyo, and in 37.0% (N=27) of items contained in a single gizzard was 3.4±2.1 of White’s Thrushes Zoothera dauma aurea resident (animal: 2.2±1.9; plant: 1.2±0.6) (N=32, mean±SD). in Tokyo (Uchida 1913). Although the study method Animal species from Gastropoda (FO=53.1%), in the present study differed from that in previous Diplopoda (53.1%), and Insecta (37.5%) frequently studies, the migratory Turdidae may share similar appeared, as did the plant species Bischofia javanica feeding behavior. (fruit/seed; 40.6%) (Table 1). Grit, parasites, and Intra- and interspecific competition, ecological plastics were also observed (Table 1). opportunity, and predation are ecological and evolu- Among the 32 samples analyzed, mean wet weight tionary factors that influence variation in individual was 689.7±802.1 (12–2945) mg. The contents of 10 diets (Araújo et al. 2011). We did not determine what (31.3%) gizzards were dominated (>80% of total wet influences the diet of the Pale Thrush among these weight) by animal species, and 17 (53.1%) gizzards factors. Further research, including identification of were dominated by plant matter (Table 2). Overall, age and sex, study in natural habitat, and long-term the contents of 13 gizzards (40.6%) were found to be surveys, are needed to elucidate the factors deter- dominated by animal food items, while others were mining individual diet variation in omnivorous birds dominated by plant species (binomial test; P=0.38). such as the Pale Thrush so as to understand their The contents of 27 (84.4%) gizzards were dominated effects on migration. (more than 80%) by a single animal item or plant species (Appendix 1 and 2). 206 Dietary habits of Turdus pallidus Table 1. Frequency of occurrence (FO) of each item. Order Family Species name Stage/Part FO Animal matters Insecta Hymenoptera Formicidae Pheidole sp. Adult 3.1 Monomorium intrudens Adult 3.1 Ochetellus glaber Adult 3.1 Tetramorium bicarinatum Adult 3.1 Tetramorium sp. Adult 3.1 Anoplolepis gracilipes Adult 6.3 Blattaria Blattellidae Adult 3.1 Coleoptera Unknown – Adult 12.5 Unknown – Larva 3.1 Lepidoptera – Lepidoptera sp. Larva 3.1 Diptera – Diptera sp. Larva 6.3 Psocodea Unknown – Adult 3.1 Unknown – – Adult 12.5 – – Larva 3.1 Insecta Total 37.5 Arachnida Araneae – Unknown Unknown 6.3 Acari – Unknown Adult 3.1 Arachnida Total 6.3 Diplopoda Polydesmida Paradoxosomatidae Unknown Unknown 28.1 Julida Unknown – Unknown 15.6 Unknown – – Unknown 28.1 Diplopoda Total 53.1 Crustacea Isopoda Armadillidae Unknown Adult 3.1 Isopoda Unknown – Larva 3.1 Crustacea Total 6.3 Gastropoda Stylommatophora Bradybaenidae Bradybaena sp. Unknown 6.3 Acusta despecta despecta Unknown 9.4 Unknown – 12.5 Subulinidae Allopeas kyotoense Adult 6.3 Mollusca Achatina fulica Larva 3.1 Unknown – – 3.1 Mesogastropoda Cyclophoridae Cyclophorus turgidus Larva 21.9 Gastropoda Total 53.1 Animal Total 78.1 Plant matters Laurales Lauraceae Machilus thunbergii Fruit/seed 3.1 Poales Poaceae Poaceae spp. Seed 6.3 Malpighiales Phyllanthaceae Bischofia javanica Fruit/seed 40.6 Clusiaceae Garcinia subelliptica Fruit/seed 3.1 Oxalidales Elaeocarpaceae Elaeocarpus zollingeri Fruit/seed 3.1 Rosales Moraceae Ficus sp.