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Effects of Dietary Differences Between Sympatric Japanese Serow and Sika Deer on Environmental Reconstruction by Means of Mesowear Analysis

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Effects of Dietary Differences Between Sympatric Japanese Serow and Sika Deer on Environmental Reconstruction by Means of Mesowear Analysis Ann. Zool. Fennici 50: 200–208 ISSN 0003-455X (print), ISSN 1797-2450 (online) Helsinki 30 August 2013 © Finnish Zoological and Botanical Publishing Board 2013 Effects of dietary differences between sympatric Japanese serow and sika deer on environmental reconstruction by means of mesowear analysis Eisuke Yamada Department of Natural Science, Graduate School of Science and Engineering, Kagoshima University, 1-21-35, Korimoto, Kagoshima, 890-0065, Japan (e-mail: [email protected]) Received 5 Nov. 2012, final version received 16 Feb. 2013, accepted 1 Mar. 2013 Yamada, E. 2013: Effects of dietary differences between sympatric Japanese serow and sika deer on environmental reconstruction by means of mesowear analysis. — Ann. Zool. Fennici 50: 200–208. Diet reconstruction using mesowear analysis has mainly been applied to extinct spe- cies and their paleoenvironments. Little is known regarding the effects of dietary dif- ferences on sympatric environments using this analysis and the limited existing knowl- edge from extant species may introduce errors when applied to fossil assemblages. I aimed to determine the sensitivity of mesowear analysis using extant ungulates with known diets. An interspecific comparison was conducted using wild populations of Japanese serow (Capricornis crispus, n = 37) and sika deer (Cervus nippon, n = 55) living in deciduous broad-leaved forest of the Nikko National Park, central Japan. One of the mesowear variables differed significantly between the two species (Fisher’s exact test: p < 0.05). According to hierarchical cluster and principal component analy- ses, Japanese serow were classified as browsers, while sika deer as mixed feeders. Pre- vious studies support these results; therefore, mesowear analysis can be used to detect dietary differences in sympatric species. Introduction Intermediate feeders have fewer environmental constraints with regard to habitat preferences Paleontologists often refer to guilds of herbivo- (Kingdon 1997). rous mammals to reconstruct paleoenvironments The physical properties of food habits are since their feeding preferences reflect the vegeta- also reflected in tooth wear. Some very gen- tion resources available in specific environments eral habitat-driven mechanisms have increased (Vrba 1985, Plummer & Bishop 1994, Fortelius the abrasiveness of food available to herbiv- et al. 1996). The relationship between food ores; for example, more grit and dust indicates habits and environment of herbivore ungulates coarser vegetation in drier environments due has been determined as follows: grazers indicate to increased water stress on plants (Kaiser & the presence of either open areas or open patches Rössner 2007). Mesowear analysis is a method in closed woody areas, while browsers indi- of dietary reconstruction based on occlusal wear cate the presence of more forested environments of cheek teeth facets of extant species with or bushy/shrubby patches in open landscapes. known diet and habitat preferences (Fortelius ANN. Zool. FENNIcI Vol. 50 • Effects of dietary differences based on mesowear analysis 201 & Solounias 2000). This method is based on visual observation and considers a large number of fossil teeth specimens over a short period of time. For hypsodont and mesodont species, mes- owear appears to be stable throughout much of the adult life (Fortelius & Solounias 2000, Rivals et al. 2007). In addition, Yamada (2012) showed that teeth have consistent diet-specific mesowear Fig. 1. occlusal relief was measured as the vertical dis- patterns. Therefore, mesowear analysis has been tance between a line connecting two adjacent cusp tips applied as a tool to reconstruct the paleodiet of and the valley bottom between them (marked 1 in the herbivorous mammals from several localities, figure) divided by the length of the whole tooth (marked from which the type of forage available could be 2 in the figure). inferred, and indirectly, the paleoenvironment of the locality (Kaiser 2003). Mesowear analysis indicates diet based on no rounded area between the mesial and distal two morphological variables of the buccal cusps facets. A round cusp has a distinct round tip of the upper second molar (M2): occlusal relief, without planar facet wear but with retained facets and cusp shape. Occlusal relief is classified as on the lower slopes (Fig. 2). A blunt cusp lacks “high” or “low” depending on how high the distinct facets altogether (Fortelius & Solounias buccal cusps are raised above the valley between 2000). Browsers tend to have a sharp cusp, and them. Occlusal relief measurements are con- grazers, which consume a more abrasive diet, ducted as follows: First, the vertical distance tend to have rounded or blunted cusps. between a line connecting two adjacent cusp tips However, this relatively new assessment and the valley bottom between them is meas- method has a few problems, one of which is the ured (Fig. 1). Second, the distance is divided by interpretation of results for the large mammal the entire tooth length. The cut-off point for the fossil assemblage horizon. Several studies on selenodont molar is 0.1 (low < 0.1, high ≥ 0.1, extinct sympatric species (MacFadden 2000, Fortelius & Solounias 2000). Relatively “high” Calandra et al. 2008) suggested they have dif- occlusal relief is typical for taxa that browse ferent diet. For example, Blondel et al. (2010) more, whereas “low” occlusal relief is typical for applied mesowear analysis and reported that fossil taxa that tend to graze more (Fortelius & Solou- bovids from the late Miocene sediments of Toros- nias 2000). Cusp shape is classified as “sharp,” Menalla (Chad) had different mesowear patterns “round,” or “blunt.” A sharp cusp has practically although this mammalian assemblage inhabited Fig. 2. Molars of two ungulates (buccal view of left M2, teeth not to scale). (a) Japanese serow (high occlusal relief and sharp cusp); (b) sika deer (high occlusal relief and round cusp). 202 Yamada • ANN. Zool. FENNIcI Vol. 50 135°E 145°E 135°E 145°E a b 40°N 40°N The Nikko NP The Nikko NP 30°N 30°N 400 km 400 km Fig. 3. Distribution of (a) Japanese serow and (b) sika deer between 1978 and 2003 (modified from Biodiversity center of Japan 2004), and the locality of the Nikko National Park (NP, Tochigi Pref, Japan). the same area (Le Fur et al. 2009). For investi- many peaks over 2000 m (the highest is 2578 m) gating the paleoenvironment, it is necessary to a.s.l. The timberline is from 2400 to 2500 m a.s.l. determine how mesowear exactly reflects the die- The areas above 1500–1600 m a.s.l. are covered tary segregation. Although Fortelius and Solou- by sub-alpine coniferous forests. Broad-leaved nias (2000) showed that mesowear replicates the forests grow below 1600 m a.s.l. In many areas, known local sequence of the Serengeti feeding up to 1800 m a.s.l. natural forests have been succession, their samples were drawn from a vari- replaced by plantations of Japanese cedar (Cryp- ety of locations. Therefore, mesowear data from tomeria japonica) (Nowicki & Koganezawa sympatric ungulates with known food habits are 2001, Takatsuki 1983). The mean annual pre- strongly recommended. cipitation and temperature at the Nikko Weather This study was focused on the dietary seg- Station (1292 m a.s.l.) in the southern part of regation of two extant ungulates in Japan: the the park are 2230 mm and 6.0 ℃, respectively Japanese serow (Capricornis crispus, Bovidae) (Nowicki & Koganezawa 2001). and the sika deer (Cervus nippon, Cervidae). In the park, Japanese serow are territorial and In general, these two species are not sympatric solitary. In contrast, sika deer are not territorial in most of Japan (Fig. 3). The Japanese serow and occur in herds. The deer scatter widely over lives in higher mountains, whereas sika deer the mountains in summer, while they concen- usually inhabit mountains and lowlands and tend trate in mid-winter (Takatsuki 1983). Their food to prefer flat habitats (Takatsuki et al. 2010). habits are also contrasting. The Japanese serow However, in the Nikko National Park (NP, central is a browser species (Takatsuki and Suzuki 1984, Japan, Fig. 3), previous ecological studies have Takatsuki et al. 1988, Takatsuki et al. 1995, shown that these two ungulates exist almost sym- Ochiai 1999, Jiang et al. 2008), whereas sika patrically despite having contrasting food habits. deer have more flexible food habits (reviewed As Nowicki and Koganezawa (2001) in Takatsuki 2009). In the park, Japanese serow described, the park is a mountainous area with feed on the leaves of deciduous trees and her- ANN. Zool. FENNIcI Vol. 50 • Effects of dietary differences based on mesowear analysis 203 baceous shrubs during the summer, while in molar. In addition, cusps that were damaged winter they feed mainly on needles of coniferous were not scored, in accordance with the methods trees (Koganezawa 1999). In contrast, grami- of Fortelius and Solounias (2000). noids (i.e., Gramineae, Cyperaceae and Jun- caceae), which dominate the forest floor, are an important food for sika deer (Takatsuki 1983). Reference mesowear data of extant Koganezawa (1999) suggested that the diets of ruminants the two species overlap in winter, although Japa- nese serow rarely fed on graminoids whereas Although Fortelius and Solounias (2000) pro- sika deer consumed them at a high rate through- vided reference data for 64 extant ungulates, they out the year. Nowicki and Koganezawa (2002) selected 27 to form a subset of species for which also reported no evidence of food competition satisfactory dietary data were available and inter- between these species. pretation was uncontroversial. Their dataset of In contrast to previous studies using meso- “typical” species form a good basis for compari- wear analysis, this study attempted a strict com- son with fossil forms, an was used in most previ- parison of two extant ungulates — the sika ous studies. In addition, morphologic differences deer and the Japanese serow — living in the of mammalian teeth could bias the method of same locality (Nikko National Park, central wear (Fortelius & Solounias 2000, Blondel et al.
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