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Environmental factors contribute to the formation and maintenance of the contact zone observed in deciduous broad-leaved tree species in Japan

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Abstract

Contact zones are defined as areas where populations from different refugia meet during a postglacial expansion and distinct DNA lineages are mixedly distributed. In Japan, contact zones of various plants and animals were reported from the Kinki-Chugoku region. These contact zones appear to be maintained without any drastic topographic barriers such as those observed in the Alps and Pyrenees Mountains. In this study, the mechanisms underlying the formation and/or maintenance of these contact zones were investigated using six deciduous broad-leaved tree species (Carpinus laxiflora, C. tschonoskii, C. japonica, Magnolia obovata, Padus grayana, and Euonymus oxyphyllus). First, the precise location of the contact zones was examined by intensive genetic analysis of the six species. Second, the relationships between the geographic location of the contact zone and various environmental factors, including climate and topography, were investigated by generalized additive models to reveal the mechanisms of the formation and maintenance of the contact zones. As a result, four of the six examined plant species clearly showed a geographically common contact zone in Hyogo Prefecture and its adjacent areas. The results of the generalized additive models indicate that the pattern of low habitat suitability estimated by ecological niche modeling was the most important factor for determining the location of the common contact zone. These results suggest that areas with low habitat suitability in Hyogo Prefecture restrict the migration and gene flow of the four species in this region, and thus, they maintain the pattern of the contact zones. This study suggests that there are major effects of habitat suitability on the formation and maintenance of the contact zones.

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Acknowledgments

We thank Dr. F. Koike (Department of Risk Management and Environmental Sciences, Yokohama National Univ.), Dr. M. Saito (The Department of General Systems Studies, The Univ. of Tokyo), Mr. H. Mitsuhashi (The Museum of Human Activities Hyogo), Dr. H. Sato (Center for Ecological Research, Kyoto Univ.), Dr. I. Saeki (The Department of World Heritage Studies, Univ. of Tsukuba), Dr. J. Suzuki, Dr. T. Kusano, Dr. M. Kohri (Department of Biological Science, Tokyo Metropolitan Univ.), Dr. T. Sugawara, Dr. H. Kato, Dr. Y. Kakugawa, Mr. M. Nakaji, Mr. S. Shimokawa (Makino Herbarium, Tokyo Metropolitan Univ.), and two anonymous reviewers for their valuable advice. We also thank Dr. A. Ebihara (Department of Botany, National Museum of Nature and Science), Dr. M. Ito, and Mr. O. Kurashima (Department of General Systems Studies, The Univ. of Tokyo) for providing the occurrence data from the locality information of herbarium specimens deposited in the herbarium of National Science Museum, Tsukuba (TNS), and the results of the National Survey on the Natural Environment by the Ministry of the Environment, Japan (NSNE). This study was partly supported by the research project “A new cultural and historical exploration into human-nature relationships in the Japanese archipelago” of the Research Institute for Humanity and Nature (to N.M and A.S).

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Correspondence to Akitaka Tono.

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10265_2015_722_MOESM1_ESM.tif

Fig. S1 The contribution of each variable to the established model evaluated by regularized training gain. Red bars indicate “alone contributions”. Blue and green bars indicate “drop contributions” and contributions when all the variables are used, respectively (TIFF 29095 kb)

10265_2015_722_MOESM2_ESM.tif

Fig. S2 Geographical distribution of the common contact zone, as well as the eastern and western regions divided by the contact zone based on the mean mixed index values (M) among the four species. The sites in the eastern and western regions are indicated in blue and red, respectively, whereas those in the contact zone are indicated in green (TIFF 26335 kb)

10265_2015_722_MOESM3_ESM.tif

Fig. S3 Median joining network relationships among the cpDNA haplotypes in a Carpinus laxiflora, b C. tschonoskii, c C. japonica, d Magnolia obovata, e Padus grayana, and f Euonymus oxyphyllus. Black circles in the network indicate missing haplotypes. Solid bars, open bars, and stars in the networks indicate nucleotide substitution, indel, and inversions, respectively. Mononucleotide repeat variants were removed from this network analyses. The color of each haplotype corresponds to that in Fig. 2a–f (TIFF 14156 kb)

10265_2015_722_MOESM4_ESM.tif

Fig. S4 Scatter plot matrix of the mixed index (M) of the six species. Regression lines are shown in green (TIFF 15564 kb)

10265_2015_722_MOESM5_ESM.tif

Fig. S5 Estimated suitable habitats using the Maxent software for the four species: a Carpinus laxiflora, b C. tschonoskii, c C. japonica, and d Magnolia obovata (TIFF 21626 kb)

10265_2015_722_MOESM6_ESM.tif

Fig. S6 Principal component analysis of the climatic variables in the Kinki-Chugoku region. PC1 (x-axis) and PC2 (y-axis) explain 60 % and 23 % of the total variation, respectively. The climatic variables highly contributing to each component are shown. The sites in the contact zone, eastern region, and western region are indicated in green, blue and red, respectively. The number beside each mark corresponds to the collection site in Table S2 (TIFF 44087 kb)

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Tono, A., Iwasaki, T., Seo, A. et al. Environmental factors contribute to the formation and maintenance of the contact zone observed in deciduous broad-leaved tree species in Japan. J Plant Res 128, 535–551 (2015). https://doi.org/10.1007/s10265-015-0722-y

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