Climate Change Impacts on Migration of Pinus Koraiensis During the Quaternary Using Species Distribution Models

Plant Ecol https://doi.org/10.1007/s11258-021-01147-z (0123456789().,-volV)(0123456789().,-volV) Climate change impacts on migration of Pinus koraiensis during the Quaternary using species distribution models Takuto Shitara . Shunsuke Fukui . Tetsuya Matsui . Arata Momohara . Ikutaro Tsuyama . Haruka Ohashi . Nobuyuki Tanaka . Takashi Kamijo Received: 27 October 2020 / Accepted: 28 May 2021 Ó The Author(s) 2021 Abstract Clarifying the influences of paleoclimate and four bioclimatic variables at 2.5 arcminute changes on the disjunct distribution formation of (approximately 5 km) spatial resolution. The simula- plants allows a historical and mechanical understand- tion revealed the excellent fit of the MaxEnt model ing of current vegetation and biodiversity. This study performance, with an area under the curve (AUC) investigated the influences of paleoclimate changes on value of 0.922 and continuous Boyce index (BCI) the present disjunct distribution formation of Pinus value of 0.925 with fivefold cross-validation. The most koraiensis (Korean pine) using species distribution important climatic factor was the minimum temper- modeling. A species distribution model (SDM) was ature of the coldest month. Suitable habitats for the built using maximum entropy principle algorithms species ranged between - 30.1 and - 4.1 °C. Pro- (MaxEnt), data from 152 occurrences of the species, jected suitable habitats under the Last Glacial Max- imum (approximately 22,000 years ago [ka BP]: LGM) period showed wide distributions in eastern Communicated by Peter le Roux. China, the southern part of the Korean Peninsula, and Supplementary Information The online version contains the Japanese Archipelago. After the mid-Holocene supplementary material available at https://doi.org/10.1007/ (approximately 6 ka BP), the suitable habitats s11258-021-01147-z. T. Shitara (&) T. Matsui Á T. Kamijo Institute of Agriculture, Tokyo University of Agriculture Faculty of Life and Environmental Sciences, University of and Technology, 3-5-8 Saiwai-cho, Fuchu-shi, Tsukuba, 1-1-1 Tennodai, Tsukuba-shi, Tokyo 183-8509, Japan Ibaraki 305-8571, Japan e-mail: [email protected] A. Momohara S. Fukui Graduate School of Horticulture, Chiba University, 648 Ministry of the Environment of Japan, 1-2-2, Matsudo, Matsudo-shi, Chiba 271-8510, Japan Kasumigaseki, Chiyoda-ku, Tokyo 100-8975, Japan I. Tsuyama T. Matsui Forest Research and Management Organization, Center for Biodiversity and Climate Change, Forestry and Hokkaido Research Center, Forestry and Forest Products Forest Products Research Institute, Forest Research and Research Institute, 7 Hitsujigaoka, Toyohira, Sapporo-shi, Management Organization, 1 Matsunosato, Tsukuba-shi, Hokkaido 062-8516, Japan Ibaraki 305-8687, Japan 123 Plant Ecol expanded northwards in continental regions and continent, such as the Japanese Archipelago (Fig. 2a). retreated from both north and southwest of Japan. Cool and cold temperate forests are among the forest This eventually formed disjunct suitable habitats in types that are most representative in these regions and central Japan. An increase in temperature after the are characterized by higher plant species richness than LGM period caused the migration of P. koraiensis other forests at the same latitude, such as those in toward new, suitable habitats in continental Northeast North America and Europe (Latham and Ricklefs Asia, while species in the Japanese Archipelago 1993; Qian and Ricklefs 2000). retreated, forming the present disjunct distributions. In these forest zones, some of the main canopy tree species such as Pinus koraiensis Siebold et Zucc. Keywords Climate change Á Disjunct distribution Á (Korean pine), Picea sect. Picea, and Betula davurica Macro fossil Á Northeast Asia Á Quaternary Á Species are widely distributed in continental regions (Kolbek distribution model 2003; Krestov et al. 2006) and are also distributed in the Japanese Archipelago, where they occur only in a few montane regions, showing present-day disjunct patterns of geographical distribution (e.g., Katsuki Introduction et al. 2004; Aizawa et al. 2012; Shitara et al. 2018). Macro-fossil and pollen fossil records indicate that The factors that contribute to the establishment of these species were widely distributed in the Japanese geographical disjunct distributions of plant species Archipelago during the Last Glacial Maximum (LGM; have long captured the interest of botanists and between ca. 27 and 19 thousand years ago [ka BP]; ecologists (e.g., Wood 1972; Wen 1999; Qian and Clark et al. 2009), and rapidly reduced from most areas Ricklefs 2000; Richard 2006). Climatic changes over in response to the global warming that occurred the glacial and interglacial periods during the Quater- thereafter (e. g., Tsukada 1983; Morita 2000; Takahara nary period (over the past 2.6 million years) have et al. 2010). However, how and why post-LGM caused the expansion and retraction of plant habitats, climate changes have influenced these distribution local extinctions, and the development of present-day shifts remains controversial. disjunct distribution patterns (Hewitt 1996; Comes Under such circumstances, species distribution and Kadereit 1998; Svenning et al. 2008; Chen et al. models (SDMs) are effective modern tools that 2012; Shitara et al. 2018). Accordingly, elucidating quantitatively determine which environmental vari- the processes whereby the disjunct distributions of ables are most strongly correlated with species distri- plant species develop in relation to paleoclimate butions and can be used to project suitable and change provides an important context for understand- vulnerable habitats under climate change scenarios ing the development of present-day vegetation and (Pearson and Dawson 2003; Elith 2006; Phillips et al. biodiversity (Wang et al. 2017). 2006; Svenning et al. 2011). However, the uncertainty Northeast Asia is composed of three landmass regarding the former habitats of species projected by elements: continental regions, such as the eastern to SDMs using paleoclimatic data should ideally be northeastern parts of China and around the southern verified by evidence such as that provided by macro- parts of Russian Far East; peninsulas extending south fossil records (Svenning et al. 2011; Varela et al. 2011; from the continent, such as the Korean Peninsula; and Roberts and Hamann 2015). the East Asian Islands running parallel to the P. koraiensis can be considered to be ideal for investigating the possible impacts of climate on the development of disjunct species distributions, because H. Ohashi numerous macro-fossil records of this species dating Department of Wildlife Biology, Forestry and Forest Products Research Institute, Forest Research and back to the LGM have been found in the Japanese Management Organization, 1 Matsunosato, Tsukuba-shi, Archipelago (Miki 1956; Tsukada 1983; Morita Ibaraki 305-8687, Japan 2000). This species occurs mainly in the Amur and Maritime provinces of the Russian Far East, northeast N. Tanaka Environment Consultant ENVI, 602-89 Akatsuka, China, the Korean Peninsula, and the Japanese Tsukuba-shi, Ibaraki 305-0062, Japan Archipelago (Okitsu and Momohara 1997; Potenko 123 Plant Ecol and Velikov 1998). However, although this species is nutcrackers (Nucifraga caryocatactes), which also widely distributed in continental Northeast Asia function as agents of seed dispersal (Miyaki 1987; (Potenko and Velikov 1998; Kolbek et al. 2003), it Hutchins et al. 1996). In this regard, the Eurasian rarely grows naturally in the Japanese Archipelago, nutcracker, which occasionally caches seeds in the where it is primarily confined to relatively small areas soil, is particularly important (Hutchins et al. 1996). in the central mountainous regions of Honshu and The timber of P. koraiensis is mainly used for Shikoku, thus indicating its present-day disjunct construction purposes, and is edible; the seeds are a distribution (Okitsu and Momohara 1997; Aizawa valuable source of local income (Thomas and Farjon and Kaneko 2009; Fig. 2a). Macro-fossil records 2013). In addition, mixed deciduous broad-leaved and indicate that this species was widely distributed in P. koraiensis forests provide important habitats for the Japanese Archipelago during the LGM, but rapidly wildlife, including the Siberian tiger (Panthera tigris retreated from most areas thereafter (Okitsu and altaica; Thomas and Farjon 2013). In recent years, Momohara 1997; Morita 2000; Okitsu 2002). More- however, the distribution range of this species has over, few previous studies have shown that P. been markedly reduced owing to logging activities koraiensis is strongly affected by climate change (Thomas and Farjon 2013). In the Japanese archipe- (Xu and Yan 2001; He et al. 2005; Zhang et al. 2014; lago, although P. koraiensis is included as a ‘‘lower Ahn et al. 2015; Belyanin and Belyanina 2019). risk species’’ in the Japanese Red Data Book (Envi- The aim of this study was to identify the climatic ronment Agency of Japan 2012), the populations of factors that have influenced the distribution of P. this species are quite small compared to that of Abies koraiensis and determine how paleoclimate change veitchii, Picea jezoensis var. hondoensis, and Tsuga has affected the distribution and development of the diversifolia which are common species in the sub- present-day disjunct distribution of P. koraiensis in alpine zone of Honshu. In addition, a P. koraiensis- Northeast Asia. To this end, we initially constructed an dominated forest is very

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