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Hypothesizing Origin, Migration Routes and Distribution Patterns of Gymnosperms in Taiwan

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Hypothesizing Origin, Migration Routes and Distribution Patterns of Gymnosperms in Taiwan Taiwania, 59(2): 139‒163, 2014 DOI: 10.6165/tai.2014.59.139 REVIEW Hypothesizing Origin, Migration Routes and Distribution Patterns of Gymnosperms in Taiwan Shing-Fan Huang No. 521, Nanda Road, Hsinchu 300, Taiwan, Department of Applied Science, National Hsinchu University of Education. * Corresponding author. Phone: 886-3-5213132#2750; Fax:886-3-5257178; Email: [email protected] (Manuscript received 22 Febuary 2013; accepted 11 March 2014) ABSTRACT: Phytogeographical study of gymnosperms in Taiwan is carried out based on reviewing data gathered from published papers on fossils, phylogeny and phylogeography. Following questions are asked. (1) How is the high degree of endemism of gymnosperm flora of Taiwan derived? (2) How many source areas of gymnosperms in Taiwan are there? (3) Is there relation between distribution pattern of endemic gymnosperms in Taiwan and those of their sister species? (4) How do gymnosperms migrate to Taiwan? In total, 28 taxa including 19 species and 9 varieties of gymnosperms are in Taiwan. Compared to the Flora of Taiwan 2nd edition, Nageia fleuryi is excluded and Pinus taiwanensis var. fragilissima is added in this paper. Species status of Calocedrus formosana and Tsuga formosana and variety status of Cunninghamia lanceolata var. konishii are retained. Scientific names are adopted for Juniperus morrisonicola instead of J. squamata and for Juniperus tsukusiensis var. taiwanensis instead of J. chinensis var. taiwanensis. According to distribution patterns, these 28 taxa may be categorized into tropical origin (TO), Southern Hemisphere origin (SMO) and Northern Hemisphere origin (NMO). Gymnosperms in Taiwan with high degree of endemism, 78.5%, may owe to woody habit, which is wider in ecological niche compared to herbaceous one and would be less sensitive to the environmental changes, and owe to temperate essence that is more easily to find shelters during temperature fluctuations. Taxa of TO and SMO are inclined to inhabit low altitudes and sporadically distributed, whereas taxa of NMO are inclined to inhabit middle to high altitudes, especially in northern and central Central Mountain Range and may be widely or restrictedly distributed. Distribution patterns of endemic taxa of NMO in Taiwan are related with those of their sister species. Taxa with sister species in higher latitudes such as Japan, northwestern China and central China are distributed in higher altitudes with midpoint of altitudinal distribution over 1800 m, while those with sister species in lower latitudes such as South China, southern South China, southeastern China are distributed in lower altitudes with midpoint of altitudinal distribution under 2000 m. Most fossil histories of endemic taxa of NMO may trace back to Asia or North America (NAM) except Juniperus morrisonicola that may trace back to Europe. For those traced back to NAM, ancestors in NAM migrated to northeastern Asia via Biringia, from where dispersed southward either to Japan, or to northern China and then to central and eastern China. From Japan, ancestors either migrated southward through the Ryukyus to Taiwan if sister species were restricted to Japan, or they might have dispersed to continental Asia and evolved when Japan was a part of continental Asia and further migrated southward via East China Sea’s land bridge to Taiwan. From central or eastern China, ancestors migrated southward either via East China Sea’s land bridge or through southeastern China via Tungshan land bridge to Taiwan. Ancestors in Europe migrated southward to the Himalayas, from where through the Yun-Kue Plateau, Nanling via Tungshan land bridge to Taiwan. Southwestern China (SWC) plus IndoChina is both refuge and dispersal center. Taxa of SMO might have dispersed from the South Hemisphere through southeastern Asia to IndoChina, from where migrated either through southern South China via South China Sea’s land bridge to southern Taiwan, or through South China via Tungshan land bridge to central Taiwan. If taxa of NMO share short genetic distance with their sister species in SWC, their migration routes would be like those of SMO. However, if taxa of NMO share longer genetic distance with their sister species in SWC, one lineage of their ancestors, possibly distributed in central China then, migrated through southeastern China via Tungshan land bridge to Taiwan while another lineage in central China further dispersed to SWC and produced disjunct distribution patterns. Taxa of gymnosperms in Taiwan distributed in higher altitudes are inclined to have sister species distributed in higher latitudes. However, horizontal distribution patterns of gymnosperms in Taiwan may be blurred by long history of colonization. Thus horizontal distribution patterns can only be explained by obtaining more data on fossils and paleogeography of such taxa in Taiwan. KEY WORDS: Distribution, endemism, gymnosperms, hypothesizing origin, migration route, Taiwan. 139 Taiwania Vol. 59, No. 2 INTRODUCTION about geology and fossils have been published. Thus it is attempted to summarize what have known about the Gymnosperms are seed plants with naked ovules phytogeography of gymnosperms in Taiwan based on (Bell and Hemsley, 2000). Extant gymnosperms are published data on fossils, phylogeny and classified into four classes, namely Cycadidae, phylogeography to further understanding the possible Ginkgoidae, Gnetidae and Pinidae, each with 2, 1, 3, 12 reasons behind the distribution patterns and to serve as a families and 10, 1, 3 and 69 genera (Christenhuszi et model for extrapolating to the flora of Taiwan. al., 2011). Distribution patterns of genera of Pinidae are Moreover, it is also to respond the idea proposed by either primarily of the Northern or Southern Huang (2011) that determining the relation between Hemisphere and both patterns are overlapped in distribution patterns of taxa in Taiwan and their source southeastern Asia, and each pattern can be further areas or distribution patterns of their sister taxa is a part distinguished into restricted or disjunct distribution (Li, of the study of historical biogeography of the Flora of 1953; Conteras-Medina and Vega, 2002). Nine areas of Taiwan. Hence following questions are asked in this endemism of gymnosperms, namely, southwestern paper. (1) How is the high degree of endemism of China, Japan, New Caledonia, western North America, gymnosperm flora of Taiwan derived? (2) How many Mesoamerica, southern South America, eastern source areas of gymnosperms in Taiwan are there? (3) Is Australia, Tasmania and southern Africa are recognized there relation between distribution pattern of (Conteras-Medina and Vega, 2002). Southeastern Asia gymnosperms in Taiwan and those of their sister plus southwestern China is considered the most species? (4) How do gymnosperms migrate to Taiwan? important diversity center of gymnosperms regarding species richness per 10,000 km2 (Mutke and Barthlow, MATERIALS AND METHODS 2005). Taiwan is situated near mainland China and Japan and its gymnosperm flora contains 5 families Taxa of gymnosperms in Taiwan were compiled (Cupressaceae, Cycadaceae, Pinaceae, Podocarpaceae mainly based on the Flora of Taiwan 2nd edition and Taxaceae, based on the classification of (Editorial Committee of the Flora of Taiwan 2nd edition, Christenhuszi et al. (2011)), 17 genera, and 28 taxa 1994) and published papers afterward. Distribution (Editorial Committee of the Flora of Taiwan 2nd edition, patterns of gymnosperms in Taiwan, mainly following 1994). They are distributed in Taiwan proper, except Liu (1966) and modified with updating data, was one species, Podocarpus costalis, in Lanyu, compiled and summarized in Table 1. Horizontal southeastern isle off Taiwan. distribution pattern of each taxon in Taiwan was Among these 28 taxa, 64.3% is endemic compared described by distribution districts recognized by Huang to 26.2% for all vascular plants in Taiwan (Hsieh, (2011), and vertical distribution pattern of each taxon 2002). Such a high degree of endemism in gymnosperm was expressed by the average of its elevational flora of Taiwan requires an explanation. Regarding distribution as midpoint altitudinal distribution. Sister distribution pattern, three distribution types have been species were determined by published phylogenies of recognized for genera of conifers in Taiwan (Liu, each taxon and topologies of phylogenies were 1966): primarily of the Northern or Southern described in Venn diagrams (cf. Kitching et al., 1998). Hemisphere, disjunct between eastern Asia and North Genetic distance (p = n/l, where n is the number of America, and restricted to East Asia. For family substitutions and l is the length of aligned sequences, distribution, most families in Taiwan are primarily of while number of indels are excluded from counting as the Northern Hemisphere except Cycadaceae of tropics variation) between gymnosperms in Taiwan and their and Podocarpaceae of the Southern Hemisphere. sister populations or species were calculated by aligning Liu (1966) carried out an intensive study on and comparing the similarity of sequences of gene phytogeography of gymnosperms in Taiwan by markers. It was carried out by uploading a sequence reviewing taxonomy of each taxon and comparing their belonging to taxon of gymnosperms in Taiwan to distribution patterns. He (Liu, 1966) concluded that GenBank (http://www.ncbi.nlm.nih.gov/) and using relationship between gymnosperm flora of Taiwan and BLAST program employed in the website to align and China was direct while that between
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