J Syst Evol ›› 1995, Vol. 33 ›› Issue (2): 105-130.

• Research Articles •     Next Articles

Studies on the Geographic Distribution, Origin and Dispersal of the Family Pinaceae Lindl.

Li Nan   

  • Published:1995-03-10

Abstract: Pinaceae Lindl., containing 10 genera and about 235 species, is the largest family in the extant conifers. It widely spreads in the Northern Hemisphere and plays a very important role in coniferous forests occurring in temperate to subtropical mountains. Numerous studies on this family have been carried out and the data dealing with many aspects of biosystematics of the Pinaceae have been accumulated. Based on the principle of unity of phylogeny and distribution of plants, and on the data from the studies of biosystematics of the Pinaceae, the present paper discusses the problems related to geographic distribution and phylogeny of the family in three respects as follows: (1) Floristic division of the Pinaceae is made based on Farjon's work (1990). Six regions and four subregions are outlined (Fig. 1). These are: I. the Mediterranean Region; II. the Eastern European and Siberian Region;III. the Eastern Asiatic Region, which can be further divided into two subregions, i. e. III a. the Northern Eastern Asiatic Subregion and III b. the Himalayas and Southern Eastern Asiatic Subregion; IV. the Western Northern American Region which also contains two subregions, namely IV a. the Northwestern North American Subregion and IV b. the Southwestern North American Subregion V. the Northern North American Region; VI. the Southeastern North American Region. The numbers of species occurring in all these floristic regions are shown in Table 1. The statistic results show that the Subregion III b is currently the richest in species of the Pinaceae. All the living genera are represented in this subregion, including three endemic genera: Keteleeria, Cathaya and Pseudolarix. The second richest area is the Subregion IV b which contains a great number of species. In fact, the two subregions are considered as counterparts. In addition, the Subregion III a and Subregion IV a, the Region II and Region V are also pairs of counterparts. The former pair has fewer but widely spread species, most of which are comparatively young probably developed from the extended refuges after the glacier period of the Quaternary. (2) The geographic distribution of all the genera are described and compared. The maps of their present ranges and their fossil localities are drawn. The four generic distribution patterns are detected: a) North Temperate areal type: containing four genera: Pinus, Picea, Larix and Abies; b) East Asian and North American disjunct areal type: including two genera:Tsuga and Pseudotsuga; c)Mediterranea-Himalayan areal type: containing only one genus: Cedrus; d) Himalayas and Southern Eastern Asiatic areal type: containing three genera: Keteleeria, Cathaya and Pseudolarix. The latter two are endemic to China. (3) The origin, differentiation and early migration of the Pinaceae are studied through the analyses of the data mainly on fossils ( including both extinct and extant genera ), paleogeography, paleoclimate and paleoflora. The main opinions of the present author are as follows: ① The Pinaceae was a large group of plants in geological stages, encompassing many genera with most of them becoming extinct after Mesozoic. The morden Pinaceae may be the offsprings of a few temperate-adapted members, However, they surpassed their ancestors and developed into the main components of current coniferous forests in north temperate zone to north subtropical mountainous regions. The modern Pinaceae is probably a derived group and its prosperity could be related to the emergence of temperate flora. ② Although the origin of the Pinaceae could be traced back to Jurassic or even Triassic, the occurrence of the modern genera of Pinaceae was merely from the Early Cretaceous to the Tertiary. ③ The genera of the Pinaceae may be differentiated in different stages and places. Pinus is possibly the earliest differentiated one among the extant genera. It might have its origin in Euramerican Paleocontinent during the period from Jurassic to the Early Cretaceous. The other genera might have not been diverged from their ancestral complex until the Late Cretaceous to the Tertiary, with one or two of them even until the Middle Tertiary. The place of the differentiation of these genera are supposed to be also restricted in Laurasia, but I intend to conside that it shifted to the North Pacific floristic region, where is currently the greatest diversity of the Pinaceae taxa. ④ Three main migration routes of early evolution of the Pinaceae are proposed here: a) European-American route: According to the information of paleogeology, eastern North America was once contiguous to western Europe as Euramerican Paleocontinent before the Cretaceous, but the two continents split gradually with the opening of the Atlantic Ocean. At the end of the Late Cretaceous, the two parts were still connected through Greenland and an Atlantic floristic region existed. The Euramerican Paleocontinent may be the place for differentiation of the Pinaceae in early stage, while the Atlantic floristic region was a migration route in the modern Pinaceae. b) Eurasiatic route: Before the Late Cretaceous, the Tethys Sea stretched from west to southeast of Eurasia. In the area north of the Tethys Sea, plants could disperse freely. By the Late Cretaceous, however, the existence of the West Siberian Sea and Turgai Straits restricted the exchanging of the Pinaceae plants between Europe and southeast Asia mainly to the coast of the Tethys Sea. Although the Tethys Sea disappeared later and the Himalayas arose, the area along the original coast of the Tethys Sea also remained as a route which played an important role in the dispersal and distribution of the modern Pinaceae. c) Paleoberingian route: At the beginning of the Late Cretaceous, eastern Asia was contiguous to the west of North America through Paleoberingia and formed “Asia-America” landmass. This situation did not cease till Pliocene. The paleoberingian route existed on the basis of this situation, playing a main role in dispersal of the morden Pinaceae between eastern Asia and western North America. There are many taxa ( generic or infrageneric ) in the modern Pinaceae with the patterns which belong to “East Asian and North American disjunct areal type” . The formation of the pattern ismostly related to the existence of the Paleoberingian route. ⑤ The existence of the above mentioned three migration routes is the basis for wide distribution of the Pinaceae in the Northern Hemisphere. In addition, the distribution patterns of the extant genera have formed as the results of the tectonic movements and the changes in paleoclimate and paleoflora since the Tertiary. 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Key words: Pinaceae, Floristic regions, Geographic distribution, Origin, Migration route