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The Eastern Asian–Eastern North American Floristic Disjunction

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The Eastern Asian–Eastern North American Floristic Disjunction Land Bridge Travelers of the Tertiary: The Eastern Asian–Eastern North American Floristic Disjunction David Yih he eastern Asian–eastern North Ameri- remains the classic disjunction. It continues to can floristic disjunction is a curious phe- stimulate new scientific papers, with each suc- Tnomenon that has fascinated botanists cessive generation applying new research tools for over 200 years: the existence of an entire to its mysteries. catalog of species and genera shared by two Recognition of the disjunction began in vastly separated regions and found nowhere the 1750s with botanists making lists of spe- else. It has inspired generations of researchers cies found in both regions. By the mid-1800s and given impetus to such fields as biogeogra- botanists had collected enough materials to phy and paleobotany. Scientists now recognize lead them to the astounding conclusion that many different disjunct patterns around the the flora of eastern North America (ENA) had world, but the eastern Asian–eastern North more in common with eastern Asia (EA) than American was the first to be discovered, and it did with western North America. Most of I C REDI L T E D ETER P ENA meets EA: A garden path separates the eastern North American species Allegheny spurge (Pachysandra procum- bens, left) from the eastern Asian species Japanese spurge (Pachysandra terminalis). The Eastern Asian–Eastern North American Floristic Disjunction 15 what were once thought to be identical spe- Sequoia (redwood) and Taxodium (baldcypress), cies are now considered congeners (distinct genera now confined to North America, have species belonging to the same genus), so the been found in eastern Asia. disjunction is more about shared genera than At present, the similarity is limited by pro- shared species, and it is now clear that east- nounced differences in biodiversity. Disjunct ern North America has more in common with genera tend to have more species in Asia than western North America than with eastern Asia. in America. The extreme example is Lindera, However, it is also clear that eastern Asia and with 80 species in eastern Asia but only 2 in eastern North America have more in common North America. Indeed, eastern Asia, with its than eastern Asia and western North America 2,753 genera of seed plants, has a biodiversity and that a remarkable disjunction phenomenon far greater than that of eastern North America, exists. Today, the list of EA-ENA botanical “dis- which has only 1,230. According to one expla- juncts” (shared taxa peculiar to the two regions) nation, the Paleocene forests of both regions includes about 65 genera, a handful of closely were equally rich in species until severe cli- related genera, and a few species (Wen 1999). matic fluctuations in North America resulted Most of the genera are temperate; only a in many extinctions. Another possibility is that few come from subtropical or tropical zones. the complex topography of eastern Asia pro- And most disjuncts are woody plants. Many of moted a greater rate of speciation there due to the herbaceous ones are early-leafing species adapted to thrive on the forest floor. Some EA- ENA disjunct genera that have familiar repre- Table 1 sentative species in the Northeast are listed in WOODY Table 1 (Li 1952), along with generalized com- mon names for the species. Campsis Trumpet vine Carya Hickory DISJUNCT REGIONS Catalpa Catalpa The majority of eastern Asian disjuncts grow in the Sino-Japanese Floristic Region, which Cornus Dogwood extends from China’s western Yunnan and Gleditsia Honey locust Sichuan provinces through central, eastern, and Hamamelis Witchhazel most of southern China to Korea and Japan. The richest association of disjunct genera occurs in Liquidambar Sweetgum central China, along the longest river in Asia: Liriodendron Tuliptree the Yangtze (Li 1952). On the American side, Lyonia Maleberry the richest disjunct area is along the Appala- chian Mountains. The two areas are the only Mitchella Partridgeberry instances globally of the mixed mesophytic for- Nyssa Tupelo est, one of the most biodiverse temperate forest Pachysandra Pachysandra types in the world. Western botanists have reported experienc- Parthenocissus Virginia creeper ing a sense of déjà vu in the forests of China. Sassafras Sassafras Add to the disjuncts the more wide-ranging spe- cies that also occur in both regions, and the HERBACEOUS level of similarity becomes quite high. A recent study found that 67% of the seed plant gen- Panax Ginseng era in Maine occur on Japan’s Honshu Island Phryma Lopseed (Qian 2002). The similarity was even greater in Podophyllum Mayapple ages past. Several genera that are now endemic to eastern Asia occur in fossil form in North Saururus Lizard’s tail America, e.g., the familiar Ginkgo and Metase- Symplocarpus Skunk cabbage quoia (dawn redwood), while fossil remains of 16 Arnoldia 69/3 • February 2012 The Sino-Japanese Floristic Region. I I C C REDI REDI L T L T E E D D ETER ETER P P Manchurian catalpa (Catalpa bungei), left, is native to China, while northern catalpa (Catalpa speciosa), right, is native to the central and eastern United States and southern Ontario. The Eastern Asian–Eastern North American Floristic Disjunction 17 H C OSE R Y CREE C AN N avid D OF Courtesy Showy flowers are a feature shared by Chinese trumpet creeper (Campsis grandiflora), left, and the familiar trumpet creeper (Campsis radicans) of North America, right. peninsula and in North America, including ayer M members of genera familiar to New England botanists: Asplenium, Lycopodium, Anemone, obert R Heuchera, and Spirea. Commercial exploitation of the phenomenon had already begun. Père Lafitau, a French Jesuit, had discovered Ameri- can ginseng (Panax quinquefolius) growing near Montreal in 1716, and French Canadians were neglecting their farms in the rush to collect wild ginseng for export to China (Kingsford 1888). In 1784, C. P. Thunberg, a Swedish botanist, included in his Flora Japonica twenty species first described for North America (Boufford and Spongberg 1983). The following year, the Ital- ian botanist Luigi Castiglioni began a two-year sojourn in the United States. Castiglioni’s mis- sion was to bring back useful seeds to Italy, and he is credited with introducing to continental Lindera obtusiloba (seen here in fall color at the Arnold Arbore- tum) is one of the many Lindera species native to eastern Asia. Europe such trees as black locust (later to show invasive tendencies), catalpa, and arborvitae. In the abundance of varied habitats and natural 1790 he published his Viaggio negli Stati Uniti barriers that could allow different populations dell’ America, with elegant botanical drawings of a species to evolve separately (Sargent 1913; of such American sights as the franklintree, Qian and Ricklefs 2000). The EA–ENA disjunc- already rare in the wild and soon to be extinct tion is now recognized not only among plants, outside of cultivation. It also contains the first but among taxa of fungi, arachnids, millipedes, explicit discussion of the floristic similarity of insects, and freshwater fish, as well (Wen 1999). eastern North America to Japan, the only part But botanists can take credit for being the first of eastern Asia for which published floras were to notice and document the phenomenon. then available. Overlooked by nearly all who would later treat the disjunction, he received A THEORY BLOOMS scant credit for his role in its discovery (Li The earliest hint came in the 1750 dissertation 1955). Brief comments in the work of Pursh and of Halenius, a student of Linnaeus. It mentions then Nuttall reveal little beyond an incipient nine species found both on Siberia’s Kamchatka recognition of the disjunction, though Nuttall PETER DEL TREDICI MICHAEL DOSMANN C virginiana blooms inlatewinterorvery earlyspringwhilecommonwitchhazel( Though theirflowerslooksimilar, ultivated 18 ), right,bloomsinlatefallorearlywintereastern North America. C Arnoldia 69/3 hinese ginseng( Panax ginseng • February 2012 C hinese witchhazel( ), left,andafruiting specimenof Americanginseng( Hamamelis mollis Hamamelis ), left, ), left, Panax quinquefolius ), right. PETER DEL TREDICI NANCY ROSE The Eastern Asian–Eastern North American Floristic Disjunction 19 KY C undertook to note the geographical distribu- ENTU K tions of North American genera. It remained OF for Asa Gray, the preeminent American bota- nist of the nineteenth century, to focus atten- tion on the disjunction, bringing it to the notice NIVERSITY , U of the wider scientific community in a series of articles beginning in 1840 and spanning ARNES B nearly 40 years. OM T OF Charles Darwin, who began an extensive cor- respondence with Gray in 1855, encouraged him to study the global distributions of the North American flora. In his second letter to Courtesy Gray he wrote, “The ranges of plants, to the east and west, viz. whether most are found in Greenland and Western Europe, or in E. Asia appears to me a very interesting point as tend- ing to show whether the migration has been eastward or westward” (Darwin 1855). In 1859, after studying new collections from Japan, Gray published his classic “Diagnostic Characters” paper that included a list of 134 species shared by eastern North America and Japan. On the list were such northeastern plants as blue cohosh (Caulophyllum thalictroides), fox grape (Vitis labrusca), ditch stonecrop (Penthorum sedoides), honewort (Cryptotaenia canadensis), hobblebush (Viburnum lantanoides), wild gin- ger (Asarum canadense), red trillium (Trillium
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