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Molecular Analysis: a New Look at Umbrella Magnolias Richard B

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Molecular Analysis: a New Look at Umbrella Magnolias Richard B Molecular Analysis: A New Look at Umbrella Magnolias Richard B. Figlar Taxonomists have long been frustrated in their attempts to decipher the complex evolu- tionary relationships within the genus Magnolia. Recent molecular research has shed new light on the problem and helped to clarify the long- standing confusion. The magnolias of section Rytido- spermum-one of sixteen categories that subdivide the 128 species of the genus Magnolia-have always been an intensely interesting group, not only for their large flowers and enor- mous whorled leaves, but because several species occur in both eastern Asia and eastern North America. Within the genus, this intercontinen- tal distribution is shared only with section Tuhpastrum, but in that case the two species involved, our native cucumber tree, Magnolia acumi- nata, and M. liliiflora, the famous Mulan magnolia from China, share few characteristics beyond the same number of chromosomes and the These three closely related magnolras share large, whorled leaves, a to two m and white presence of reduced outer tepals. rangmg from foot feet length, large, flowers with diameters m the six to twelve mches. The The Rytidospermum section, range of flowers, which open after the leaves have developed, are strongly scented, to most according taxonomists, Magnolia tnpetala, above, unpleasantly so. A natme of the consists of six species: MagnoliaAllegheny region of the eastern Umted States, it seldom exceeds tripetala, the umbrella magnolia; M. ;forty feet and is uncommon both m the mld and m cultivation. fraseri, the mountain magnolia; M. At top mght Is the Japanese Magnolia obovata. It grows to macrophylla, the big-leaf magnolia; eighty feet m its native damp, mch, highland forests, and is one of M. obovata (M. hypoleuca), M. the hardiest Asian magnohas (zones 6 to 9). Its shghtly less hardy Chmese M. officinalis var. at bottom also and M. rostrata-the first sister, biloba, right, officinalis, ’ at to ‘grows altitudes from 2,000 5,500 feet, and achieves heights three native to southeastern Umtedup to seventy feet. Its bark is so highly valued as medicine that States into Mexico and the latter the. tree has been nearly extmpated m its native provmces of Hubei three native to eastern Asia, from theand Sichuan 23 24 represent true whorls, the leaves of Rytidospermum magnolias are arranged in false whorls; that is, the individual leaves actually emerge in alternate fashion but with very little stem growth (internodes) between successive leaves. The pattern of many leaves emerg- ing almost simultaneously is called flushmg. Apparently, Rytidosper- mum magnolias adapted this flush- type leaf-emergence pattern in order to compete effectively in the gaps of forest understory during early spring. By producing more leaves more or less simultaneously, such plants are better able to compete with other species for scarce sunlight. And since little stem growth is produced, the process itself is very energy efficient. Later in the spring, the growth reverts to the more typical pattern, where leaves are produced one at a time along longer stem shoots, as in other magnolias. Flush-type leaf- emergence patterns are common in many other plant species of the understory; for instance, some of the deciduous azaleas, although because of their much smaller leaves, the umbrella effect is less noticeable than in the Rytidosper- mum magnolias.2 Clearly, among the magnolias this Like other members the three within of subgenus Magnolia (one of trait is and for that reason the the M. tend to be red and unique, genus), frmts of tripetala bmght taxonomists have showy They persist for several weeks in late summer. suggested that, despite their intercontinental distri- Kurile Islands and Japan westward to southwest bution, they all form a natural group and should China. Among the various morphological char- be very closely related. This provokes several acteristics shared by members of this group, the questions. Did today’s species evolve from a most distinctive are the enormous whorls of common ancestor? If so, how and when did its deciduous leaves, which are crowded m parasol descendants cross the Pacific Ocean? Which one fashion at the ends of the branches. For this of the North American species is the most reason, the Rytidospermum magnolias are closely related to its Asian counterpart(s)? often referred to as umbrella trees. Indeed, to the Using modern molecular systematics, uninitiated, the first impression of these plants researchers Yin-Long Qiu, Clifford Parks, and is often more reminiscent of the houseplant Mark Chase analyzed the chloroplast DNA known as the umbrella tree-the giant tropical (cpDNA) of all section Rytidospermum species. Schefflera-than it is of a Magnoha. However, (CpDNA is the part of the DNA chromosome unlike Schefflera, whose compound leaves that is reponsible for photosynthesis. ~ By com- 25 paring the differences in the cpDNA of the (between all combmations of pairs of species) various species, they were able to quantify the over the entire chloroplast genome. The analy- amount of evolutionary change, measured as sis counts the number of site changes encoun- molecular divergence, that had taken place tered, then calculates the cpDNA sequence between them. The underlying assumption or divergence (as a percentage of sequence diver- theory in this method is that the amount of gence/ between all species pairs. The results are genetic difference is proportional to the amount shown in Table 1. of time elapsed since the species diverged from This analysis clearly shows that Magnolia their common ancestor, relative to other pairs tripetala from eastern North America has or groups of organisms being compared. The diverged far less from the Asian species M. results of the study team were published in two obovata and M. officinalis var. biloba than it separate papers in the American Journal of has from other North American species. It also Botany, both in 1995. This article attempts to indicates that the other North American species summarize the findings of these researchers and have diverged just as much from each other to interpret how molecular data, when used in (including M. tripetala) as they have from the conjunction with traditional morphological two Asian species. studies, can lead to better understanding of the evolutionary relationships among plants. No Allozyme Electrophoresis attempt will be made in this article to decode The study team used a second method, allo- the complexities of their analytic techniques, zyme electrophoresis, to examine genetic varia- the details of which are treated in the study tion of enzyme-coding genes. This analysis team’s original papers.4-s results in the calculation of a parameter called Nei’s unbiased genetic identity for each of the Restriction Site CpDNA Analysis species pairings. The numbers are from zero to Qui, Parks, and Chase used three different one, with one being a perfect genetic match. laboratory techniques to assess the divergence One of the authors, Clifford Parks, suggests that among Magnolia obovata, M. tripetala, M. as a rule of thumb, readings greater than 0.90 fraseri, M. macrophylla, and M. officmalis suggest populations of the same species, while var. biloba (a variety of M. officmalis with readings less than 0.67 indicate distinctly differ- notched or bilobed leaves; shown in the tables ent species. The results can be seen in Table 2. as M. biloba). The first method, cpDNA restric- Though not shown in the table, it should also be tion site analysis, randomly samples changes noted that Nei’s genetic identity for mtraspe- TABLE 1 TABLE 2 26 cific comparisons was nearly 1.000, as would extremely low-the lowest divergence ever be expected: the values ranged from 0.993 for reported for any eastern Asia-eastern North M. obovata vs. M. obovata to 0.932 for M. America disjunct taxa." For example, the macrophylla vs. M. macrophylla. sequence divergence over the entire chloro- The results of this second method almost plast genome (cpDNA) between Limodendron mirror the results of the restriction site analysis, tulipifera and L. chinense was found to be 1.24 giving very strong evidence of a close relation- percent (as compared to 0.083 percent between ship between Magnolia tripetala and the Asian M. obovata and M. tripetala/,3 which is a species and relatively distant relationships remarkable difference in that many taxonomists among the rest of the species. It is interesting long considered both Liriodendron taxa to be to note that in both analyses the relationship varieties of the same species. between M. fraseri and M. macrophylla is The study team speculated how and when the most distant of any of the pairs. Iromcally, Magnolia tripetala and its sister species became some texts on North American trees refer separated from their common ancestor. One to these two species as closely related on hypothesis is that the common ancestor could account of their similar auriculate (earlobe- have migrated between the contments via the shaped) leaf bases. Bering land bridge during one of the earth’s warm periods in the middle Miocene (17 to 15 Gene rbcL Chloroplast Sequencing million years before the present) or early Finally, the study team comparedll Pliocene (6 to 5 million years before the tripetala, M. macrophylla, and M. TABLE 3 obovata to each other by analyzing (i.e., sequencing) a specific segment of the chloroplast gene called rbcL. This analysis involves comparing the 1,432 base pairs of the rbcL gene for each pair of species in the analysis, which in this case is three (M. macrophylla vs. M. obovata, M. macrophylla vs. M. tripetala, and M. obovata vs. M. tripetala). The results, once again, confirm the find- ings of the first two analyses, which suggest that M. tripetala and the two Asian species form a clade, or "sister group." In fact, the sequenc- ing of the chloroplast gene rbcL yielded no divergence between M.
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