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2 Department of Biology, Sungkyunkwan University, Suwon 440–746, Korea; and 3 Department of Biology, Colorado State University, Fort Collins, Colorado 80523 USA
Received for publication August 17, 1999. Accepted for publication March 9, 2000.
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ABSTRACT |
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 TOP ABSTRACT INTRODUCTIONMATERIALS AND METHODSRESULTSDISCUSSIONLITERATURE CITED |
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Key Words: Amygdaloideae • ITS sequences of nrDNA • phylogeny • Prunus • Rosaceae
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INTRODUCTION |
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TOPABSTRACTINTRODUCTIONMATERIALS AND METHODSRESULTSDISCUSSIONLITERATURE CITED |
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200 species with most species in the temperate zone and some in the tropical and subtropical regions (Rehder, 1940
; Willis, 1985
; Yü et al., 1986
; Ghora and Panigrahi, 1995
; Mabberly, 1997
). Prunus is economically important because many species are sources of fruits (e.g., plums, peaches, apricots, cherries, and almonds), oil, timber, and ornamentals. Prunus is generally defined based on a combination of characteristics: a solitary carpel with a terminal style, a fleshy drupe, five sepals, five petals, and solid branch pith (Rehder, 1940
). The most widely adopted classification of Prunus (Rehder, 1940
) divided it into five subgenera, although some workers recognized six to ten genera (de Tournefort, 1700
; Roemer, 1847
; Yü et al., 1986
; Takhtajan, 1997
) within the generic concept of Prunus s.l.
The subfamily Amygdaloideae differs from other rosaceous subfamilies by having a drupe, a fleshy fruit with a stony endocarp or stone (de Candolle, 1813
). Traditionally four genera (Maddenia Hook. f. & Thomson, Oemleria Reichb., Prinsepia Royle, and Prunus) were included in the Amygdaloideae (Rehder, 1940
). Exochorda Lindley has a capsule and was previously placed in the Spiraeoideae (Rehder, 1940
). Several lines of evidence have suggested its close affinity with the Amygdaloideae (Goldblatt, 1976
; Zhang, 1992
; Morgan, Soltis, and Robertson, 1994
). Recent workers have thus placed it in the Amygdaloideae (Takhtajan, 1987, 1997
; Thorne, 1983, 1992a, b
). An additional genus Pygeum Gaertner was previously recognized (Gaertner, 1788)
, but synonymized to Prunus subgen. Laurocerasus (Kalkman, 1965
). Oemleria was previously known as Nuttalia Torr. & Gray (Hooker and Arnott, 1839)
and Osmaronia Greene (Hutchinson, 1964
). Landon (1975)
showed that Oemleria is the correct name of the western North American monotypic genus and this name has been recently accepted (e.g., Hickman, 1993
). Plagiospermum Oliver was recorded by Oliver (in Hooker, 1886)
but later treated as Prinsepia (Bean, 1909
; Rehder, 1915
). On the other hand, Prinsepia (including Plagiospermum) and Oemleria were sometimes resurrected to the subfamilial or tribal levels, e.g., the subfamily Prinsepioideae (Sterling, 1963
), the tribe Osmaronieae (Rydberg, 1918
; Sterling, 1964b
), and the subfamily Osmaronieae (Hutchinson, 1964
; Kalkman, 1988
).
The taxonomy of Prunus, especially concerning the generic delimitation, has been controversial (Table 1). Based primarily on fruit morphology, de Tournefort (1700)
recognized six distinct genera within Prunus s.l.: Amygdalus L., Armeniaca Miller, Cerasus Miller, Laurocerasus Duhamel, Persica Miller, and Prunus s.s. (sensu stricto). The de Tournefort treatment was accepted to some degree but amended by later workers. Linnaeus (1753)
recognized three genera: Amygdalus, Padus Miller, and Prunus s.s. and later (1754)
accepted four genera: Armeniaca, Cerasus, Padus (including Laurocerasus), and Prunus s.s. The Tournefort and Linnaean generic concepts were adopted by several workers, e.g., Miller (1754)
, Komarov (1971)
, Yü et al. (1986)
, and Ghora and Panigrahi (1995)
. Minor changes were often made. For example, Laurocerasus was synonymized under Padus (Miller, 1754)
; Armeniaca was included in Prunus (Ghora and Panigrahi, 1995
) or in Persica (de Jussieu, 1789
); and Laurocerasus was treated in Cerasus (de Candolle, 1825
).
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were the first to unite the six genera of de Tournefort (1700)
into a single genus Prunus, which was subdivided into seven sections: sects. Amygdalopsis, Amygdalus, Armeniaca, Cerasoides, Cerasus, Laurocerasus, and Prunus. Koehne (1893)
initially divided the genus Prunus into seven subgenera, but later (Koehne, 1911
) recognized four: Amygdalus, Cerasus, Padus, and Prunophora (= Prunus). Rehder (1940)
reviewed the previous treatments and divided the genus into five subgenera: Amygdalus, Cerasus, Laurocerasus, Padus, and Prunophora (= Prunus). Rehder's system was followed by many workers, e.g., Groh and Senn (1940)
, Fernald (1950)
, and Radford, Ahles, and Bell (1968)
. On the other hand, Hutchinson (1964)
recognized three genera (Laurocerasus, Padus, and Prunus s.s.) within Prunus s.l., and synonymized the other previously recognized genera (Aflatunia Vassilcz, Amygdalopsis Carr., Amygdalus, Armeniaca, Cerasus, Emplectocladus Torr., Microcerasus M. Roem., and Prunophora Neck) to Prunus s.s.
The controversy on the classification of Prunus s.l. largely results from the lack of a phylogenetic framework. Previous workers mostly emphasized a few characters, especially fruit morphology, inflorescence type, and leaf duration in formulating the classification (McVaugh, 1951
). The character-based classification systems of Prunus s.l. and the Amygdaloideae need to be evaluated within a phylogenetic framework. A few studies attempted to construct the evolutionary relationships of the genus (e.g., Mowrey and Werner, 1990
; Uematsu, Sasakuma, and Ogihara, 1991
; Badenes and Parfitt, 1995
). But the taxon sampling in these studies was limited. The analysis by Mowrey and Werner (1990)
remains the most comprehensive phylogenetic study of the genus. These workers examined the isozyme variation of 34 species belonging to three (subgenera Amygdalus, Cerasus, and Prunus) of the five subgenera (Rehder, 1940
). This study recognized three groups, which roughly correspond to the three subgenera with a few exceptions.
The present study aims to: (1) construct the phylogenetic relationships among the genera of the subfamily Amygdaloideae; (2) delimit the major evolutionary lineages within Prunus s.l., and (3) evaluate the infrageneric classification of Prunus s.l. Sequences of the internal transcribed spacers (ITS) of nuclear ribosomal DNA (rDNA) were employed for this phylogenetic study.
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MATERIALS AND METHODS |
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TOPABSTRACTINTRODUCTIONMATERIALS AND METHODSRESULTSDISCUSSIONLITERATURE CITED |
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) and all genera of the Amygdaloideae (Thorne, 1992a, b
). Two subspecies of Lyonothamnus floribundus of the Spiraeoideae were used as outgroups. We tried to use Amelanchier and Malus of the Maloideae as additional outgroups, however sequences of these two genera can be hardly aligned with those of the Amygdaloideae.
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. DNA amplifications were performed in 100-µL reactions following Wen and Zimmer (1996)
. The PCR products were purified using millipore columns (Ultrafree-MC Filter Unit, 30 000 NMWL, Millipore, Bedford, Massachusetts, USA). The ITS regions were sequenced using the Cycle Sequencing Kit (188403, Amersham, Cleveland, Ohio, USA) and 33P-dNTPs as radioactive tracers. Four of the six sequencing primers (C5.8S, ITS3, N5.8S, C26A, N-nc18S10, and N18L18; Wen and Zimmer, 1996
) were used to obtain the entire ITS and 5.8S regions from both directions.
The DNA sequences obtained were assembled, and the boundaries between the coding and spacer regions were determined by comparing with the sequences of carrot (Daucus carota L.; Yokota et al., 1989
). The assembled sequences were exported to PAUP* (version 4.0; Swofford, 1999
). Most mutations were base substitutions or deletions, thus allowing manual alignment. All the sequences were deposited at GenBank (see Table 3 for accession numbers).
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) using the maximum parsimony (Swofford et al., 1996
), the maximum likelihood (Felsenstein, 1981
), and the neighbor-joining (Saitou and Nei, 1987
) methods. Parsimony analysis was performed using a heuristic search with MULPARS and furthest addition sequence options. Decay values (Bremer, 1988
) were calculated by using a reverse constraint and comparing the length differences to the maximally parsimonious trees (MPTs). The maximum likelihood analysis was performed with the input order of sequences randomized and the transition/transversion ratio set at 1.8 based on the observed frequencies in the MPTs of the parsimony analysis. A neighbor-joining tree was constructed using the Kimura two-parameter distance (Kimura, 1980
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RESULTS |
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TOPABSTRACTINTRODUCTIONMATERIALS AND METHODSRESULTSDISCUSSIONLITERATURE CITED |
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Of the 662 aligned positions of ITS and 5.8S regions, 218 bases were variable: 114 bases in ITS1, 12 bases in 5.8S, and 92 bases in ITS2. Among the variable sites, 193 positions were phylogenetically informative.
The Kimura two-parameter distances (Table 3) ranged from 0.00149 to 0.08596 among the species of Prunus s.l., whereas they were 0.11387–0.21739 between taxa of the Prunus s.l. and other genera in the Amygdaloideae (Maddenia excepted). Maddenia had a relatively low level of sequence divergence with taxa of Prunus s.l., ranging from 0.02371 to 0.07803. An identical ITS profile was found between P. umbelleta and P. nigra, and Prinsepia sinensis and P. unifolia. No infraspecific variation was found in the two accessions of Prunus americana, whereas the two accessions of P. maximowizii and P. maackii exhibited a distance of 0.00148 and 0.01407, respectively.
Phylogenetic analysis
Treating gaps as new characters, the parsimony analyses generated more than 15 000 MPTs with a length of 630 steps, a consistency index (CI) of 0.632, a retention index (RI) of 0.808, and a rescaled consistency index (RC) of 0.510. The strict consensus tree of 16 383 MPTs is presented in Fig. 1. The analysis of treating gaps as missing data produced congruent topologies with Fig. 1.
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). The latter is paraphyletic. Prunus besseyi and P. tomentosa of sect. Microcerasus of subgen. Cerasus are also nested within the clade of the subgenera Prunus–Amygdalus complex. Within the subgenera Prunus–Amygdalus group, the two species of subgen. Amygdalus sampled are basal. Most species of subgen. Cerasus form a group. Taxa of subgenera Padus and Laurocerasus are mostly basal within Prunus, and taxa of the two subgenera are intermingled with each other. Furthermore, the sectional segregation (indicated by numbers after the subgeneric name in Figs. 1–3) of subgenera Prunus and Cerasus is mostly not supported.
Exochorda, Oemleria, and Prinsepia form a monophyletic group. The relative position of the three genera varied among the parsimony, distance, and the maximum likelihood analyses (cf. Figs. 1–3).
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DISCUSSION |
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TOPABSTRACTINTRODUCTIONMATERIALS AND METHODSRESULTSDISCUSSIONLITERATURE CITED |
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; Ghora and Panigrahi, 1995
) and possesses a dry dehiscent capsule. This genus has a controversial taxonomic position. Traditionally it was placed in the Spiraeoideae (e.g., Bentham and Hooker, 1865
; Focke, 1894
; Rehder, 1940
; Schulze-Menz, 1964
; Cronquist, 1981
). However, as early as 1918, Juel placed Exochorda in the Amygdaloideae, and most recent treatments have allied it to taxa of the Amygdaloideae (Goldblatt, 1976
; Thorne, 1983, 1992a, b
; Takhtajan, 1987, 1997
).
The alliance of Exochorda with the Amygdaloideae has been well supported by several lines of evidence. A cladistic analysis using wood anatomical characters and chromosome numbers (Zhang, 1992
) showed that Exochorda formed a monophyletic group with genera of the Amygdaloideae. This relationship was supported by three synapomorphies (intervessel pit shapes, crystals in ray cells, and same chromosome base number). In Zhang's analysis, Prinsepia was suggested to be basal within the Amygdaloideae—Exochorda clade (Zhang, 1992
). The rbcL phylogeny (Morgan, Soltis, and Robertson, 1994
) also supported the close alliance between Exochorda and the Amygdaloideae. Morgan, Soltis, and Robertson (1994)
reported a sister-group relationship between Exochorda and Oemleria. The NJ tree (Fig. 2) of the ITS phylogeny also suggested this relationship. The close relationship between Exochorda and Oemleria was previously proposed based on the fact that they have five carpels and similar carpel vasculature (Sterling, 1963, 1964a, b, 1966
), and lack stipules at maturity (Morgan, Soltis, and Robertson, 1994
).
The parsimony and the maximum likelihood analyses of the ITS sequences (Figs. 1 and 3) suggest a sister-group relationship between Oemleria and Prinsepia. This relationship is supported by the fact that only Oemleria and Prinsepia of the Amygdaloideae produce cyanogenetic substances (Hegnauer, 1973
). Some workers treated Oemleria and Prinsepia as independent subfamilies (Sterling, 1963
; Hutchinson, 1964
; Mai, 1984
; Kalkman, 1988
). However, the two genera exhibit a relatively low level of ITS sequence divergence (6.848–6.937%). On the other hand, both Oemleria and Prinsepia show a relatively high level of sequence divergence from taxa of Prunus (17.295–23.877%). Thus, treating Oemleria and Prinsepia in separate subfamilies may be inappropriate.
Phylogenetic position of Maddenia
The parsimony and distance analyses of the ITS sequences suggest that Maddenia is nested within Prunus, specifically among taxa of the subgenera Larocerasus–Padus complex (Figs. 1–2). The ML tree shows that it is basal to the Prunus clade (Fig. 3). Maddenia consists of five species from the Himalayas and China (Yü et al., 1986
; Ghora and Panigrahi, 1995
). The generic status of Maddenia is widely accepted (Focke, 1894
; Rehder, 1940
; Hutchinson, 1964
; Kalkman, 1988
; Mabberley, 1997
). Morphologically, Maddenia is similar to subgenera Laurocerasus and Padus in that they all have a racemose inflorescence. It differs from Prunus s.l. in that (1) it has ten sepals (vs. five in Prunus); and (2) it has no or only highly reduced petals (vs. prominent petals in Prunus). Previously the most significant difference of Maddenia from Prunus was thought to be the possession of unisexual flowers in Maddenia (vs. bisexual flowers in Prunus; Hooker and Thomson, 1854)
. Focke (1894)
described Maddenia as dioecious and possessing a two-carpellate ovary. Sterling (1964b)
, however, reported that all specimens of Maddenia under his study had bisexual flowers with a predominance of one carpel. He also showed that Maddenia had similar carpel anatomy and fruit morphology to those of Pygeum, which was synonymized to Prunus (Kalkman, 1965
). Our ITS phylogeny thus corroborates Sterling's observations of a close relationship between Maddenia and Prunus. Additional investigations are needed to confirm that Maddenia should be included in Prunus s.l.
Relationships within Prunus s.l
Rehder (1940)
subdivided Prunus into five subgenera (Table 1). The ITS phylogeny (Figs. 1–3) showed that subgen. Cerasus and subgen. Prunus each generally form a clade. Taxa of subgenera Laurocerasus and Padus are mingled with each other and together form a large basal paraphyletic group. The two species sampled for subgen. Amygdalus form a paraphyletic group basal to the subgen. Prunus clade. Prunus besseyi and P. tomentosa of subgen. Cerasus are nested within the clade of subgenera Amygdalus and Prunus.
The alliance of subgenera Prunus and Amygdalus in the ITS phylogeny is supported by their sharing of several morphological characters. Taxa of both subgenera have a sulcate (grooved or furrowed) endocarp and a glaucous, sometimes pubescent fruit surface. The isozyme study by Mowrey and Werner (1990)
, however, reported that most taxa of subgen. Amygdalus form a group. Only P. triloba of subgen. Amygdalus is grouped with taxa of subgenus Prunus. Our sampling of subgen. Amygdalus is limited at present and additional analysis is planned to evaluate the position of subgen. Amygdalus.
Prunus besseyi and P. tomentosa belong to sect. Microcerasus of subgen. Cerasus (designated as Ce1 in Figs. 1–3) in Rehder's system (1940)
. These two species are nested within the subgenera Prunus and Amygdalus clade. Subgenus Amygdalus and sect. Microcerasus do share several characters such as three axillary buds, sessile or only a short pedicel, and conduplicate leaves. Like taxa of subgen. Amygdalus, P. tomentosa sometimes has pubescent fruits. Mowrey and Werner (1990)
also found P. besseyi and P. tomentosa grouped with taxa of subgen. Prunus and P. triloba of subgen. Amygdalus. The phylogenetic position of taxa in sect. Microcerasus therefore should be reevaluated.
Subgenus Cerasus excluding sect. Microcerasus formed one group with taxa of subgenera Padus and Laurocerasus (Figs. 1–3). This Cerasus–Padus–Laurocerasus complex is somewhat well separated from the taxa of the Prunus–Amygdalus complex morphologically as discussed above. Within the Cerasus–Padus–Laurocerasus complex, taxa of subgen. Cerasus (excluding sect. Microcerasus, and P. mahaleb) form a clade. Subgenus Cerasus differs from subgenera Padus and Laurocerasus in that the former has one to few flowers in an umbellate cluster (vs. a raceme with 12 or more flowers) and each flower has a conspicuous bract.
Subgenus Cerasus was divided into six sections by Koehne (1893)
and Rehder (1940)
with only minor differences between the two systems. In this study, samples of only four sections sensu Rehder (1940)
were included. Prunus mahaleb of sect. Mahaleb is nested within taxa of the subgenera Padus and Laurocerasus. Prunus mahaleb, like taxa of subgenera Padus and Laurocerasus, has a racemose inflorescence and a small bract below each flower. Like other taxa of subgen. Cerasus, P. mahaleb and other species in sect. Mahaleb do have fewer flowers per inflorescence than those in subgenera of Padus and Laurocerasus.
The separation of subgenera Padus and Laurocerasus (Koehne, 1893
; Schneider, 1912
; Rehder, 1940
; Hutchinson, 1964
; Komarov, 1971
) is not supported in this study. The ITS phylogeny shows that species of the two subgenera are intermingled with each other. Morphologically, these two taxa share common characteristics of an elongate raceme with 12 or more flowers, and conduplicate vernation (Koehne, 1893)
. Major differences of the two subgenera are the deciduous habit of subgen. Padus (vs. evergreen of subgen. Laurocerasus) and bracteate racemes in subgen. Padus (vs. bractless racemes in subgen. Laurocerasus). Yü et al. (1986)
added another difference between the two groups: terminal (subgen. Padus) vs. axillary (subgen. Laurocerasus) inflorescence. The lack of congruence of the ITS phylogeny with the traditional delimitation of the two subgenera may be due to character convergence. The differences in deciduous or persistent nature of leaves and the presence or absence of bracts within the Padus–Laurocerasus group may be caused by adaptation to the temperate mesic vs. tropical humid environments, respectively (Webb, 1968
; Davis and Taylor, 1980
; Wilf, 1997
).
Some workers (e.g., Hutchinson, 1964
; Komarov, 1971
; Yü et al., 1986
; Takhtajan, 1997
) recognized several genera within the present concept of Prunus s.l. The monophyly of Prunus s.l. (plus Maddenia) is well supported (Figs. 1–3). The general lack of congruence of the clades in the ITS phylogeny with the narrowly defined genera such as Prunus s.s., Amygdalus, Cerasus, Pygeum, Laurocerasus, and Padus supports the generic concept of a broadly defined Prunus. Furthermore, the ITS sequence divergence within Prunus s.l. is comparable to that of many other plant genera (cf. Baldwin et al., 1995
).
Roughly two major groups are recognizable within Prunus s.l. (Figs. 1–3): (1) the Amygdalus–Prunus–P. besseyi complex, and (2) the paraphyletic Cerasus–Laurocerasus–Padus group. Additional data are needed to arrive at a phylogenetically well-supported infrageneric classification of Prunus s.l.
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FOOTNOTES |
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4 Author for correspondence, current address: Department of Botany, Field Museum of Natural History, Chicago, Illinois 60605 USA (e-mail: jwen{at}fmnh.org
).
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