Further Evidence for Paraphyly of the Celtidaceae from the Chloroplast Gene Matk

Plant Syst. Evol. 228: 107±115 42001) Further evidence for paraphyly of the Celtidaceae from the chloroplast gene matK B.-H. Song1,2, X.-Q. Wang1, F.-Z. Li2, and D.-Y. Hong1 1Laboratory of Systematic and Evolutionary Botany, Institute of Botany, the Chinese Academy of Sciences, Beijing, China 2The Key Laboratory of Plant Stress, Shandong Normal University, Jinan, China Received January 22, 2001 Accepted March 23, 2001 Abstract. Based on the chloroplast matKgene Urticales, have been controversial in taxono- sequence, a phylogenetic analysis of the Urticales my for decades 4Engler and Prantl 1893; in its traditional circumscription and its putative Thorne 1968, 1983, 1989; Dahlgren 1983, anities produced three equally most parsimonious 1989; Cronquist 1981, 1988; Ueda et al. 1997; trees with tree length 1527 steps, CI 0.6863 and Wiegrefe et al. 1998). The arguments mainly RI 0.6352, indicating that the Ulmaceae s. l. are focused on: 4a) Are the Ulmaceae s. l. mono- polyphyletic while the Celtidaceae are paraphyletic, phyletic or polyphyletic? 4b) Are the Celtida- and particularly, Cannabis and Humulus in the Cannabaceae are consistently nested within the ceae and Ulmaceae independent families or Celtidaceae. Therefore, the present data strongly subgroups 4subfamily or subtribe) of the suggest that the Cannabaceae should be merged Ulmaceae s. l.? 4c) What is the relationship with the Celtidaceae to form a monophyletic group. between the Celtidaceae and the other families According to the present study, the Celtidaceae of Urticales? 4d) Where should some problem- including Cannabaceae are more closely related to atic genera such as Gironniera Gaud., Apha- the Moraceae and Urticaceae than to the Ulmaceae nanthe Planch. be assigned to? In the past s. str.. Gironniera and Aphananthe are both basal years, studies on fossil 4Zavada and Crepet clades of the Celtidaceae rather than members of 1981, Manchester 1989), distribution 4Grud- Ulmaceae s. str.. The Rhamnaceae and Rosaceae zinskaya 1967), ¯avonoid chemistry 4Giannasi are the closest relatives of the traditional Urticales, 1978), wood anatomy 4Sweitzer 1971), leaf which is very congruent with the newest system of venation, fruit type and embryology 4Hutch- ¯owering plants put forward by APG. inson 1967), vernation patterns 4Terabayashi Key words: Celtidaceae, Ulmaceae, Rosales, matK, 1991), pollen morphology 4Zavada 1983, Ta- molecular phylogeny. kahashi 1989), seed coat morphology 4Takaso and Tobe 1990), karyomorphology 4Oginuma et al. 1990), phylogenetic analysis 4Zavada and Introduction Kim 1996) and molecular analyses 4Sytsma The two angiosperm families, Celtidaceae and et al. 1996, Ueda et al. 1997, Wiegrefe et al. Ulmaceae, usually treated as two subgroups of 1998) have come to some conclusions: the the Ulmaceae s. l. 4Cronquist 1981) in the Ulmaceae s. l. are polyphyletic rather than 108 B.-H. Song et al.: Further evidence for paraphyly of the Celtidaceae Table 1. The materials Taxa Sources GenBank References Accession No. Ulmaceae Ulmus parvifolia Jacq. Botanic Garden, Institute of AF345321 Present study Botany, Beijing Zelkova schneideriana Botanic Garden, Institute of AF345328 Present study Hand. -Mazz. Botany, Beijing Hemiptelea davidii 4Hance) Botanic Garden, Institute of AF345322 Present study Planch. Botany, Beijing Celtis sinensis Pers. Jinan, Shandong, China AF345316 Present study Pteroceltis tatarinowii Maxim. Jinan, Shandong, China AF345324 Present study Gironniera subaequalis Planch. South China Botanic Garden, AF345319 Present study Guangzhou Aphananthe aspera 4Thunb.) South China Botanic Garden, AF345320 Present study Planch. Guangzhou Trema tomentosa 4Roxb.) Hara South China Botanic Garden, AF345325 Present study Guangzhou Moraceae Morus alba Linn. Botanic Garden, Institute of AF345327 Present study Botany, Beijing Broussonetia papyrifera 4Linn.) Botanic Garden, Institute of AF345326 Present study L'Herit. ex Vent Botany, Beijing Cannabaceae Cannabis sativa L. Jinan, Shandong, China AF345317 Present study Humulus lupulus L. Botanic Garden, Institute of AF345318 Present study Botany, Beijing Urticaceae Boehmeria platanifolia Yantai, Shandong, China AF353579 Present study Franch. et Sav. Malvaceae Hibiscus syriacus L. Botanic Garden, Institute of AF345329 Present study Botany, Beijing Ranunculaceae Cimicifuga acerina Jinfo Mountain, Nanchuan, AF353578 Present study 4Sieb. et Zucc.) Tanaka Sichuan, China Juglandaceae Juglans nigra Linn. U92851 Manos and Steele, 1997 Betulaceae Betula papyrifera Marsh. U92853 Manos and Steele, 1997 Fagaceae Fagus grandifolia Ehrh. U92861 Manos and Steele, 1997 Rhoipteleaceae Rhoiptelea chiliantha Diels & U92852 Manos and Hand. -Mazz. Steele, 1997 Hamamelidaceae Liquidambar orientalis Mill. AF015651 Li et al., 1997 Rosaceae Rosa stellata Wooton AB012000 Unpublished Rhamnaceae Ziziphus obtusifolia Gray AF049848 Unpublished B.-H. Song et al.: Further evidence for paraphyly of the Celtidaceae 109 monophyletic although the Ulmaceae s. str. even distribution at three codon positions are a monophyletic group with little doubt; the 4Wang and Shu 2000; Wang et al. 1998, Celtidaceae and Ulmaceae s. str. should be 2000; Johnson and Soltis 1995). The matK treated as independent families, moreover, the gene has been used eectively in addressing Ulmaceae s. str. are the sister group of the systematic questions in the families Pinaceae Celtidaceae plus all other families in the Urti- 4Wang et al. 2000), Cupressaceae s. l. 4Gadek cales. Nevertheless, there are still some ques- et al. 2000, Kusumi et al. 2000), Taxaceae and tions unresolved, and some new interesting Cephalotaxaceae 4Wang and Shu 2000, Cheng questions appeared. The Cannabaceae, tradi- et al. 2000), Brassicaceae 4Koch et al. 2001), tionally placed in the Moraceae, were nested Saxifragaceae 4Johnson and Soltis 1994, 1995), within the celtidaceous clade while the Celti- Polemoniaceae 4Steel and Vilgalys 1994, John- daceae and Ulmaceae were undoubtedly son and Soltis 1995), and Poaceae 4Hilu et al. included in the Urticales in the molecular 1999). phylogeny 4Sytsma et al. 1996, Ueda et al. Here, we report a matKgene phylogeny of 1997, Wiegrefe et al. 1998). Furthermore, the the Urticales or Rosales, and mainly discuss current system of ¯owering plants put forward the following questions: What should the by APG 41998) implied that the Celtidaceae Celtidaceae and Ulmaceae be assigned to? and Ulmaceae might be closely related to the Are the Celtidaceae monophyletic or paraphy- Rosaceae and Rhamnaceae in the Rosales even letic? How to treat the genera Gironniera and though the putative relatives of the Urticales Aphananthe based on molecular analyses? are still controversial. 4Berg 1977, Cronquist 1981, Dahlgren 1983, Thorne 1983, Judd et al. Materials and methods 1994). Although the previous molecular phyloge- Materials. We sampled three of six genera of the netic studies of the Ulmaceae and their relatives Ulmaceae, ®ve of nine genera of the Celtidaceae, have resulted in signi®cant contributions to the representatives of the other three families 4Mora- recognition of some generic relationships of the ceae, Urticaceae and Cannabaceae) in the Urticales. Ulmaceae s. l., the phylogeny generated from Since the closest relatives of the Urticales, including those studies was not well resolved due to the Hamamelidae 4Cronquist 1981), Malvales and Eu- slow evolution rate of rbcL gene 4Ueda et al. phorbiales 4Dahlgren 1989, Thorne 1989), are still being debated vigorously 4Berg 1977, Dahlgren 1997) or poor sampling of Urticales and its close 1983, Thorne 1983, Judd et al. 1994), and the relatives in the cpDNA restriction site analysis members of Urticales have been merged with of Wiegrefe et al. 41998). Therefore, further Rosales by APG 41998) although previous molec- studies on the molecular phylogeny of the ular analyses 4Chase et al. 1993, Gunter et al. 1994, Celtidaceae and Ulmaceae or Urticales are still APG 1998, Swensen 1996) and morphological data needed. Among the dierent markers frequent- 4Huord 1992) suggest that Rhamnaceae and ly used in recent molecular systematics, the Rosaceae are better outgroup candidates, we also matKgene 4 1500 bp size) which is located sampled a representative from each of the two within the intron of the chloroplast gene trnK orders Malvales 4Hibiscus) and Ranunculales 4Cimi- has been demonstrated to be highly potential cifuga), and used Cimicifuga acerina 4Ranuncula- for providing insight into evolutionary and ceae) as the outgroup. Additional seven matKgene systematic problems 4Hilu and Liang 1997). sequences used in the present study were recovered from GenBank 4Table 1). Particularly, at lower taxonomic levels, such as DNA extraction, ampli®cation and sequencing. genus, family, the matKgene provides better Total DNA isolation followed the protocol of resolution than the most widely used chlorop- Rogers and Bendich 41988), using fresh or silica gel last gene rbcL. Moreover, it has been revealed dried leaf tissue from a single individual. that the variation of the matKgene has similar Polymerase Chain Reaction 4PCR) was used to rate in closely related lineages and a relatively amplify the matKgene with the primers 110 B.-H. Song et al.: Further evidence for paraphyly of the Celtidaceae Table 2. Pairwise distances between taxa Below diagonal: Total character dierences. Above diagonal: Mean character dierences 12345678910 1 Ulmus ± 0.01327 0.04018 0.11823 0.11715 0.10610 0.09643 0.11799 0.10033 0.10786 2 Zelkova 16 ± 0.04975 0.12935 0.12050 0.11000

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