Molecular Phylogenetics and Evolution Vol. 16, No. 1, July, pp. 96 –112, 2000 doi:10.1006/mpev.1999.

0769, available online at on

The Phylogeny of the Asteridae sensu lato Based on Chloroplast ndhF Gene Sequences
Richard G. Olmstead,* Ki-Joong Kim,† ,1 Robert K. Jansen,† and Steven J. Wagstaff‡
*Department of Botany, University of Washington, Seattle, Washington 98195; †Section of Integrative Biology and Institute of Cellular and Molecular Biology, University of Texas, Austin, Texas 78712; and ‡Manaaki Whenua, Landcare Research, P.O. Box 69, Lincoln 8152, New Zealand
Received July 23, 1999; revised December 21, 1999

A phylogenetic study of Asteridae sensu lato was conducted based on chloroplast ndhF gene sequences for 116 ingroup and 13 outgroup species. Prior molecular studies based on rbcL sequences identified terminal groups corresponding to families, but were unable to resolve relationships among them. These results are largely consistent with earlier rbcL studies, but provide much greater resolution and stronger bootstrap support throughout the tree. The parsimony analysis found eight equally parsimonious trees, all of which recognize four major clades with the following relationship: (Cornales (Ericales (Euasterids I, Euasterids II))). Euasterids I includes (Garryales ((Solanales, Boraginaceae) (Gentianales, Lamiales))), although with weak support for relationships among the named clades. Euasterids II includes (Aquifoliales (Asterales (Apiales, Dipsacales))) with strong support for these relationships. Relationships within Ericales are weakly supported and merit further attention. © 2000
Academic Press

Key Words: Asteridae; phylogeny; ndhF; angiosperm classification.

INTRODUCTION A growing body of molecular evidence has accrued to support the recognition of a major lineage of eudicot angiosperms called the Asteridae sensu lato (Olmstead et al., 1992, 1993; Chase et al., 1993; Soltis et al., 1997). The Asteridae sensu lato (henceforth simply called Asteridae) includes the families placed in the subclass of the same name by Cronquist (1981), plus many other families in his system assigned to subclasses Dilleniidae, Hamamelideae, and Rosideae, and comprises approximately one-third of all angiosperm species. This group is characterized predominantly by the presence of unitegmic and tenuinucellar ovules and the widespread occurrence of iridoid compounds, which seem
Present address: Department of Biology, Young-Nam University, Kyung San City, Kyung Buk, South Korea.
1055-7903/00 $35.00 Copyright © 2000 by Academic Press All rights of reproduction in any form reserved.

likely to represent synapomorphies for the group (Olmstead et al., 1993). Another common and widespread characteristic shared by members of this group is the occurrence of fused corollas. Ontogenetic studies have shown that there are two developmental paths to floral fusion, “early sympetaly” and “late sympetaly” (Erbar and Liens, 1996). Polypetalous representatives examined exhibit early sympetaly, suggesting that corolla fusion may be another synapomorphy of the group (Erbar and Liens, 1996). Initial molecular systematic studies of the Asteridae based on rbcL sequences succeeded in circumscribing the group and identifying its major lineages (Olmstead et al., 1992, 1993; Chase et al., 1993). However, the degree of resolution among those lineages was limited and the strength of support for some of the identified groups was weak. It was clear that additional data would be needed to confirm the monophyly of the lineages identified by rbcL studies, to determine the relationships among those lineages, and to make recommendations for changes in classification that reflect our current understanding of the phylogeny of this group. This study of Asteridae phylogeny based on sequences of the chloroplast gene ndhF is parallel to those based on rbcL. Initial applications of ndhF sequences to systematic studies centered on relationships within families, making use of its greater length and higher average substitution rate to generate ca. three times as many parsimonyinformative characters (Olmstead and Sweere, 1994; Scotland et al., 1995; Clark et al., 1995; Kim and Jansen, 1995; Olmstead and Reeves, 1995; Scotland et al., 1995; Neyland and Urbatsch, 1996; Catalan et al., 1997; Terry et al., 1997a,b). As more sequences from different families became available, it became apparent that ndhF sequences would have application to phylogenetic questions at much greater phylogenetic depth than just within families (Olmstead et al., 1998). The sampling for this study is similar to that of previous rbcL studies with respect to family diversity, but is not universally parallel at the generic level; therefore, combining these data with

Of these. Also. and 627 were third codon positions.250. 319 were variable but uninformative for parsimony analysis. Olmstead and Reeves. and 1350 were parsimony informative. however. PCR and DNA sequencing methods were as described in Olmstead and Sweere (1994).379 steps (CI ϭ 0.0 (vers.532). Rice et al. and Kim and Jansen (1995). 1) encompasses the range of possible most-parsimonious tree topologies.303. ordinallevel classification of angiosperms (Angiosperm Phylogeny Group. in PAUP*). which were included in the data set as binary characters. (1993). in which 105 rbcL sequences were analyzed. The goals of this study were to corroborate the results of prior molecular studies based on rbcL sequences and to achieve greater resolution of relationships among groups within Asteridae. in the study of Olmstead et al. A 152-nucleotide region in the middle of the Hydrostachys sequence could not be aligned readily with the other sequences.551) than among first (CI ϭ 0. No shorter trees or additional trees at the same length were found using the inverse constraint procedure.600 vs 0. The gap characters. (1993) was conducted using the same parameters as for the ndhF analysis to provide bootstrap support values for comparison between studies for clades found by both. The strategy for parsimony analysis enabled the discovery of multiple islands. so that 23 nucleotides (nt) at the 5Ј end of the gene and 108 nt at the 3Ј end of the gene were not obtained for many sequences (Olmstead and Sweere. 2 to illustrate representative branch lengths (using accelerated transformation optimization. 1998). 1995). the character defined by the shorter gap was scored as unknown for the taxon with the longer gap (see Table 1 for further information about individual gap characters). lending further support to the inference that the strict consensus tree of the eight trees (Fig. RESULTS The PCR strategies used by the two collaborating labs differed in the proportion of the ndhF gene sequence obtained. including gaps.534).213. Parsimony analyses were conducted with all nucleotide and gap changes weighted equally using the computer program PAUP* 4. each with a random taxon addition starting tree with TBR branch swapping and MULTREES on.. . 417 occupied first codon positions. 1994) and adjusted by eye. 785 were invariant among the sampled taxa. ACCTRANS. consistency with a representative shortest tree depicted in Fig. Olmstead and Reeves (1995). The total aligned sequence length. 1995). One of the most-parsimonious trees is depicted in Fig. making a total of 1374 cladistic characters. For comparison with rbcL. given these data. A total of eight trees representing two islands of four trees each was recovered with lengths of 13.251. The results presented here (along with many other studies) already have contributed to a newly proposed. MATERIALS AND METHODS This study includes ndhF sequences for 129 angiosperm species.882 vs 0. 94 sequences are first reported in this study (see Appendix 1 for GenBank Accession Nos. there were only 495 parsimony-informative nt characters and no gap characters. RI ϭ 0. These regions and occasional missing data points in the middle of sequences account for approximately 2% missing data. A total of 100 replicate tree searches was undertaken. 1997. Initial sequence alignments were constructed with the assistance of the computer program CLUSTAL W (Thompson et al. Gaps provide partial support for 19 clades. 1994. Of the informative positions. 1999).CHLOROPLAST PHYLOGENY OF THE ASTERIDAE 97 data from previous studies is inadvisable until considerably more work is done to fill in gaps in both data sets. including 116 species of Asteridae s. including 24 shared by two or more taxa. Bootstrap analysis with 500 replicates was carried out using a heuristic search with TBR branch swapping and MULTREES off (DeBry and Olmstead. was 2454 nucleotides. Primers external to the gene were used in the Jansen lab (and for some sequences in the Olmstead lab). Swofford. 500 replicate heuristic searches were undertaken using the strict consensus tree of the mostparsimonious trees as a constraint. Many of the plant accessions and 35 of the ndhF sequences have been used in previously published molecular systematic studies.l. RI ϭ 0.286. RI ϭ 0. a bootstrap analysis of the rbcL data set of Olmstead et al. so that the entire sequence of the gene was obtained (Kim and Jansen. RI ϭ 531) codon positions. so that only those trees that did not match the constraint were retained in an effort to find shorter or other equal-length trees (Catalan et al.. whereas internal primers located near the ends of the gene were used for most taxa in the Olmstead lab.. were found to be more highly consistent with the shortest tree topology than were the nucleotide characters (CI ϭ 0. though few in number. A total of 82 gaps were inserted to align the sequences (Table 1).521) or third (CI ϭ 0. Subsequent to these searches. In cases in which gaps of different length shared a start or end point or in cases in which one gap entirely overlapped the other in different taxa. Among the nucleotide characters. and 13 outgroup species (Appendix 1). 306 were second codon positions. 1997).). so was excluded from analysis. The aligned sequences begin at the start codon and end at the location of the stop codon in Asteraceae (6 nt before the stop codon in Nicotiana and many other taxa). Gaps shared by two or more taxa were scored as additional binary characters. 2000). or only 36% as many characters as ndhF provided for this study. 16 of which have bootstrap values of 97% or greater. RI ϭ 0. representing a phylogenetic distribution very similar to that represented here. 2 was greater on average among second codon positions (CI ϭ 0. b2.

Rhododendron. Hedera. Boopis. Pentas Gelsemium Phlox Veronica. Villarsia. Symphoricarpus. Exacum Ipomoea Clerodendrum. Phyllonoma. Styrax. Impatiens. Corokia. Campanula. Quercus. Hydrangea. Ilex. Phyllonoma Veronica Acer Justicia Gentiana. Panax. Griselinia. Carpenteria. Halesia. Lonicera. Scaevola. Angelica. Hydrostachys. Plantago. Cercidiphyllum. Viburnum Boopis Schizanthus Paeonia Phlox. Hedera. Helwingia. Griselinia. Dasyphyllum. Eucnide Gentiana. Panax Cephalanthus. Cornus. TABLE 1 Asteridae ndhF Sequence Alignment Gaps Gap a 1 2A 3A 4A 5A 6A 7A 8A 9A 10 A 11 A 12 13 14 15 16 17 Character No. Erithalis Pentaphragma Anagallis Selago Gelsemium Montinia Aquilegia. Schizanthus Impatiens Callitriche Sambucus Pentas Anagallis Garrya. Lycopersicon. Heuchera. Digitalis. Succisa. Camellia. Petunia. Nolana. Phyllonoma. Alangium. Exacum. Lobelia. Dipsacus. Abelia. Helwingia. Valeriana.98 OLMSTEAD ET AL. Viburnum. Gerbera. Pittosporum. Eucnide. Sambucus. Ajuga Impatiens Sambucus. Carpenteria. Guettarda. Rhamnus. Anagallis. Nicotiana. Stylidium. Codonopsis. Cyananthus. Trachelium. Aster. Scabiosa. Osmorhiza Borago Selago Phytolacca Ilex. Lactuca. b Location 657/658 1371–1394 1393–1398 1396–1404 1401/1402 1401–1403 1415–1420 1426/1427 1439/1440 1440/1441 1448/1449 1449–1452 1449–1488 1460/1461 1462/1463 1467–1475 1467/1468 Size c 6 24 6 9 6 3 6 6 3 6 9 12 c 42 c 3 3 15 9 Polarity Insertion Deletion Deletion Deletion Insertion Deletion Deletion Insertion Insertion Insertion Deletion Deletion Deletion Insertion Insertion Deletion Insertion Taxa Codonanthe Hydrolea Dicentra Acanthopanax. Pentaphragma. Rogiera. Diospyros. Nymphoides. Dampiera. Acanthopanax. Phytolacca Ipomoea Rhododendron Quercus Spigelia 1 2 3 41 18 19 B 20 B 21 B 22 B 23 B 24 B 25 B 26 B 27 B 28 B 29 B 30 B 31 B 32 B 33 B 34 B 35 B 36 B 37 B 38 B 39 B 40 B 41 B 42 B 43 B 44 B 45 B 51 61 71 82 1467/1468 1468–1475 1470–1478 1470/1471 1475/1476 1479/1480 1476–1484 1482–1484 1485/1486 1486–1488 1486–1491 1488–1490 1489–1494 1494/1495 1495–1500 1498–1506 1498–1509 1504–1506 1504–1509 1506–1514 1513–1521 1515/1516 1518–1526 1519/1520 1525–1590 1527/1528 1533/1534 1534–1536 3 9 9 6 15 6 9 3 6 3 6 3 6 6 6 9 12 3 6 9 9 21 9 6 66 3 3 3 Insertion Deletion Deletion Insertion Insertion Insertion Deletion Deletion Insertion Deletion Deletion Deletion Deletion Insertion Deletion Deletion Deletion Deletion Deletion Deletion Deletion Insertion Deletion Insertion Deletion Insertion Insertion Deletion 9 10 11 12 13 3 . Coriandrum. Phlox. Helianthus. Barnadesia. Menyanthes. Paeonia Aquilegia Hellwingia. Tetraclea. Eucnide Lobelia Ilex. Acer.

Tetraclea. Buddleja. due to overlap with deletion 62. missing for Liriodendron and Magnolia. Codonopsis. due to overlap with 70. . for the complete sequence of Hydrostachys. 1986.D 62 E 63 F 64 E 65 E 66 E 67 E 68 E. 3. Selago. Drymonia Montinia. as corrected by Olmstead et al. due to unalignable region in Hydrostachys. Verbascum. Thunbergia. to indicate the actual size of the deletion. Aquilegia. missing for Callitriche due to overlap with deletion 13. Campanula. wherein gaps are coded 0/1 for absence and presence of sequence. Lantana. missing for Boopis and Paeonia due to overlap with deletions 38 and 40. the best fit alignment (not included in the PAUP data set) suggests a 3-nt insertion somewhere in the 6-nt gap in 37.F 75 G 76 G 77 78 79 80 81 82 15 5 16 4. Paulownia. 6. C. 5. Codonanthe. positions 1468 –1582 in Nicotiana.F 73 E. Retzia. Digitalis. 9.F 70 E. Character number refers to the phylogenetic analysis. Helianthus.F 74 E. missing for Ipomoea.5 6 1544/1545 1546–1551 1549–1554 1551/1552 1557/1558 1558–1560 1560/1561 1561–1566 1567–1569 1569/1570 1569/1570 1581/1582 1585–1587 1603–1608 1686–1688 1687/1688 1690–1698 1691–1699 1691/1692 1693–1698 1695/1696 1699–1707 1700–1708 1701/1702 1703/1704 1703–1708 1705–1710 1743–1766 1759–1764 1878–1886 1904/1905 1906/1907 1911/1912 1912/1913 1926/1927 18 6 6 9 6 3 6 6 6c 6 3 6 3 6 3 9 9 9 6 6 6 9 9 6 6 6 6 24 6 9 6 6 6 6 6 Insertion Deletion Deletion Insertion Insertion Deletion Insertion Deletion Deletion Insertion Insertion Insertion Deletion Deletion Deletion Insertion Deletion Deletion Insertion Deletion Insertion Deletion Deletion Insertion Insertion Deletion Deletion Deletion Deletion Deletion Insertion Insertion Insertion Insertion Insertion 17 7 18 19 8 20 8 21 22 23 9 24 Note. Ajuga. and Lactuca. Jasminum. F. 1993). Magnolia Gentiana. missing for Aster.F 71 E. Cyananthus. respectively. Vitex. Drymonia. b Character scoring for taxa with completely overlapping gaps is as follows: 1. missing for Ipomoea due to overlap with deletion 42. Borago Justicia Pentaphragma Ipomoea Lactuca Aquilegia Pentas Camellia Justicia Rhamnus. Campsis. Location of gaps is in reference to the ndhF sequence of Nicotiana tabacum (Shinozaki et al. Nematanthus. Justicia.F 72 E. but alignment difficulties make coding this region as missing a conservative choice). respectively. due to overlap with 56. Codonanthe. Olea. Quercus. Callitriche. Nematanthus. Phacelia.CHLOROPLAST PHYLOGENY OF THE ASTERIDAE 99 TABLE 1—Continued Gap a 46 47 B B Character No. Helianthus. Aloysia. due to overlap with deletion 64. Mentha. Ligustrum Aster. Exacum Hydrostachys (in hypervariable region not included in analysis) Dicentra Staphylea Hydrolea Ipomoea Plantago. 7. Phyla. missing for Callitriche and Lobelia due to overlap with deletions 13 and 29. Stylidium. D. G. Exacum. Callitriche. Veronica. Dicentra. Hydrostachys (the complete sequence for Hydrostachys has been determined through this region. A “–” indicates the range of nucleotides deleted and a “/” indicates the location of an insertion relative to the sequence of Nicotiana.. Lactuca Phyllonoma Plantago Hydrophyllum. 8. c Includes a portion of a gap in Nicotiana. Cyrtandra Cephalanthus Cyananthus Acer Rhamnus Vitex Cyrtandra Succisa Cercidiphyllum Liriodendron. Rhododendron.. Scrophularia. 2. Euthystachys. Hydrolea Mentha Dipsacus. Tecoma. B. Sesamum. 4. a Missing sequence in the region of these gaps as follows: A. Scabiosa. b 14 4 Location 1535/1536 1544/1545 Size c 6 6 Polarity Insertion Insertion Taxa Sambucus Clerodendrum. Catalpa.F 69 E. respectively. missing for Hydrostachys. Acer 48 B 49 B 50 B 51 B 52 B 53 B 54 B 55 B 56 B 57 B 58 B 59 B 60 61 C. Lobelia Gentiana Nyctanthes. Sizes of deletions that overlap insertions in the alignment for other taxa are exclusive of the insertion length. missing for Montinia and Hydrolea. E. Myoporum. Succisa Trachelium.

FIG. DISCUSSION These results agree with those of previous cpDNA studies in finding a monophyletic Asteridae (Olmstead et al. 1993. 1. 1992. although the outgroup sampling was not designed for a rigorous test of this hypothesis. Cornales. Numbers above branches indicate bootstrap support values (numbers in parentheses in (a) represent values when Hydrostachys is not included). Classification follows the Angiosperm Phylogeny Group (1998) ordinal classification.. Alignment of the most remote out- .100 OLMSTEAD ET AL.. 1993). Strict consensus tree of eight most-parsimonious trees based on ndhF sequences. and Euasterids II. Shaded boxes enclose the Euasterid I and Euasterid II clades. (a) Outgroups. Ericales. Chase et al. (b) Euasterids I.

1994) and evidence suggests that it may be one in much of the rest of Convolvulaceae also (R. For this reason. Stefanovic. but they should be sufficient to root the asterid portion of the tree. Ipomoea had four unique deletions. Olmstead and S. including one of 66 nt in length. . some of the outgroup relationships may be questionable (although relationships among outgroups are consistent with rbcL results wherever bootstrap values are Ͼ50%). 1—Continued group sequences with the rest of the sequences was problematic in places. but alignment of outgroup sequences closest to Asteridae was straightforward. Sequences of two taxa (Ipomoea and Hydrostachys) were particularly divergent.CHLOROPLAST PHYLOGENY OF THE ASTERIDAE 101 FIG. ndhF is known to be a pseudogene in Cuscuta (Haberhausen and Zetsche.

including a region in the middle of the sequence that could not be aligned with confidence. for which Ipomoea exhibits no rate acceleration.102 OLMSTEAD ET AL. However. insertions. The Hydrostachys sequence was ex- tremely divergent. Cornales. (a) Outgroups. Hydrostachys also . and deletions (scale lower right). and Euasterids II. One of the eight most-parsimonious trees based on ndhF sequences drawn with branch lengths proportional to the number of inferred nucleotide substitutions. FIG. hollow bars indicate homoplastic insertions and deletions. 2. unpublished). The placement of Ipomoea as sister to the Solanaceae is consistent with published results based on rbcL. Ericales. it did not contain excessive deletions or any indication of being a pseudogene. Solid bars indicate nonhomoplastic insertions and deletions. (b) Euasterids I.

1993). 1998). but are more fully . Olmstead et al. 1994) and 18S rDNA (Albach. who cited cpDNA studies (e. 1993) and other. However.g.CHLOROPLAST PHYLOGENY OF THE ASTERIDAE 103 FIG. Chase et al..g. unpublished studies of 18S sequences that did not find that result. Also.. finding Caryophyllidae to be nested within Asteridae was considered anomalous by the authors of that study (Soltis et al. Within Asteridae the relationships inferred from ndhF sequences are highly consistent with prior rbcL results (e. 2—Continued exhibits highly divergent sequences for rbcL (Hempel et al... 1997)... 1998). which found Caryophyllidae (including Phytolacca) to be nested within Asteridae (Soltis et al. One member of Caryophyllidae (Phytolacca) was included in the outgroup sampling and fell outside the Asteridae.. 1997). in contrast to the results of an 18S rDNA analysis of angiosperms. combined cpDNA and 18S rDNA analyses support the exclusion of Caryophyllidae from Asteridae (Soltis et al..

1997. resolved and typically with greater support for clades found therein (Fig. Ebenales. (1993) within the Asteridae. 1981. The strength of support for the inclusion of Hydrostachys within Cornales cannot be considered strong on the basis of ndhF sequences alone.. There is moderate bootstrap support for this clade (64%). Kron. with weak support for any of the possible relationships among them. Taxon sampling in the rbcL study was different from that of this study. 3. 1). Cullings. and Asteridae (Hydrostachyaceae). but the remainder of the clade. indicating that the position of Cornales as the basal-most branch is not firmly resolved. The FIG. Euasterids I. 1997). A study of Ebenales (Morton et al. Most of the plant families belonging to the Ericales (sensu Angiosperm Phylogeny Group. Polemoniaceae (Steele and Vilgalys. 1997). Additional molecular systematic studies within this group have focused on Cornaceae. 1998). Balsaminaceae).. 1996. Convolvulaceae. Morton et al. bootstrap val- . as well as differences in relationships. 1980) into two groups. 1993.. so.. but support for the monophyly of Euasterids with Ericales is weak (30%). 1995). Johnson and Soltis. 1993. not previously circumscribed in any authority-based classification (e. 1993.. Boraginaceae. The group is weakly supported here (31%). 1996). This clade is the most poorly represented of the four major clades in our study. Hydrangeaceae (Soltis et al. However. but sampling is limited in some groups (e. 1998) were assigned to subclass Dilleniidae. 1998). 1993) and ndhF trees showing bootstrap support for the Asteridae and the 11 clades recognized by Olmstead et al. Solanales. Hydrangeaceae).. that are paraphyletic with respect to Gentianales and Solanales and the inclusion of Caryophyllidae within Ericales. 1997). 1995).. circumscribed here to include Solanaceae. 1994. 1). respectively).. Cornales. 1993. which correspond roughly to the Euasterids I and Euasterids II (in part)..g. 1997.104 OLMSTEAD ET AL. Johnson et al. 1 and discussed below is consistent with the newly proposed classification of the Angiosperm Phylogeny Group (1998). 1). Gentianales. 1994. rbcL sequences also place Hydrostachys in this clade close to Hydrangeaceae with a similarly long branch (Hempel et al. Ericales. Cornales) and there are numerous small differences in circumscription. Theales. and Sarraceniaceae (Bayer et al. and see Grant. In many bootstrap samples.. including all or part of the orders Diapensiales. Euasterids I. which may account for some of the difference in resolution and magnitude of bootstrap support. but relatively weak support for relationships within the clade (Fig. Olmstead et al.. Ericales. 1998). are weakly supported (25%). 1996. Four main groups. and Violales (Cronquist. This clade is the smallest of the four major clades and contains families assigned by Cronquist (1981) to the Rosidae (Cornaceae. Polemoniaceae) and Rosidae (e. and Lamiales. bootstrap support was not applicable. Thorne. Four main groups are identified within this portion of the clade. probably due to the numerous alternative placements of Hydrostachys in various bootstrap replicates.. but the Solanales. 1996) have helped resolve relationships within those families.. Fig. with Hydrostachys removed from the analysis. due in part to the very long terminal branch leading to Hydrostachys.. comprise the Asteridae.. The classification depicted in Fig. Hydrostachys falls outside the Cornales. Numbers in parentheses in the ndhF tree represent bootstrap values when Hydrostachys is not included (Hydrostachys was not included in the rbcL analysis). with Garryales (represented only by Garrya here) as a basal branch (consistent with rbcL evidence. Cronquist. Olmstead et al. 1992). 1993.g. exclusive of Garryales.. Takhtajan (1987) split his former Asteridae (Takhtajan..g. However. 1996. Euasterids I is supported moderately (53%). Cornales. and Loasaceae (Hempel et al. Molecular systematic studies of Ericaceae (Kron and Chase. including Nyssaceae and Alangiaceae (Xiang et al. The 18S rDNA sequences find similar groups (Soltis et al. a pattern observed in rbcL studies as well (Kron et al. The Gentianales and Lamiales each are well supported (95 and 100%. as do values for major divisions in the Asteridae. 1997). 1995) and 18S rDNA sequences suggest a similar placement near Hydrangeaceae (Albach. Lecythidiaceae (Morton et al.. Rice et al. Nepenthales. Ericales... respectively. Dilleniideae (Loasaceae). Porter. but relationships among families remain poorly known. 1981). Kron and King. Chase et al. 1996) has shown that the traditionally circumscribed order is polyphyletic. 1993. is more strongly supported (75%. and Euasterids II. Lamiidae and Asteridae. and a small clade represented by Montinia and Hydrolea in this study. 1995. ues for all of the more inclusive clades in the Cornales rise dramatically (Fig. including Lamiales. with a few elements from the Asteridae (e. Only one taxon was included in Garryales.g. 3). Support for the monophyly of a group composed of the two Euasterid groups is moderate (70% bootstrap support). Primulales. Reduced cladograms of rbcL (Olmstead et al.

Cosner et al. Hippuridaceae. (1994) based on rbcL and includes Sphenocleaceae (Cosner et al. and Phrymaceae (Wagstaff and Olmstead... Fay et al.. unpublished).. Verbenaceae (100%). with the exception that ndhF strongly supports (88%) a relationship between the Loganiaceae and the Gentianaceae (Fig. The most thorough previous study of the Euasterids I clade based on rbcL was unable to resolve the placement of the Boraginaceae and recognized a separate Boraginales (Olmstead et al. The location of Gesneriaceae near the base of the Lamiales is consistent with the results of other molecular studies (Olmstead and Reeves. 1995. 1997. Olmstead et al. Gesneriaceae form a well-supported clade (100%). Soltis et al.. Olmstead. Olmstead et al. 1991. The large family Rubiaceae has been subjected to several molecular systematic studies (Bremer and Jansen. Bremer. However. Scotland et al. but comparable work is less advanced in Convolvulaceae (S. Molecular studies provide firm evidence to support the inclusion of Hydrophyllaceae within Boraginaceae (Ferguson. and Oleaceae (Kim and Jansen. Chase et al.. 1993. can be identified (dePamphilis et al. and Plantaginaceae (in agreement with Olmstead and Reeves. Support for monophyly of Solanaceae with Convolvulaceae is modest (52%)...l. Molecular studies have contributed to the realignment of the Lamiaceae and Verbenaceae (Cantino et al.. Oxelman et al. 1999). Andreasen et al.. 1997). including Retzia. 1999. Molecular studies have resolved the placement of several genera sometimes assigned to the Gentianales to other orders.. resolution of interfamilial relationships in the rest of the Lamiales is poor. McDade and Moody. Support for the order as a monophyletic group (Fig. 1997). Oxelman. a family with 1 species in New Zealand and 1 in Patagonia. 1992. which forms a monophyletic group (100%) with Buddlejaceae (Oxelman et al. 1999). which forms the sister group to the rest of the clade (99%). hampered by the difficulty in obtaining ndhF sequences in the family (R. 1997. 1 and 2).. Thevetia) and separation of Gelsemiaceae from Loganiaceae. Tetrachondraceae. Matelia) within Apocynaceae (Nerium. including Lamiaceae (99%). Spooner et al. Olmstead and D. (79%). Relationships also are congruent with those based on rbcL (Olmstead et al. Sennblad and Bremer. Bremer et al. 1995. Cyclocheilaceae (Nesogenaceae in Wagstaff and Olmstead. but this stems in part from the highly divergent ndhF sequence of Ipomoea (Figs. Representatives of Scrophulariaceae included here form two well-supported groups. South America) are not included here. 1994. with Verbenaceae restricted to the former subfamily Verbenoideae and with most of the rest of Verbenaceae and Symphoremaceae transferred to Lamiaceae.. 1992. 1) and based on rbcL (Cosner et al. 2000).. 1994. 1997.. Scrophulariaceae s. albeit weakly in all cases. 1995. 1993. 1997. unpublished). 1993). 3). Wolfe and dePamphilis.. unpublished). Wagstaff and Olmstead. 1995. unpublished). as is support for monophyly of Oleaceae (100%). 1998. Young et al.. Wagstaff et al. presented here (Fig. Bremer et al. R. and Polypremum (monotypic.. 1) is strong (100%)...g. also in Montiniaceae (B. 1994. 1997. which includes Callitrichaceae. Stefanovic and R. Olmstead et al. 1999). in agreement with molecular studies of the family (Smith et al. Mione et al. . The Lamiales are the largest order within the Euasterids I. Nematanthus.CHLOROPLAST PHYLOGENY OF THE ASTERIDAE 105 Boraginaceae (including Hydrophyllaceae) are well supported (97%) but weakly placed as sister to the Solanales (15%). The classification adopted by the Angiosperm Phylogeny Group (1998) takes a conservative approach. Several molecular systematic studies have substantially resolved relationships within Solanaceae (Olmstead and Palmer.. (Bremer et al.. 1998. 1993. Bignoniaceae (97%). 1992. 1998) and relationships within the family based on limited sampling here are congruent with those based on greater sampling with rbcL (Bremer et al. 1993.. 1998).. All prior studies of rbcL and 18S rDNA sequences support monophyly of Solanaceae plus Convolvulaceae (Olmstead et al... et al. 1994) and Grevea.. Nickrent et al. 1998.b). Hedren et al. Wallander and Albert.. 1995. etc. 1995.. but may belong at or near the base of the order (Wagstaff and Olmstead. and Stilbaceae (100%). with approximately 23 families and over 20. Desfontainia. Buddlejaceae (100%). Acanthaceae (80%). 1997). 1996). Olmstead and Sweere. 1997). 1995) in finding Rubiaceae as sister to the rest of the order and in finding poorly resolved relationships among the other families. 1994.. 1993. 1994. Individual families are well resolved.. 1). and Veronicaceae (97%). including the achlorophylous holoparasites conventionally assigned to the family and the green hemiparasites assigned to Scrophulariaceae s. Oxelman et al. 1995. Chase et al. several other studies of rbcL sequences with different taxonomic sampling have suggested a close relationship with the Solanales (e. ndhF results support inclusion of Asclepiadaceae (Araujia.. 1993. The clade composed of Montinia (Montiniaceae) and Hydrolea (Hydroleaceae) was first identified by Cosner et al. 1997. Olmstead. 1992.. Ferguson.. 1994) provide weak support (25%) for the monophyly of the Montinia clade with the Solanaceae and Convolvulaceae.. unpublished). Bremer. Within Gentianales. Buddleja.000 species. Cryptostegia. leaving Boraginaceae unassigned to an order within the Euasterids I. Evidence from ndhF.s. Bignoniaceae (Spangler and Olmstead. 1995). Manen et al. as indicated by rbcL studies (Olmstead et al. Cosner et al.. with Cyrtandroideae (Cyrtandra) and Gesnerioideae (Codonanthe. 1998). 1992. Steane et al.. 1999). 1997a.. Other families that have been subject to molecular systematic study include Acanthaceae (Scotland et al. Support for this clade is stronger with rbcL (Fig. 1998. However.. 1998. 1998. 1999). Additional work on Scrophulariaceae suggests that a third major group.. Wolfe et al. including Avicenniaceae. 1999). Wagstaff and Olmstead. 1997. Drymonia) forming sister groups. Reeves and Olmstead. Molecular data support the separation of several small families (not represented here) often assigned to Verbenaceae. Orobanchaceae. 1995. 1995). 1998.

with a few exceptions. and Grant DEB-9020171 to R. 1996) and other studies have focused on the relationships among families most closely related to Asteraceae (Michaels et al. Other studies of rbcL (Olmstead et al. . 1993. Jansen and Kim. Eriksson and Donoghue. All families with sampling of two or more species were strongly supported monophyletic groups. 1997. which together form the sister group to Asteraceae (Olmstead et al. Olmstead and Reeves. Reeves and two anonymous reviewers for helpful comments. 1993. which make up most of the Apiales. Thanks to the many individuals and Botanical Gardens who helped by providing plant tissue or DNA for analysis. 1993. Gustafsson et al. subfamily Hydrocotyloideae (Apiaceae). Aquifoliaceae and Helwingiaceae are strongly supported (92%).g.106 OLMSTEAD ET AL. 1997) data sets find that this relationship is unresolved. ndhF results concur with rbcL studies (Donoghue et al.. relationships among orders in this clade are clear. 1995. The congruence between the phylogenetic results from these two gene sequences provides a foundation for a revised classification of Asteridae at the ordinal level (Angiosperm Phylogeny Group. M. and Phyllonomaceae. 1997). 1) also is supported moderately (68%) and comprises four well-supported orders: Apiales (100%). monophyly of Calyceraceae with Asteraceae is strongly supported (98%) and the family Goodeniaceae is sister to that clade (Fig. 1996.. was not sampled in this study. Helwingiaceae. Eriksson. Euasterids II. However. Most studies based on rbcL supported monophyly of Calyceraceae with Goodeniaceae. Kim and Jansen.J.. in this study. Backlund and Bremer. However. The Caprifoliaceae without Sambucus and Viburnum still are paraphyletic with respect to Dipsacaceae and Valerianaceae. 1996. in addition to the shared characteristic of epiphyllous inflorescences. and Jansen and Kim (1996). 1993. and T. 1995. which has been found to be basal and paraphyletic (Plunkett et al. which is sister to Asterales (90% support for the inclusive clade). whereas both alternate relationships have been suggested by different rbcL studies. and Dipsacales (83%).g. Adoxaceae (including Sambucus and Viburnum) are sister to the rest of the order. The greater ability of ndhF sequences to resolve intrafamilial relationships (e. Unlike Euasterids I. and segregating a narrowly defined Asterales (Asteraceae only) would leave a paraphyletic grade of small groups often comprising individual families.... Aquifoliaceae. Morgan and Soltis. 1998). Thanks to P. This broad concept of Asterales is accepted here. Kim et al.. Within Dipsacales. Goodeniaceae (97%). 1995). 1987. The Aquifoliales comprise a group of three small families. based on ndhF.. each appears to be monophyletic (Fig. R. This also was found.. Jansen and Kim. means that data from studies of individual families based on ndhF may be integrated more effectively into a broad-scale study of Asteridae phylogeny and may ultimately help to resolve the remaining familylevel problem areas (e. Kim and Jansen.. 1991. 1992. The second Euasterid clade (Fig. 1996). Olmstead. which find Pittosporaceae to be sister to a combined Apiaceae/Araliaceae. With limited sampling in the two large families Apiaceae and Araliaceae.. and Aquifoliales being sister to the clade comprising the other three orders (Fig. though with weak support. These two taxa also share a 9-bp deletion in their ndhF sequence. 1). unpublished). 1992. Jansen et al. Ericales). but with less support. Our results suggest that Pittosporaceae and Apiaceae form a clade (81%). 1992. Cosner et al. and Campanulaceae (100%). including Asteraceae (80%)..O. 1).. Calycerales. 1998). With minor exceptions. in contrast to some rbcL studies (Chase et al.. Relationships inferred within this clade correspond closely to those based on rbcL studies. 1993. Takhtajan’s (1987) Asteridae sensu stricto corresponds very closely to the group recognized as the Asterales here and includes the Asterales. based on combined rbcL and ndhF sequences. Backlund and Bremer. ACKNOWLEDGMENTS This research has been supported by NSF Grant BSR-9107827 to R.. 1997. 1996.. which was included in this analysis as a binary character weighted equally with nucleotide substitutions. 1997). Backlund and Bremer (1997). which suggest that the traditional Caprifoliaceae are paraphyletic with respect to the other families in the order (Figs. 1993. Menyanthaceae (95%). with Apiales and Dipsacales forming a clade (65%). Molecular phylogenetic studies have examined Adoxaceae (Baldwin et al. 1995.G. Plunkett et al. 1997) and Valerianaceae (Backlund and Bremer. 1993... 1996) and chloroplast (Downie et al. Asterales (99%). Downie et al.. relative to rbcL. Gustafsson et al. 1). 1996) DNA. In this study. 1995.. based on rbcL. Morgan and Soltis (1993) note a shared leaf venation pattern in Helwingia and Phyllonoma. Michaels et al. the ndhF results confirm previous hypotheses of relationship based on rbcL sequences and provide greater resolution and increased support for those hypotheses. 1997). 1993. Lamiales. Asteraceae have been the focus of extensive molecular systematic study (Jansen and Palmer. Backlund and Bremer.. Aquifoliales (100%).. in the studies of Kim and Jansen (1995). 1 and 2). The large subfamily Apioideae has been the subject of additional study with both nuclear (Downie and Katz-Downie. Donoghue. Helwingiaceae/Phyllonomaceae—Chase et al. 1996). and Campanulales of Cronquist (1981). 1993. Olmstead et al. 1993) and matK and combined matK/rbcL (Plunkett et al. because segregating a narrowly defined Campanulales (Campanulaceae plus Stylidiaceae) would leave a poorly supported clade (37%). albeit with lower bootstrap support (Aquifoliaceae/Phyllonomaceae—Olmstead et al.. Chase et al.K.

a Matelea biflora (Rafin. coll. Kim 13777 (YNUH) G..-J. MI. no voucher R. Panero s. & Ernst a a Impatiens capensis Meerb. a Plantago lanceolata L. Kim 13748 (YNUH) K. ex Lindl.-J.-J. b Hydrostachys multifida A. I. Kim 12809 (TEX) J. no voucher K. M.-J. Kim 13743 (YNUH) K. Olmstead 93-45 (WTU) K.-J. f. MI. no voucher H. ex Regel b Gentianales/Gentianaceae Gentianales/Loganiaceae Gentianales/Rubiaceae Gentianales/Rubiaceae Gentianales/Rubiaceae Gentianales/Rubiaceae Gentianales/Rubiaceae Gentianales/Rubiaceae Lamiales/Acanthaceae Lamiales/Acanthaceae Lamiales/Veronicaceae Lamiales/Veronicaceae Lamiales/Veronicaceae Gentiana procera Holm Spigelia marilandica L. Kim 12776 (TEX) K. MI. and GenBank Accession Nos. Ceratophyllum demersum L.-J. 3413 (MO) Powell 1707 (TEX) Chase 114 K. & S. Kim 13785 (YNUH) K. Kim 13593 (YNUH) Rancho Santa Ana B. a Rhamnus davurica Pall.-J. Erithalis fruticosa Urb. Ann Arbor. Kim 12807 (TEX) Matthaei B.-J. a Guettarda uruguensis Cham.-J. no voucher K. a Ixora parviflora Vahl.G. Kim 13943 (YNUH) K. b Callitriche hermaphroditica L. a Thevetia peruviana K. a Staphylea emodi Wall. Hay & M. a Phytolacca americana L. Liriodendron tulipifera L. a Aquilegia bicolor Ehrh.n. a Phlox drummondii Hook. Kim 13840 (YNUH) Soltis and Soltis 1949 WS AA-478-78. b a a Araujia sericifera Brot. G.-J. Schum. Schatz et al. Kim 13754 (YNUH) Kron 2115 K.-J. Juss... a Eucnide rupestris (Baill. Magnolia sieboldii Koch a Paeonia tenuifolia L. a Anagallis arvensis L. Kulik 4061 (A) UC Campus.) Thomps. Kim 13790 (YNUH) T. Acer negundo L. & Schltdl. no voucher K. Gilmore 4 (A) UC Campus. a a K.) Lem.-J. Br. a Cercidiphyllum japonicum Siebold & Zucc.-J.-J. no voucher K.-J.-J. a Nerium oleander L. Kim 13693 (YNUH) Soltis and Soltis 2478 WS K. Westland. a Halesia carolina L.-J.) Ait. a Dicentra spectabilis (L. b a a Cephalanthus occidentalis L. Camellia japonica L. Kim 13700 (YNUH) K. a Carpenteria californica Torrey a Hydrangea macrophylla Ser. Hay. no voucher Bremer 2712 (S) UC GH 850161. a Gelsemium sempervirens (L. Kim 12898 (TEX) K. no voucher UC GH 850807. G. no voucher UC GH. Diospyros texana Scheele a Rhododendron mucronulatum Turcz.-J. (TEX) K. no voucher Beal B. ex Lindl. no voucher K.-J. Cryptostegia grandiflora R.. Kim 13736 (YNUH) K. Michaels. Harrison. Philbrick 3022 (CONN) AF130227 AF130232 AF130224 AF130226 AF130230 AF130231 AF130223 AF130234 AF130229 AF130233 AF130225 AF130222 AF130228 AF130221 AF130220 AF130217 AF130218 AF147712 AF130219 AF130210 AF130213 AF130209 AF130211 AF130212 AF130214 AF130215 AF130216 AF147714 AF130165 AF130167 AF130166 AF130168 AF130169 AF130170 AF147710 and AF147711 L36400 AF147713 AF130173 AF130174 AF130172 AF130176 AF130175 AF130171 AF130155 U12667 AF130150 AF130151 L36441 Alangium platanifolium Harms Cornus florida L. Order/family Outgroups Sapindaceae Ceratophyllaceae Cercidiphyllaceae a a Species DNA source/voucher GenBank Accession No.) Deflers a Rogiera suffrutescens (Brandegee) Borhidi a Justicia carnea Lindl. Kim 13766 (YNUH) UC Campus.-J.-J. a Pentas lanceolata (Forssk.) Woods.CHLOROPLAST PHYLOGENY OF THE ASTERIDAE 107 APPENDIX 1 List of Taxa Included in Study. no voucher K. a Heuchera micrantha Dougl. Kim 13602 (TEX) AA-23185A. Kim 12894 (TEX) AA-22308A I. a a Styrax americana Lam. Sources of Plant Material. a a a Fagaceae Magnoliaceae Magnoliaceae Paeoniaceae Papaveraceae Phytolaccaceae Ranunculaceae Rhamnaceae Saxifragaceae Staphyleaceae Cornales Cornaceae Cornaceae Hydrangeaceae Hydrangeaceae Hydrostachyaceae Loasaceae Ericales Balsaminaceae Ebenaceae Ericaceae Polemoniaceae Primulaceae Styracaceae Styracaceae Theaceae Euasterids I Garryales/Garryaceae Gentianales/Apocynaceae Gentianales/Apocynaceae Gentianales/Apocynaceae Gentianales/Apocynaceae Gentianales/Apocynaceae Gentianales/Gelsemiaceae Gentianales/Gentianaceae Quercus rubra L. b Thunbergia alata Bojer a Digitalis purpurea L. Kim 12498 (TEX) Meagher 990 (FTG) Gillis 9575 (FTG) Bremer 3104 (UPS) UC GH 850651. . b Exacum affine Balf. Garrya elliptica Dougl. Kim 12897 (TEX) K. East Lansing.

AF130198 AF130199 AF130200 AF130202 AF130203 AF130204 AF130205 AF130201 AF130206 AF130207 a a Osmorhiza claytoni (Michx. b Selago thunbergii Choisy b Verbascum thapsus L. a Panax ginseng C.-J. no voucher C.n. DC. Olmstead 92-144 (WTU) K. coll. s. Kim 13948 (YNUH) UC Pharmacy Garden. no voucher M.n. Hay & M.G. (AUK) K.. a Pittosporum tobirum Ait. no voucher R. Boulder.G. Petunia axillaris (Lam.n.) B.. S. coll. Forst. coll.) Wiehler Nematanthus ‘Butterscotch’ (CV. b Sesamum indicum L. no voucher UC GH. Wagstaff 89-026 (BHO) R. W. C. a Helwingia japonica (Thunb. b Hydrolea ovata Nutt. India) UC GH 850974.Sun. Kim 12808 (TEX) K. Bremer 3099 (UPS) UC GH 920093. APPENDIX 1—Continued Order/family Lamiales/Veronicaceae Lamiales/Bignoniaceae Lamiales/Bignoniaceae Lamiales/Bignoniaceae Lamiales/Buddlejaceae Lamiales/Buddlejaceae Lamiales/Gesneriaceae Lamiales/Gesneriaceae Lamiales/Gesneriaceae Lamiales/Gesneriaceae Lamiales/Lamiaceae Lamiales/Lamiaceae Lamiales/Lamiaceae Lamiales/Lamiaceae Lamiales/Lamiaceae Lamiales/Myoporaceae Lamiales/Oleaceae Lamiales/Oleaceae Lamiales/Oleaceae Lamiales/Oleaceae Lamiales/Paulowniaceae Lamiales/Pedaliaceae Lamiales/Scrophulariaceae Lamiales/Scrophulariaceae Lamiales/Scrophulariaceae Lamiales/Stilbaceae Lamiales/Stilbaceae Lamiales/Verbenaceae Lamiales/Verbenaceae Lamiales/Verbenaceae Solanales/Convolvulaceae Solanales/Hydroleaceae Solanales/Montiniaceae Solanales/Solanaceae Solanales/Solanaceae Solanales/Solanaceae Solanales/Solanaceae Solanales/Solanaceae Unassigned/Boraginaceae Unassigned/Boraginaceae Unassigned/Boraginaceae Euasterids II Apiales/Apiaceae Apiales/Apiaceae Apiales/Apiaceae Apiales/Araliaceae Apiales/Araliaceae Apiales/Araliaceae Apiales/Griseliniaceae Apiales/Pittosporaceae Aquifoliales/Aquifoliaceae Aquifoliales/Helwingiaceae b a Species Veronica catenata Pennell Campsis radicans Seem.-J.-J. no voucher Cult. Paulownia tomentosa Steud. no voucher K. Clarke Acanthopanax sessiliflorus Seem..n.. Herbarium.-J.0224..K. no voucher K. Olmstead 88-008 (WTU) Cult. Kim 13594 (YNUH) K. b Nicotiana tabacum L. . Gray a Vitex agnus-castus L. Olmstead 88-003 (WTU) Cult. J. of N.) Tronc.-J. Dave s. a b b Nolana spathulata Ruiz & Pav. B. Schizanthus pinnatus Ruiz & Pav.-J.-J. Kim 13594 (YNUH) K. no voucher K. a Lantana horrida H.) F. RGO S-72 (WTU) Cult. Meyer a Griselinia lucida G. MI. a Nyctanthes arbor-tristis L.) a Clerodendrum trichotomum Thunb. Aloysia gratissima (Gill & Hook.-J. Harrison 5579 (A) GenBank Accession No. no voucher Ka ¨ llersjo ¨ 0401191 (BOL) K. W. O.S. Kim 13555 (CNUH) Cameron s.. dePamphilis s.G. MI. 1984-4220.B.B. Kim 12803 (TEX) K.) Huds. in Hawaii Tropical Garden. L36419 AF130144 AF130145 LL36397 AF130143 L36405 AF130156 AF130159 AF130158 AF130157 AF130146 U78706 U78707 L36391 U78696 L36403 AF130162 AF130164 AF130161 AF130163 L36406 L36413 L36411 L36412 L36417 AF147715 AF147716 AF130154 AF130152 AF130153 AF130177 AF013999 AF130178 L14953 U08925 U08926 U08929 U08921 AF130179 L36393 AF019646 Drymonia strigosa (Oerst. Kew No.. Olmstead.P. b Scrophularia californica Cham. I.108 OLMSTEAD ET AL. Kim 12811 (TEX) AA-912. a a Angelica gigas Nakai Coriandrum sativum L. Kim 13557 (YNUH) K. a Tecoma stans Juss. Ann Arbor. Ann Arbor. MI.Kim 12899 (TEX) Matthaei B. a Phyla incisa Small a Ipomoea batata Lam. Mey) A. b Retzia capensis Thunb. Kim 10026 (TEX) K. Kim 12801 (TEX) K. Ann Arbor.-J. b Euthystachys abbreviata (E. Dietr.0367.-J.-J. Kim 12802 (TEX) K. Wen 850 (GH) K. no voucher UC GH 850103.-J. a Tetraclea coulteri A. dePamphilis s. Phacelia patuliflora (Engln. Dillon 3767 (F) Seed BIRM/s.n. Africa. Wagstaff 89-07 (BHO) S. in NYBG. b Ajuga reptans L. s. Kim 2804 (TEX) S. a Hedera helix L. Manoa. Clarke a a DNA source/voucher R. a b Hydrophyllum virginiana L. a b Olea europeaa L. longipes DC. no voucher B. (Sardar Patel Univ. no voucher J. RGO S-60 (WTU) Seed BIRM/s. ex Choisy a Montinia caryophyllacea Thunb.-J.n.-J. Olmstead. b Myoporum mauritianum A.. a a Jasminum mesnyi Hance Ligustrum vulgare L. a Ilex crenata Thunb.-J. R. Kim 12805 (TEX) R. J. b Catalpa speciosa Warder a Buddleja davidii Franchet b Nicodemia diversifolia (Vahl) Tenore a Codonanthe digna Wiehler a Cyrtandra hawaiensis C. no voucher R. Kim 13844 (YNUH) R. (CNUH) K.-J. no voucher B. Kim 12810(TEX) AA 486-88. Olmstead 89-009 (WTU) Williams 2833 (MO) Matthaei B. DC. CO no voucher McDonald & Rourke.) C. B. b Mentha rotundifolia (L. b b Solanum lycopersicum L. & Schldl. & Gray) Gray b Borago officinalis L. no voucher Y.

(1992). a Nymphoides indica (L. Syst.Nesom 7741 (TEX) R... Wagstaff. J. Univ. G. S. Pullman. WA.n. Am. (1997). Corokia cotoneaster Raoul DNA source/voucher D. (1998). Note. a Lobelia cardinalis L. a Dampiera diversifolia De Vriese a Scaevola frutescens Krause a Menyanthes trifoliata L. Kim 13768 (YNUH) K. Washington State University.. Baldwin. Gard. Harley. B. Bremer. K. B. based on rbcL sequences. rbcL sequences support exclusion of Retzia. Gard.” M. Biol. 82: 383–397. Ed. (1997)..) Honck. in NYBG s. no voucher K. 46: 313–331. Mo. Ann. Bremer. M. (1998). Cantino. R. Bot. B. (1998). Morgan 2124 (WS) Strybing Arb. B. Arnold Arobretum. Bremer. P. no voucher Cult.. Mo. L. Hufford. J. Kim 12841 (TEX) K. Richmond.) Blake Dasyphyllum argenteum Kunth a Gerbera jamesonii Bolus a Helianthus annuus L. 217: 119 –135. 82: 247–277... pp. Phylogeny of the Rubiaceae (Chiococceae) based on molecular and morphological data—Useful approaches for clas- sification and comparative ecology. K.. (1992)..-J. a Scabiosa mansenensis Nakai a Succisa pratensis L. 85: 531–553. Classification follows Angiosperm Phylogeny Group (1998). Struwe. Evol. D. R. Cosner 174 (OS) K. Ann. no voucher MBG 19426. D.. A. Bayer. 21: 121–134.-J. Mo. D. a Villarsia calthifolia F. no voucher Stuessy & Viteri 12464 (OS) R. L. B. . J. and Sweere.Muell. M.. Comparative restriction site mapping of chloroplast DNA implies new phylogenetic relationships within Rubiaceae. Harley. Phylogenetic utility of the nuclear rDNA ITS region in the subfamily Ixoroideae (Rubiaceae): Comparisons with cpDNA rbcL sequence data. Angiosperm Phylogeny Group. Backlund. (1995). B.-J.. G. J. no voucher Cult.. a Symphoricarpos orbiculatus Moench a Dipsacus sativus (L. 78: 198 –213.n. of Singapore) M. J. str. with particular reference to the Dipsacales. 74-211. Cosner 176 (OS) UC GH 980112. J. A comparison of phylogenetic nomenclature with the current system: A botanical case study. Kim 13839 (YNUH) GenBank Accession No. no voucher M. F. Combined and separate analyses of morphological and molecular data in the plant family Rubiaceae. J. Phylogeny of the Asteridae s. In “Advances in Labiate Science” (R..-J. s. Bremer.. Jansen 931 (MICH) K. Plant Syst. Gard. B.n. Ann.) Engl. and Bremer. (1996). and Bremer. Kim 12887 (TEX) K. R. Bot. a Campanula ramulosa Wall. Bot. UK. M.. (1996). Plant Syst. Barnadesia caryophylla (Vell. D. and Soltis.. Bremer. Jansen 915 (MICH) Price Ha89 (TAES) Cult. in NYBG. Evol. D. B. The ITS region of nuclear ribosomal DNA: A valuable source of evidence on angiosperm phylogeny. Andreasen. Andreasen. Royal Botanic Gardens..). University of Connecticut. Ann. Cantino. UC. a Lactuca sativa L. and Olsson. S.CHLOROPLAST PHYLOGENY OF THE ASTERIDAE 109 APPENDIX 1—Continued Order/family Aquifoliales/Phyllonomaceae Asterales/Argophyllaceae Asterales/Asteraceae Asterales/Asteraceae Asterales/Asteraceae Asterales/Asteraceae Asterales/Asteraceae Asterales/Asteraceae Asterales/Calyceraceae Asterales/Campanulaceae Asterales/Campanulaceae Asterales/Campanulaceae Asterales/Campanulaceae Asterales/Campanulaceae Asterales/Goodeniaceae Asterales/Goodeniaceae Asterales/Menyanthaceae Asterales/Menyanthaceae Asterales/Menyanthaceae Asterales/Pentaphragmataceae Asterales/Stylidiaceae Dipsacales/Adoxaceae Dipsacales/Adoxaceae Dipsacales/Caprifoliaceae Dipsacales/Caprifoliaceae Dipsacales/Dipsacaceae Dipsacales/Dipsacaceae Dipsacales/Dipsacaceae Dipsacales/Linnaeaceae Dipsacales/Valerianaceae a a Species Phyllonoma laticuspis (Turcz. Bot. C. Sambucus canadensis L. and Nicodemia (Buddlejaceae) from the Gentianales. Cantino. Jansen 906 (MICH) Kew 001-76-0038. Desfontainia. (1995). 207: 225–254. a Boopis anthemoides Juss.. B. Gard. G. D. Evol. “Phylogeny of the Asteridae s. G. Kim 13889 (YNUH) UC Pharmacy Garden. K.. Phylogenetic relationships in Sarraceniaceae based on rbcL and ITS sequences. and Donoghue. 511–522. and Wagstaff. Kim 13942 (YNUH) M. a Trachelium caeruleum L.. Jansen 984 (MICH) K. Gard. An ordinal classification of the families of flowering plants.-J. (1991).S. Plant Syst. no voucher Hunziker 25258 (CORD) R. Syst. Genera of Labiatae: Status and classification. New York Bot. (1998). (1994).. Ornduff 9726 (UC) Hugh Tan s. Porter. a Pentaphragma elliptica Poulsen a a Stylidium graminifolium Sw. no voucher R. and Jansen. a Cyananthus microphyllus Benth. Bot. 79: 380 – 387.. P. (Natl.. C. Kim 13767 (YNUH) G.-J. R. Wojciechowski. J. thesis.. and Wagstaff. b Sequences determined in Olmstead lab. a Sequences determined in Jansen lab. no voucher R. G. Cosner 173 (OS) Kew 420-84-04494. A. Olmstead. J. 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