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Phylogenetic classification of Ericaceae: molecular and morphological evidence.

Resumen

Se presenta una clasificacion nueva de las Ericaceae basada en analisis filogeneticos, empleando secuencias de ADN nuclear y de cloroplasto, morfologia, anatomia y embriologia. Se reconocen ocho subfamilias y 20 tribus. En esta clasificacion las Epacridaceae son incluidas como Styphelioideae y las Empetraceae como la tribu Empetreae dentro de Ericoideae. Los grupos herbaceos previamente reconocidos como Pyrolaceae y Monotropaceae por algunos autores tambien se incluyen dentro de las Ericaceae, en la subfamilia Monotropoideae. Se dan una clave, descripciones morfologicas e imagenes representativas de todos los grupos nombrados. Se hacen dos combinaciones nuevas en Kalmia (K. buxifolia y K. procumbens), y se describen tres nuevos taxones: Oligarrheneae, Richeeae y Cosmelieae (todos dentro de Styphelioideae).

II. Introduction

Prior to 1993, evolutionary relationships among the Ericaceae, Empetraceae, and Epacridaceae were addressed using a traditional "evolutionary taxonomic" approach. Classification was based on selected characters thought to be significant and on general notions of the primitive or advanced conditions in angiosperms. Cronquist (1981) considered Ericaceae, Empetraceae, and Epacridaceae to be closely related, and he placed them in the Ericales (also including Clethraceae and Cyrillaceae). Cronquist also recognized the non-green ericad taxa as Monotropaceae, and the green herbaceous taxa as Pyrolaceae. Subdivisions within Ericaceae and the recognition of the segregate families Monotropaceae and Pyrolaceae varied with different authors (e.g., Drude, 1889; Stevens, 1971; Thorne, 1992; Wallace, 1976; Watson et al., 1967) and are discussed in detail in Stevens (1971). However, cladistic relationships among members of the Ericales (sensu Cronquist, 1981) were not addressed until the studies by Anderberg (1992, 1993) and Judd and Kron (1993), who analyzed morphological data, and by Kron and Chase (1993), who analyzed rbcL sequence data. These explicitly phylogenetic studies showed not only that Ericaceae were paraphyletic in the traditional sense but also that Empetraceae and Epacridaceae were derived from within Ericaceae.

The inclusion of empetrids and epacrids within the ericads was the impetus for an in-depth molecular and morphological analysis of the entire ericad clade. The First International Ericaceae Conference was held in Gainesville, Florida, at the University of Florida to organize and coordinate an international collaborative effort that would include A. A. Anderberg (Sweden), D. M. Crayn (Australia), P. A. Gadek (Australia), W. S. Judd (United States), K. A. Kron (United States), C. J. Quinn (Australia), and P. F. Stevens (United States). The first meeting determined the list of proposed morphological characters, taxon sampling for both the molecular and morphological analyses, and methods of data acquisition for the morphological portion of the study. The Gainesville meeting also developed a timetable and general outline of goals and purpose of the project. Current work by the Australian group (Crayn, Gadek, Quinn) on epacrids, work on Enkianthus and empetrids by Anderberg, morphological work by Judd and Stevens, and molecular work in non-epacrids by Kron was coordinated to prevent duplication of effort. Sharing of DNA sequences and consistent exchange of information regarding preliminary results helped to maximize the number of taxa that could be sampled and allowed a focus on problematic areas early in the project. Naming of groups was deferred until essentially all of the data were in and at least preliminary analyses had been performed on individual and combined data sets. At this point the Second International Ericaceae Conference was held in Sydney, Australia. The primary purpose of this meeting was to develop a new classification of the ericads based on all available data (Fig. 1). In addition to the original participants, R. Fuller (United States), J. L. Luteyn (United States), and E. Brown (Royal Botanic Garden, Sydney, Australia, as outside observer) contributed their insight into the classification process.

Formally named groups were based on the following criteria: only monophyletic groups are recognized, no paraphyletic or metaphyletic groups; named clades are robustly supported or at least the clade should repeatedly occur in the strict consensus of each individual analysis. The new classification was developed using Wiley's (1981) sequencing convention for naming and followed the current system of botanical nomenclature (Greuter, 1994). The recommendations for naming groups discussed in Backlund and Bremer (1998) were also used when appropriate. All names subsequently used in the text, referring to Ericaceae or subgroups therein, follow the new classification (Fig. 1) unless otherwise noted.

III. Materials and Methods

A. SAMPLING

Taxa were chosen for inclusion in the study based on their representation of morphological diversity and geographical distribution (Appendix 1). Additional samples were used to represent clades previously identified in other cladistic studies (Anderberg, 1993; Judd & Kron, 1993; Kron & Chase, 1993) as well as major groups traditionally recognized within the Ericaceae, Epacridaceae, and Empetraceae. Subfamilies and tribes for which there was reason to suspect para- or polyphyly (e.g., traditionally recognized groups for which there were no known consistently diagnostic characters) were sampled more intensively than were those likely to be monophyletic as shown in previous studies (Anderberg, 1992; Crayn et al., 1998; Kron, 1996, 1997; Kron & Judd, 1997). The taxon sampling for rbcL, matK, and morphology were coordinated so as to maximize the number of taxa sampled in common among the three data sets. Additional taxa were sampled for each individual data set as was appropriate based on the amount of variation o bserved in preliminary analyses of subsets of the data. Terminal taxa were exemplar species in both molecular and morphological analyses (Kron & Judd, 1997; Kron et al., l999b). The same accession was used for all genes or regions sequenced for a particular species.

B. MORPHOLOGICAL DATA

Compared with many groups of plants the morphological, anatomical, and Phytochemical features of ericads have been well studied (Anderberg, 1992, 1993, 1994a, 1994b; Artopoeus, 1903; Baas, 1985; Bocher, 1981; Chou, 1952; Copeland, 1933, 1935, 1937, 1938, 1939, 1941, 1954; Cox, 1948; Dorr, 1980; Freudenstein, 1999; Furman & Trappe, 1971; Harborne & Williams, 1973; Hegnauer, 1966; Hermann & Palser, 2000; Hersey & Vander Kloet, 1976; Judd, 1979, 1981, 1982, 1984, 1986, 1990, 1995a, 1995b, 1995c; Judd & Hermann, 1990; Leins, 1964; Lems, 1962, 1964; Luteyn, 1991, 1995a, 1995b, 1995c, 1996; Matthews & Knox, 1926; Melvin, 1980; Middleton, 1991a, 1991b; Middleton & Wilcock, 1990a, 1990b; Niedenzu, 1890; Odell & Vander Kloet, 1991; Palser, 1951, 1952, 1954, 1958, 1961a, 1961b; Paterson, 1961; Rao & Chakraborti, 1985; Safijowska, 1960; Samuelsson, 1913; Sleumer, 1959, 1966, 1967; Smith-White, 1948, 1955, 1959; Stace et al., 1997; Stevens, 1969, 1970a, 1970b, 1971, 1995; Villamil & Palser, 1981; Wallace, 1975, 1976; Wat son, 1962, 1965; Watson et al., 1967; Wood, 1961). These previous studies provided a basic reference point for deciding which features would likely be informative across the family. However, character scoring was based on first-hand observations of living, herbarium, and liquid-preserved material whenever possible.

The preliminary list of morphological characters was assembled at the First International Ericaceae Conference. At this meeting a general procedure for obtaining data was developed. Characters were described in detail on standardized data sheets. The completed data sheets were sent to the senior author, who did the initial screening for obvious continuously variable characters and state delimitation. Further refinement of the data matrix resulted in 91 potentially informative characters (Table 1) for 80 taxa of ericads (Appendix 2).

Because of the size and complexity of the data matrix, heuristic approaches to tree construction were used. The morphological matrix was analyzed in the following manner using the parsimony criterion in PAUP* 4.0 (Swofford, 1999) [Enkianthus designated as outgroup based on Anderberg (1993), Judd & Kron (1993), Kron & Chase (1993)]: 10 searches of 10 random replicates each were performed using nearest neighbor interchange branch swapping (NNI) and saving the best tree from each replicate. The 100 trees obtained were used as the starting trees for a search that used tree bisection reconnection (TBR) branch swapping and saved all of the most parsimonious trees. The search was allowed to run until the computer's memory was exhausted. Trees one step longer were also investigated; however, because of the enormous number of trees only a rough estimate of branch support could be obtained.

Results of the parsimony analysis were inspected and characters that were highly homoplasious on the tree (>10 steps) were identified. A second analysis was performed under the same conditions as the first, but with the highly homoplasious characters removed.

C. MOLECULAR DATA

Total DNA was extracted using techniques based on the CTAB procedure of Doyle and Doyle (1987) and modified accordingly for ericads. Amplification and sequencing was performed by modified protocols outlined in Olmstead et al. (1992) and Kron et al. (1999b). Nuclear ribosomal (18s) and chloroplast genes (matK, rbcL) were sequenced for various subsets of Ericaceae (Appendix 1). These sequences were used to construct phylogenetic trees for purposes of determining the following: sister groups to the Ericaceae; relationships of Empetraceae and Epacridaceae to Ericaceae; tribal relationships within Ericaceae.

Nuclear ribosomal sequences were obtained for 16 taxa of Ericaceae (18s) and 8 outgroup taxa. Within the chloroplast genome the rbcL and matK genes were sequenced for 90 and 130 taxa (species) within Ericaceae, respectively (89 species in common). Sequences were edited using Sequencher 3.0 (Gene Codes Corporation, Ann Arbor, MI, U.S.A.), aligned manually and analyzed using PAUP* 4.0 (Swofford, 1999). Measures of branch support were assessed using parsimony jackknife (Farris et al., 1996; Kallersjo et al., 1998). Indels (insertion/deletions) were scored as missing data. Each data set was analyzed separately. Transition/transversion ratios and changes by codon position (where applicable) were estimated using MacClade 3.0 (Maddison & Maddison, 1992). Comparison of individual analyses showed little or no conflict (as determined by inspection of strict consensus trees and comparison of parsimony jackknife support), and the data were combined for additional analyses. A combined analysis of 18s, matK, and rbcL was p erformed for 16 taxa representing major groups within the ericadclade as well as potential sister groups. Combined data matrices were analyzed for rbcL + matK (Ericaceae s.l.).

Analyses were performed with all characters and character state changes equally weighted. Because of the large size of the data sets, heuristic approaches were used to search for the shortest trees. For the individual matK and rbcL analyses the results of parsimony jackknife analysis (Farris et al., 1996; Kallersjo et al., 1998) were used to construct a constraint tree (clades with >60% parjac support). The data were analyzed using the parsimony jackknife--based constraint and random addition sequence (10 replicates). NNI swapping was used, and all of the most parsimonious trees were saved. These trees were used as the starting pool of trees for the second phase of the search. In this phase, the parsimony jackknife--based constraint was removed, and TBR branch swapping was performed. Combined analyses of molecular data were performed using standard heuristic search methods (100 random additions, TBR, MULPARS).

D. COMBINED ANALYSIS OF MORPHOLOGICAL AND MOLECULAR DATA

The total evidence approach (Ernisse & Kluge, 1993) to analyzing the data was used in this study, and a combined analysis (100 random additions, TBR, MULPARS) of morphology, matK, and rbcL was performed for 59 taxa of Ericaceae using PAUP* 4.0 (Swofford, 1999). Parsimony jackknife and bootstrap values were also obtained using the XAC program (Farris et al., 1996; pers. comm.) and the bootstrap option of PAUP*4.0. Morphological character-state transformations were mapped onto a representative tree obtained in the combined analysis using MacClade 3.0 (Maddison & Maddison, 1992).

IV. Results

A. DELIMITATION OF ERICACEAE

The combined analysis of nrl8s, rbcL, and matK( for 16 taxa representing major clades of ericads and several outgroups [Diospyros was used to root the tree (Morton et al., 1996)] resulted in three most parsimonious trees [uninformative characters removed, L (length) = 1558, CI (consistency index) = 0.53, RI (retention index) = 0.59]. The strict consensus (Fig. 2) indicates that Cyrilla (Cyrillaceae) is sister to Ericaceae, but support for this is less than 50% (bootstrap). The relationship of Cyrilla and Clethra to Ericaceae in this analysis differs from that obtained in a larger rbcL analysis (Morton et al., 1996). lathe large rbcL analysis, Cyrilla is more closely related to Actinidia than to Ericaceae, and Clethra is more closely related to Styrax than to Ericaceae. However, these results have low parsimony jackknife support (Morton et al., 1996). Because of the uncertainty of placement of Cyrilla and Clethra with respect to Ericaceae, Actinidia is used as an outgroup in the matK and rbcL analyses included in this study. Enkianthus is sister to the remaining Ericaceae. Arbutoideae (Arbutus) are the next to branch after Enkianthus, followed by Pyrola (representing Monotropoideae). The position of Arbutoideae is strongly supported (100%); however, only one representative of each clade was used in this analysis.

The sister relationship of Styphelioideae to Vaccinioideae found in this analysis is strongly supported. In the Ericoideae clade, Cassiope is sister to remaining Ericoideae, but this is not strongly supported (<50% bootstrap). The relationships within Ericoideae (i.e., Bejaria, Calluna, Ceratiola, and Rhododendron) are unresolved in this analysis.

B. RELATIONSHIPS WITHIN ERICACEAE

1. Morphological Relationships

Parsimony analysis (incomplete due to memory limit) of the 91 morphological characters resulted in 82,000 trees of 472 steps each (autapomorphies excluded); CI = 0.25, RI 0.66. Trees one step longer were obtained to estimate the relative branch support for particular relationships indicated in the most parsimonious trees. A strict consensus of the most parsimonious trees plus the mp + 1 trees showed significant loss of resolution among groups, but most smaller groups (corresponding roughly to tribes) remained intact (tree not shown). However, this is merely a rough estimate, as it is likely that not all of the most parsimonious trees were found. Of the 91 characters used in the analysis, 9 characters were identified as highly homoplasious (i.e., >10 changes) and likely contributing more noise than signal. These characters were excluded from a second analysis: pith condition (char. #8), leaf persistence (char. #17), inflorescence (char. #31), pedicel bracteoles (char. #35), abaxial calyx stomata (char. #43), i ndumentum of interior surface of corolla (char. #49), filament hairs or texture (char. #55), anther texture (char. #58), and basic chromosome number (char. #89). Of these nine characters only basic chromosome number (char. #89) was scored with more than 3 states.

The second analysis (Figs. 3A, 3B) resulted in 86,600 most parsimonious trees (L = 346,01 = 0.28, RI 0.70), but due to computer memory limitations not all of the most parsimonious trees were found. Bootstrap values (fast bootstrap) were generally low, with the deeper nodes of the tree poorly supported. This analysis found several clades that correspond to named subfamilies and tribes in Ericaceae. The base of the tree is unresolved with the Pyroleae, the non-green taxa (i.e., Monotropeae, Sarcodes, and Piero spora), and the remaining ericads forming a trichotomy. The non-green taxa are supported as a clade (56% bootstrap) by characters #3 (lacking chlorophyll), #6 (branched trace in petiole or leaf base), #15 (more or less palmate leaf venation), and #76 (partial to distinctly intruded parietal placentae). The Pyroleae are supported by the lack of multicellular hairs on the leaves (char. #22), erect to more or less horizontal corollas (char. #45) that are choripetalous (char. #44), and the absence of anther a ppendages (char. #59). In some of the most parsimonious trees the Pyroleae and non-green taxa form a clade supported by the herbaceous habit (char. #2), fibers associated with the midrib lacking (char. #19), anther appendages present on the anther filament junction (char. #60), and undeveloped embryos with only 30-40 cells (char. #85).

The Arbutoideae clade is strongly supported (86% bootstrap), with synapomorphies that include the presence of adaxial calyx stomata (char. #42), conspicuously asymmetrically dilated anther filaments (char. #54), and an indehiscent fruit (char. #78). Arbutoideae also possess a bony endocarp (char. #81).

The remaining Ericaceae form a monophyletic group, with the synapomorphy of anthers inverting early in the development of the flower (char. #56). The results also indicate pedicel articulation (char. #37) as a synapomorphy for this clade, although this character state has been subsequently lost many times. Within this large clade (early anther inversion clade), several groups correspond to clades also found in the molecular and combined analyses. Ericoideae are supported by characters #45 (corolla erect more or less horizontal), #69 (presence of viscin threads associated with the pollen), and #79 (fruit a septicidal capsule). Of these characters, the septicidal capsule is the most consistent, with a single reversal to loculicidal in the Ericeae and subsequently back to septicidal in Calluna. Vaccinieae are supported as monophyletic in the morphological analysis by the inferior ovary (char. #72). Lyonieae are monophyletic in this analysis (Kron & Judd, 1997; Kron et al., 1999b), as indicated by the presence of bands of fibers in the secondary phloem (char. #7). Styphelioideae are supported as monophyletic by the presence of bisporangiate anthers (char. #57) and a persistent corolla (char. #48). The Rhodoreae dade is supported by the presence of zygomorphic flowers (char. #33) and the presence of the leaf flavonoid gossypetin (char. #87), although both of these characters are homoplasious.

2. Molecular Relationships

a. matK Analysis

The aligned matrix required 20 insertion/deletions (indels) that are restricted to the 5' half of the gene. Most of the size variation was found within Styphelioideae. These include some very large deletions in Needhamiella (45 bp, 237 bp) and Oligarrhena (36 bp, 108 bp). Of the 20 indels, 6 were potentially informative. A 12-base insertion is common to all Ericaceae except Enkianthus and several members of Vaccinioideae. The most parsimonious explanation for this distribution is an insertion event in the ancestor of the pollen tetrad dade with multiple losses in the Vaccinieae, Gaultherieae, and Lyonieae. A 6-base deletion is a potential synapomorphy of Cosmelieae and Styphelieae, but this insertion also appears inPrionotes. Also within Styphelioideae, Richea and Dracophyllum share a unique 6-base insertion. Other clades with shared insertions include Kalmia, with a 3-base insertion. This insertion is apparently also present in the Lyonia clade, as well as in Chamaedaphne, Pieris nana, and Agarista populifol ia. A different 3-base insertion is shared by Orthilia secunda and Pyrola picta.

Parsimony analysis of the matK data matrix (127 taxa) resulted in more than 50,200 trees (L = 4138, CI = 0.36, RI = 0.68, uninformative characters removed, transition/transversion ratio = 0.85). Because of the size of the matrix the analysis did not swap to completion. The strict consensus (Figs. 4A, 4B) demonstrates that matK data allow for sufficient resolution of many relationships within the ericads, primarily among the more recent clades within the group. Deeper branches are poorly supported (<60% parjac) in this analysis. Groups that are well supported include Pyroleae, Arbutoidene, Empetreae, Rhodoreae, Prionoteae, Styphelioideae, and Vaccinieae (Table I). Within Ericoideae, there is strong support for the sister relationship of Therorhodion to Rhododendron + Menziesia + Diplarche and for the dade including Phyllodoce, Kalmiopsis, Rhodothamnus, and Epigaea. Cassiope is sister to Ericoideae (63% parjac), and Harrimanella is sister to Styphelioideae + Vaccinioideae (<60% parjac). In this analysis the Arb utoideae, Pyroleae, and Monotropeae branch sequentially near the base of the tree. However, this branching order lacks parsimony jackknife support above 60%.

b. rbcL Analysis

Analysis of rbcL data from 89 taxa resulted in 68,700 trees (L = 1515, CI = 0.36, RI 0.65, uninformative characters removed; transition/transversion ratio 1.0). It is likely that, as in the matK analysis, this does not represent all of the most parsimonious trees because memory limits did not allow this analysis to swap to completion. The strict consensus (Figs. SA, 5B) shows that many of the same groups are resolved based on the rbcL data as were found using matK. In this analysis several of the deeper branches were moderately to strongly supported by the parsimony jackknife analysis. Groups that receive strong support include pollen tetrad dade (i.e., sister group to Enkianthus), Pyroleae, Arbutoideae, early anther inversion clade, Ericoideac, Empetreae, and Styphelioideae. The general pattern of relationships is the same in both matK and rbcL analyses, although there are some notable differences. In the rbcL analysis the Pyroleae branch first after Enkianthus, followed by the Arbutoideae dade, This is the reverse of the relationship indicated in the matK analysis, in which the Arbutoideae branch before the Pyroleae. However, the rbcL analysis does not include the representatives of Monotropeae, and there is very weak support for the relationships indicated by the matK data. One strongly supported relationship found in the rbcL analysis is Diplarche sister to Empetreae. This is different from the matK analysis, in which Diplarche is placed within the Rhodoreae clade. This is the only case of strongly supported disagreement between the rbcL and matK analyses.

c. Combined Analysis of matK and rbcL

Combined analysis of rbcL and matK for 70 taxa resulted in 448 trees (L = 4435, CI = 0.52, RI = 0.65). Support for the relationships in this tree is generally strong (Fig. 6). All of the deepest branches within the tree are supported at the 99% or 100% level (parjac), with the exception of the branching order of the Pyroleae, Arbuteae, and the remaining Ericaceae (i.e., early anther inversion clade), which is unresolved. Arbutoideae, Ericoideae, Styphelioideae, and Vaccinioideae are each supported at 100%. Other named groups with strong support include Pyroleae, Rhodoreae, Ericeae, Empetreae, Lyonieae, and Vaccinieae. The Phyllodoceae form a dade but are not well supported in this analysis. Other groups that receive strong support include a dade containing Tepuia, Diplycosia, Gaultheria, and Pernettya (100% parjac) and Therorhodion as sister to Rhododendron + Menziesia + Diplarche. As in the individual analyses of rbcL and matK, Cassiope is sister to Ericoideae, and Harrimanella is sister to Styphelioideae + Vaccinioideae; but now, in the combined analysis, support for these relationships is strong (99% parjac each for the position of Cassiope and Harrimanella).

3. Combined Analysis of Morphological and Molecular Data

Parsimony analysis of rbcL, matK, and 91 morphological characters for 59 taxa (Fig. 7) resulted in 96 trees (L = 3562, CI = 0.41, RI = 0.62). Glade support throughout most of the tree is high (bootstrap), with 32 of the 45 clades identified with a bootstrap value above 80%. In addition, the relationship of Pyroleae (representing Monotropoideae) and Arbutoideae to the remaining ericads is resolved in this analysis (73% bootstrap). Some morphological characters that appeared to be homoplasious in the first morphological analysis are consistent in the combined analysis. Early inversion of anthers in the bud is the morphological synapomorphy (unique) for the large dade that includes Ericoideae, Styphelioideae, and Vaccinioideae (in contrast to the late anther inversion present in Arbutoideae and Monotropoideae). This "early anther inversion clade" has 100% bootstrap support in the combined analysis. In the first morphological analysis this dade was not recovered, whereas in the second morphological analysis it wa s consistently present but without bootstrap support (Fig. 3). In this analysis Vaccinieae are strongly supported as closely related to Gaultherieae plus Andromeda and Zenobia (i.e., Andromedeae). Individual analysis of molecular and morphological data also indicate a close relationship of Vaccinieae to Gaultherieae and associated taxa, but not in a sister relationship, as indicated in the combined morphological + rbcL + matK analysis.

When the molecular and morphological data were analyzed after the removal of the nine characters found to be highly homoplasious in the morphological analysis, 20 trees were obtained (L = 3461, CI = 0.42, RI = 0.63). The strict consensus of these trees was nearly identical with the first combined morphological and molecular analysis, with the exception that relationships among the tribes within Ericoideae are resolved in the second analysis. However, none of the resolved branches is supported by more than 50% bootstrap.

V. Discussion

A. DELIMITATION OF ERICACEAE

Previous studies of the limits of Ericaceae (Anderberg, 1992, 1993; Judd & Kron, 1993; Kron, 1996; Kron & Chase, 1993) indicated that Epacridaceae and Empetraceae are nested within the traditionally defined Ericaceae. A large-scale study that addressed the monophyly of Ebenales using rbcL sequences (Morton et al., 1996) supports this hypothesis. The sister relationship of Enkianthus to the remaining Ericaceae s.l. was also indicated by rbcL data (Morton et al., 1996). The combined analysis of 18s, rbcL, and matK data in this study indicates strong support for the inclusion of Epacridaceae and Empetraceae within an expanded Ericaceae.

Cyrillaceae and Clethraceae were included in Cronquist's (1981) order Ericales; in addition, Takhtajan (1997) placed Actinidiaceae in Ericales, and subsequent morphological and molecular phylogenetic studies have included representatives of Actinidiaceae, Glethraceae, and Cyrillaceae. Morton et al. (1996) found Actinidiaceae + Cyrillaceae as sister to Ericaceae s.l. based on rbcL data. However, support for this relationship was weak. The combined analysis of 18s, rbcL, and matK presented here indicates Cyrilla (Cyrillaceae) as sister to Ericaceae, but Actinidia (Actinidiaceae) is not sister to Cyrilla in this analysis. The relationships of potential sister groups, Actinidia, Clethra, and Cyrilla, found in this analysis of 18s, matK, and rbcL have been supported in a recent study by Anderberg et al. (2002) of the Ericales dade (APG, 1998), using 5 genes.

Trees in the combined analysis of 18s, matK, and rbcL obtained the same general relationships of subfamilies found in the larger individual matK and rbcL analyses. The sister relationship of the Vaccinioideae to the Styphelioideae has also been shown in a more detailed analysis of the relationships of Styphelioideae and traditionally defined "Vaccinioideae" (e.g., Stevens, 1971) using matK (Kron et al., 1999b).

B. RELATIONSHIPS WITHIN ERICACEAE

1. General Relationships

Results of the individual analyses of molecular data generally correspond well. In both analyses Enkianthus branches first, followed by the Monotropoideae clade and the Arbutoideae clade. The order of branching of the latter two clades is different in the rbcL and matK analyses, but support in both analyses is weak. In the matK analysis Allotropa and Monotropa hypopithys (not present in the rbcL analysis) form a clade and branch sequentially after Pyroleae. However, support for this relationship is weak. Other molecular studies of the non-green Ericaceae (Monotropeae and Pterosporeae) using 18s, nrITS (Kron, in prep.), and 26s (Cullings, 1994) indicate a close relationship of the herbaceous Ericaceae (Pyroleae, Monotropeae, Pterosporeae) to Arbutoideae.

In both rbcL and matK analyses two large sister clades (1 and 2; Figs. 3 and 4) are identified. The only difference in clade membership between the two analyses is the expanded sampling present in the matK analysis. In both analyses Cassiope and Harrimanella are not closely related. Cassiope is sister to the remaining Clade 1 (i.e., Ericoideae), and Harrimanella is sister to the remaining Clade 2 (i.e., Styphelioideae + Vaccinioideae). The morphological similarities found in Cassiope and Harrimanella are thus likely symplesiomorphies or homoplasies and not indicative of recent common ancestry.

In general, relationships among smaller clades within Clade 1 differ with respect to levels of support or resolution in the matK and rbcL analyses. In both analyses Ericeae and Empetreae are clades. With the exception of Elliottia, Phyllodoceae form a clade in the rbcL analysis (weakly supported). Within the Phyllodoceae the Kalmia group and the Phyllodoce clade (Phyllodoce, Rhodothamnus, Kalmiopsis, Epigaea) are well supported. In the matK analysis the Phyllodoceae do not form a clade, but the Kalmia group and the Phyllodoce group are strongly supported. In the rbcL analysis there is no support for the Rhodoreae clade, in contrast to the matK analysis. Diplarche is placed within the Rhododendron clade in the matK analysis, but in the rbcL analysis it is placed as sister to Empetreae.

Within Clade 2 Styphelioideae are sister to Vaccinioideae. Vaccinieae are supported as monophyletic in both analyses. Andromeda and Zenobia are sister taxa in the rbcL and the matK analyses, supporting the monophyly of Andromedeae (as defined in this study). This relationship was not anticipated based on morphological characters but is strongly supported in the analyses presented here, as well as in a more detailed analysis discussed in Kron et al. (1999b). The paraphyly of Gaultheria is indicated in the matK analysis, but in the rbcL tree Gaultheria splits into two clades, and these are unresolved with respect to each other and to Pernettya and several other vaccinioid taxa. The Styphelioideae clade is present in both analyses, with Prionotes + Lebetanthus sister to the remaining Styphelioideae.

The differences between the rbcL and matK analyses may reflect differences in sampling, but variability in rbcL was approximately 33% less than in matK, making a direct test of this hypothesis difficult. Although the number of taxa sampled was different (127 in the matK analysis, 89 in the rbcL), relationships of major groups were generally the same, and levels of homoplasy in both analyses were very similar. The matK analysis showed strong support for many of the smaller clades but low (parjac) support for the larger clades in Ericaceae. In contrast, rbcL data showed strong parjac support for several of the larger clades, but support for many smaller clades was weak. In the matK analysis groups within Ericoideae were poorly resolved, but relationships within Lyonieae were well resolved. Conversely, resolution within the Lyonieae was poor in the rbcL analysis, and the relationships between the Kalmia clade and the Phyllodoce clade were better resolved than in the matK analysis. Neither the matK nor the rbcL a nalyses swapped to completion. However, parsimony jackknife analyses indicated significant phylogenetic signal in both data sets. In addition, the general topological congruence found in the two separate analyses lends support to the indicated relationships.

The combined analysis of rbcL and matK resulted in fewer trees, and in these trees most clades showed strong parjac support. In this analysis Lebetanthus and Prionotes are sister taxa (Clade 2), and Therorhodion is sister to Rhododendron (including Diplarche). Deeper branches within the tree are strongly supported, with the exception of the unresolved relationship of Arbutoideae and Monotropoideae with respect to the remaining Ericaceae (except Enkianthus). The relationships of the tribes of Ericoideae remain unresolved, as do the relationships of Lyonieae to Gaultherieae + Andromedeae and Oxydendrum.

Morphological features analyzed in combination with rbcL and matK data resulted in highly resolved trees with strong bootstrap support for most of the clades. The addition of morphological data helped to resolve the relationship of the Monotropoideae and Arbutoideae to remaining Ericaceae. Monotropoideae branch after Enkianthus, making the herbaceous green (and, by extension, non-green) taxa an early branch in the evolution of Ericaceae. Arbutoideae branch next and are sister to the clade containing Clade 1 and Clade 2. Analysis of the morphological data alone for the 59 taxa included in the combined molecular and morphological analysis resulted in four most parsimonious trees (L = 360, CI = 0.29, RI = 0.63, not shown). Relationships in these trees differed substantially from the combined analysis trees, but bootstrap was low (below 50%) for 32 of the 45 clades obtained. Constraining the morphological tree (MacClade 3.0 [Maddison & Maddison, 1992]) to the topology found in the combined molecular and morpholog ical tree (Fig. 7) resulted in an increase of 42 steps. Similarly, when constraining a most parsimonious tree obtained from the larger morphological analysis (91 characters, 80 taxa; Fig. 3) to fit the topology found in the combined molecular and morphological analysis, an increase of 43 steps was required. Thus, at least in this instance, the smaller taxon sampling used in the combined analysis appears sufficient for representing general aspects of morphological evolution among the major groups of Ericaceae.

The combined analysis has fairly strong support (73% bootstrap; Fig. 7) for a clade containing Arbutoideae, Clade 1, and Clade 2. Morphological support for this grouping includes leaves scattered along the stem (char. #12), pedicel with two bracteoles (char. #35), loss--or reduction of--an endothecium (char. #66), and pollen shed in tetrahedral tetrads (char. #68). Bracteole number, however, is rather homoplasious (Fig. 8). A fibrous endothecium has also been lost in many non-photosynthetic Monotropoideae and is present in Bejaria. It is noteworthy that tetrads also occur in Moneses and Pyrola (within Monotropoideae); the presence of tetrads in these taxa is most parsimoniously interpreted as a parallelism (Fig. 9). However, if tetrads are considered to have evolved only once, then the feature becomes synapomorphic for all Ericaceae except Enkianthus.

Significantly, the combined analysis also strongly supports the existence of a large clade (Clade 1 + Clade 2 = early anther inversion clade; Fig. 7) that contains all in-group taxa except Monotropoideae and Arbutoideae. This clade is supported by the morphological feature of anthers inverting early in development (char. #56; Fig. 10). Once this feature evolved in Ericaceae, it has apparently never been lost. Early anther inversion is a remarkable morphological feature (Matthews & Knox, 1926), though not unique to Ericaceae, and correlates with a grouping strongly supported in the molecular analyses (Figs. 2, 4, 5, and 6).

Within the early anther inversion clade, Clade 2 comprises Vaccinioideae + Styphelioideae + Harrimanella, and Clade 1 is composed of Ericoideae + Cassiope. Morphological support for either Clade 1 or Clade 2 apparently is lacking, although Clade 1 (Ericoideae + Cassiope) could be supported by the presence of leaves revolute in the bud (char. # 10; Fig. 11), depending on how the character is optimized. This lack of morphological support argues against the formal recognition (i.e., naming) of either Clade 1 or Clade 2.

2. Enkianthoideae

The Enkianthoideae comprise a single genus, Enkianthus, with about 15 species occurring in Japan, China, Taiwan, and Indochina. This genus is the sister group of all other taxa of the Ericaceae (Anderberg, 1993; Judd & Kron, 1993; Kron & Chase, 1993). Cox had earlier noted that "the primitive species of Enkianthus are without doubt the most primitive in the Ericaceae" (1948, 495). Enkianthus was included in the Andromedeae (Drude, 1889; Hooker, 1876; Watson et al., 1967) or Cassiopeae (Cox, 1948) but was placed in its own tribe, Enkiantheae, of the Vaccinioideae, by Stevens (1971).

The species of Enkianthus differ from other Ericaceae in having anthers with fibrous endothecium, monadinous pollen without viscin threads, and seeds with vascular bundles in the raphe, but these features may represent symplesiomorphies, and as such they are not diagnostic of the genus. The monophyly of Enkianthus and the phylogenetic interrelationship among its species were investigated by Anderberg (1994a). Evident synapomorphies are difficult to find, but some character states have been mentioned in the literature as being unique in the genus; that is, trichomes of moderately long cells with a distal end that become necrotic long before the proximal end (Stevens, 1971), a nucellar pedestal on which the embryo sac is attached, earlike projections from embryo sac, and ventral carpellary bundles in the septal plane (Palser, 1951, 1952, 1961b). Given the position of the Enkianthoideae as the sister group of all other Ericaceae, a character such as perulate buds emerges as a potential synapomorphy of Enkianthus . This feature is characteristic of the Rhodoreae but is generally not found in Arbutoideae or the herbaceous Pyroleae, Monotropeae or Pterosporeae of the Monotropoideae. It is most parsimonious to assume that perulate buds have evolved in parallel in Enkianthus and in the Rhodoreae, especially since axillary branches arise from the axils of the perulae, a feature found elsewhere only in a very few species of Rhododendron.

The species of Enkianthus differ in leaf texture, inflorescence structure, corolla shape, and seed morphology, but they are rather similar in most other respects. The leaves are flat and fairly large. They are spiral but often borne in clusters toward the end of the branches. Flowers are arranged in raceme-like or umbel-like inflorescences and have corollas that are campanulate, tubular, or urceolate. The stamens are twice the number of the corolla-lobes and have anthers that are smooth to distinctly papillose, opening with longitudinal slits by means of a fibrous endothecium. The anthers have awned appendages, which gradually narrow from the thecae, and the filaments are variously papillose to hairy. The ovary is superior, and the fruit develops into a loculicidal capsule opening with five valves. The seeds in most species are provided with winglike ridges. Based on the cladistic analysis by Anderberg (1994a) Enkianthus seems to have originally been shrubs with deciduous leaves, racemose inflorescences, camp anulate corollas, and winged seeds, whereas the more derived species possess coriaceous leaves, umbellate inflorescences, flowers with tubular or urceolate corollas, and unwinged seeds.

In the early evolution of the Ericaceae two evolutionary lineages emerged. One of these evolved into the Enkianthoideae, of which Enkianthus is the only known representative today. The other evolved into the ancestral group that has diversified into the rest of the family.

3. Monotropoideae

Monotropoideae are united by their more or less herbaceous habit (char. #2) and reduced embryo (char. #85). Many members of this clade lack multicellular hairs on the leaves (char. #22), and this rather homoplasious condition is also resolved as a synapomorphy for members of Monotropoideae in the combined analysis (Fig. 7). However, in the morphological analysis multicellular hairs are hypothesized to have been lost twice within Monotropoideae, with Pterospora and Sarcodes retaining the plesiomorphic condition (i.e., possessing such hairs), and this character optimization is more likely, given the greater sampling of members of Monotropoideae in the morphological analysis.

Monotropoideae comprise three tribes: the autotrophic Pyroleae and the mycotrophic Pterosporeae and Monotropeae. Pyroleae are characterized by the lack of multicellular hairs on their leaves (char. #22), separate petals (char. #44), and lack of stamen appendages (char. #59). Phylogenetic relationships within Pyroleae recently have been clarified by Freudenstein (1999). Both Monotropeae and Pterosporeae have lost chlorophyll, becoming strongly mycotrophic (char. #3) with reduced leaves; this condition likely has evolved in parallel in these two clades. Although the Pterosporeae (Pterospora and Sarcodes) are not monophyletic in the morphological analysis (Fig. 3), they are monophyletic in analyses that include nrITS and 18s data (Kron, unpubl.). The Pterosporeae are fairly generalized; they have retained many plesiomorphic characters but show the apomorphies of non-verticillate leaves (char. #12) and more or less parietal placentation (char. #76). Monotropeae are a distinctive clade and can be diagnosed by thei r black coloration upon drying (char. #5), non-verticillate leaves (char. #12), lack of multicellular hairs on the leaves (char. #22), lack of stamen appendages (char. #59), lack of an endothecium (char. #66), more or less parietal placentation (char. #76), and an extremely reduced embryo (char. #85). Only the black coloration and extremely reduced embryo are uniquely derived.

4. Arbutoideae

The monophyly of the Arbutoideae is also well supported in the combined analysis. The following morphological characters are likely synapomorphic for members of this clade: inside of corolla with unicellular hairs (char. #49), filaments conspicuously asymmetrically dilated (char. #54), indehiscent fruit (char. #78), fleshy fruits, brightly colored (char. #82), with a bony or fibrous endocarp (char. #81), and plants with high levels of ellagic acid (char. #86). Only the last is not homoplasious (but it has been insufficiently investigated). The others also occur in a few or several other groups within the Ericaceae. The presence of a bony or fibrous endocarp, however, occurs elsewhere only in Empetreae and in some Styphelioideae, although sclerids occur in the endocarp of the bony capsules of Craibiodendron. Although not coded in the analysis, the Arbutoideae may also be united by the reduction in the number of ovules per locule. In this clade the ovule number varies from few to one, whereas most Ericaceae hav e more or less numerous ovules per locule.

5. Cassiopoideae

Within Clade 1, the sister-group relationship of Cassiope and Ericoideae is well supported. The monophyly of Cassiope (11 spp.) is supported by several homoplasious characters: Calluna-type pith (char. #8), lack of bud scales (char. #9), decussate leaves (char. #11), fibers absent or poorly developed in leaf midrib (char. #19), and reduced axillary inflorescences (char. #29). It is noteworthy that axillary inflorescences are very uncommon in Clade 1 (Ericoideae + Cassiope), occurring more commonly in Vaccinioideae (Fig. 12). Ericoid leaves (char. # 13) may be an additional apomorphy of Cassiope, but such leaves commonly occur within Clade 1 (Fig. 13). It is also possible (though less parsimonious) to assume that ericoid leaves evolved in the common ancestor of Clade 1 or even in the common ancestor of the core Ericaceae, but the ericoid leaf of Cassiope differs in morphology from that of Calluna and other ericoid members of Ericoideae (Stevens, 1970b). A distinctive synapomorphy of the species of Cassiope, wh ich was not included in our analyses, is the bisporic embryo sac. Cassiope is the only member of the Ericaceae known to have other than a Polygonum type embryo sac (Palser, 1952). The unusual fasciculate hairs may also be synapomorphic.

6. Ericoideae

Morphological support for Ericoideae (as here delimited; i.e., including the traditional Rhododendroideae; Fig. 7) includes the erect to more or less horizontal position of the flowers (char. #45), loss of stamen appendages (char. #59), and septicidal capsules (char. #79). All these characters, however, show homoplasy (Figs. 14, 15, and 16). The last is probably most useful, but it is important to note that loculicidal capsules have evolved in Erica, a very species-rich group. Viscin threads (char. #69; Fig. 17) are restricted to this group (among extant Ericaceae) and may represent another synapomorphy for these plants.

Within Ericoideae, five major clades--Bejarieae, Phyllodoceae, Ericeae, Empetreae, and Rhodoreae--can be recognized, although their interrelationships are unclear (Fig. 7). Ericeae, Empetreae, and Rhodoreae have strong bootstrap support. Bejarieae and Phyllodoceae lack support above 50%, but these clades are found in all of the most parsimonious trees obtained in both the combined molecular and molecular and the morphological data analyses. All of the tribes in Ericoideae can be recognized by means of morphological synapomorphies.

a. Bejarieae

Bejarieae can be diagnosed by their separate petals (char. #44), a feature they share with a few other Ericaceae (Fig. 18). In fact, this character may be a synapomorphy of Bejaria, Ledothamnus, and Bryanthus, but the placement of Bejaria is not, at present, strongly supported. Ledothamnus and Bryanthus also share ericoid leaves (char. #13). Possible additional synapomorphies for Ledothamnus and Bryanthus are the presence of adaxial calyx stomata (char. #42) and a style articulated with the ovary (char. #73). Finally, Ledothamnus and Bryanthus differ from Bejaria in the lack of an endothecium. The Bejarieae may be sister to Phyllodoceae (Fig. 7), although this relationship is not supported in some of the most parsimonious trees; both clades are characterized by homogeneous pith (char. #8), which may be synapomorphic.

b. Ericeae

Ericeae may be supported by a gradually tapering style (char. #73; Fig. 19). However, this character is quite homoplasious. A persistent corolla (char. #48; Fig. 20) is a synapomorphy for Calluna plus remaining Ericeae, but this character is lacking in Daboecia. As pointed out by Stevens (1971), Daboecia is rather similar to Erica in its anatomy, having flanges of lignifled tissue in the petiole; it also has small revolute leaves, which are nearly ericoid in form. Some South African species of Erica are wind pollinated and have dramatically expanded, cup-shaped or funnel-shaped stigmas.

c. Empetreae

The tribe comprises three genera: Ceratiola, Corema, and Empetrum. Samuelsson (1913) was the first to show that the Empetreae were closely related to the Ericaceae, and later Anderberg (1993), Judd and Kron (1993), and Kron and Chase (1993) demonstrated them to be nested within the Ericaceae. Prior to Samuelsson's (1913) investigation the relationships of empetrids to other plants were unclear, with various proposed systematic positions; for example, in the orders Euphorbiales (Don, 1827), Sapindales (Pax, 1891), or Celastrales (Hutchinson, 1969).

A position as a section of Conifers was put forward early in the nineteenth century (Nuttall, 1818). In recent years, most authors (e.g., Cronquist, 1981; Takhtajan, 1997) have placed empetrids close to the Ericaceae, albeit as a separate family.

Empetreae are easily diagnosed because this dade has numerous distinctive apomorphic characters, most of which relate to a shift from insect to wind pollination. Unlike most Ericaceae, Empetreae exhibit a high degree of dioecy (found elsewhere in Ericaceae only in Epigaea and possibly some Gaultherieae). Unique--or nearly unique--synapomorphies of Empetreae include imperfect flowers (char. #32, evolving independently in Epigaea), flowers with persistent tepals (chars. #39 and 48, although persistent corollas are found in most Ericeae), stigma flabellate to pinnatifid (char. #70), and drupaceous fruits (char. #77 and 81, occurring elsewhere only in Arbutoideae, some Styphelioideae, and some Vaccinioideae). Additional, more homoplasious synapomorphies of Empetreae are their lack of multicellular hairs on the leaves (char. #21), flowers with separate perianth parts (char. #44), stamens less than two times the number of petals (char. #51; Fig. 21), and indehiscent (fleshy) fruits (char. #78). In many-but not all- of the most parsimonious trees resulting from the combined morphology, marK, and rbcL analysis, the Empetreae and Ericeae form a dade, which is supported by the absence of bud scales (char. #9, also in some Phyllodoceae, Diplarche, Ledothamnus, Cassiope, and Harrimanella) and smooth filaments (char. #55, also quite homoplasious). In addition, the presence of 4- (or 2-) merous flowers may be synapomorphic, but this feature is also homoplasious. Both Empetreae and Ericeae have ericoid leaves (char. #13). However, it is noteworthy that the occurrence of gossypetin (char. #87) links Empetreae with Rhodoreae.

Although the data strongly indicate that Empetreae are derived from within Ericoideae, the exact relationships to the other tribes are not fully understood. In the analysis of morphological data set (Fig. 3) Empetreae are placed in the same clade as Erica and Calluna, but the support for this is very low. The matK data (Fig. 4) indicate a position for the Empetreae as sister to the Rhodoreae, but again with low support. In the combined analysis of morphological and molecular data sets, the position of Empetreae remains unresolved in relation to the Bejarieae, Ericeae, Rhodoreae, and Phyllodoceae. Relationships among the genera and species of Empetreae have been studied by Anderberg (1994b).

d. Rhodoreae

Morphological synapomorphies supporting the Rhodoreae include flowers on shoots of the previous season (char. #30), ovoid to cylindric fruits (char. #83), and the presence of gossypetin (char. #87). Rhodoreae have zygomorphic flowers with spots or a blotch (chars. #33 and 47), and it is probably significant that both of these features are found in Therorhodion, which is the sister taxon to the rest of the clade. It is probable that zygomorphic corollas evolved only once--in the common ancestor of Rhodoreae--and have been lost several times within this clade. Although this hypothesis is less parsimonious, based on the results presented here, than an assumption of parallelism, the hypothesis of multiple losses is preferred based on an analysis employing more taxa within this group (Kron & Judd, 1990). A major subgroup of Rhodoreae is diagnosed by scarious inflorescence bracts (char. #38), forming the perulate inflorescence characterizing the species-rich Rhododendron clade (which includes Tsusiophyllum and Menz iesia). The base chromosome number of 13 also characterizes the Rhododendron clade.

e. Phyllodoceae

The Phyllodoceae clade is not strongly supported by the data (Figs. 4-8). The molecular analyses indicate two strongly supported clades: Kalmia s.l. (including Leiophyllum and Loiseleuria) and a Phyllodoce clade (also including Epigaea, Kalmiopsis, and Rhodothamnus). In the rbcL analysis, these two clades are sisters, but without parsimony jackknife support. The matK data failed to resolve the Kalmia clade and the Phyllodoce clades as sisters. In both of the individual analyses Elliottia is unresolved with respect to the remaining Ericoideae. But in the combined molecular analysis Elliottia is sister to the Kalmia + Phyllodoce clade. This relationship is also indicated in the combined morphology and molecular tree (Fig. 7), but without bootstrap support. With the exception of Elliottia and Epigaea, the taxa that fall within the Phyllodoceae clade in this study have traditionally been considered closely related (e.g., Stevens, 1971). Further studies of the relationships of the Kalmia, Phyllodoce, and especiall y the Elliottia clades are in progress.

As here defined, Phyllodoceae are characterized by their articulated pedicels (char. #37, but this character also occurs in Bryanthus and may therefore be a synapomorphy for a Phyllodoceae + Bejarieae clade) and lack of abaxial calyx stomata (char. #43, but such stomata are present in Elliottia and Kalmia buxifolia). The monophyly of this group, like that of the Bejarieae, is thus not well supported by morphology. Kalmiopsis and Phyllodoceae are linked by their distinctive multicellular hairs with biseriate stalks (chars. #23 and 26), whereas members of the Kalmia group are possibly linked by the presence of an exothecium (char. #67). The monophyly of a clade comprising Leiophyllum, Loiseleuria, and Kalmia (i.e., Kalmia s.l.) also has been strongly supported in a detailed phylogenetic analysis of this group (Kron & King, 1996).

7. Harrimanelloideae

Within Clade 2, the sister-group relationship of Harrimanella with the Styphelioideae + Vaccinioideae clade is well supported (Fig. 7). However, this relationship is only supported by molecular data, and there are no discernable morphological synapomorphies for Glade 2. The Styphelioideae + Vaccinioideae dade is diagnosed by only a single homoplasious morphological feature--a persistent, non-withering calyx (char. #41), but the group is strongly supported by molecular characters.

8. Styphelioideae

The Styphelioideae include 35 genera and some 420 species. They are found throughout the Australasian region but are the most diverse and abundant in southwestern Western Australia, Tasmania, and southeastern Australia. Outliers extend the range to Tierra del Fuego (Lebetanthus), Hawaii (Styphelia), and Southeast Asia (Leucopogon).

The monophyly of the Styphelioideae is very well supported, and the group is easily diagnosed. Morphological apomorphies for Styphelioideae include the lignified leaf epidermis (char. #21; Fig. 22), scarious inflorescence bracts (char. #38), persistent corolla (char. #48), stamens less than two times the number of corolla lobes (char. #51; Fig. 21), smooth filaments (char. #55), bisporangiate, usually monothecal anthers (char. #57), and probably a branched vascular bundle in the petiole or leaf base (char. #6). Only the presence of monothecal anthers is not homoplasious (Fig. 23). Epidermal leaf lignification is uncommon in Ericaceae (Fig. 22), occurring outside Styphelioideae only in the Lyonieae. Anthers opening by longitudinal slits may also be synapomorphic. Core Styphelioideae--that is, all genera except Prionotes and Lebetanthus--have several additional synapomorphic features. These include parallel-(or more or less palmate-) veined leaves (char. #15) and a lack of multicellular hairs (char. #22). Epipe talous stamens (char. #52) has a single origin below Archerieae (Fig. 7). The stamens are free from the corolla in Andersonia, Lysinema, Richea, and Woollsia (Crayn et al., 1998), suggesting reversals of this character. For example, in Richea the corolla is deciduous, falling off at anthesis as a cap, precluding epipetaly. Outside Styphelioideae, only Diplarche (Rhodoreae) possesses truly epipetalous stamens. Traditionally (e.g., Bentham & Hooker, 1876; Brown, 1810), styphelioids have been divided into two subgroups, comprising the capsular-fruited and the fleshy-fruited taxa. However, leaf and stem anatomical characters also have been used as the basis for the primary division in the group (Watson, 1967).

More recently, Powell et al. (1996, 1997) conducted the first cladistic studies of the group and recognized four tribes, namely Richeeae (equivalent to Richeoideae sensu Watson, 1967), Cosmelieae, Styphelieae and Epacrideae, although the last appeared paraphyletic in their analysis. Parsimony analyses of nucleotide sequence data (Crayn & Quinn, 2000; Crayn et al., 1996, 1998) have supported the monophyly of some of these named groups but have clearly demonstrated others to be artificial. The results of the matK analysis in this study support the recognition of Prionoteae, Oligarrheneae, Richeeae, Epacrideae, Styphelieae, and Cosmelieae as named clades (Crayn & Quinn, 1998). In some of the most parsimonious trees, Archeria (i.e., Archerieae) branches after the Prionoteae but before the remaining epacrids. However, this relationship is not supported in the strict consensus, in which Archeria, Oligarrheneae, and remaining styphelioids (except Prionoteae) form a trichotomy. Sampling of epacrids in the rbcL and mo rphological analyses was not sufficient to test these relationships, but recent work using alternative DNA regions has given broad support to this classification (Crayn & Quinn, 2000; Crayn et al., 1998; Owens et al., in prep).

a. Prionoteae

The taxonomic history of Prionotes (one species, P. cerinthoides) and Lebethanthus (one species, L. myrsinites) exemplifies the power of cladistic analysis and the weakness of an overall similarity approach to classification and evolutionary relationships. Stevens (1971) discussed the characteristics that led to these taxa being alternately placed in Epacridaceae or Ericaceae (as previously circumscribed). Based on the concepts of overall similarity, these genera could not be satisfactorily placed in either group because they possessed a combination of characters of both "Ericaceae" and Epacridaceae. However, cladistic analysis shows that Prionotes and Lebetanthus possess many plesiomorphic characters with respect to the Styphelioideae (as here defined) while also possessing distinct synapomorphies of the group (such as a lignified leaf epidermis and monothecal anthers; Figs. 22 and 23, respectively). Prionotes and Lebetanthus form a dade that is sister to the remaining styphelioids in both the individual and combined molecular analyses in this study (see also Anderberg, 1993; Crayn et al., 1998).

b. Archerieae

The Archerieae comprise the small genus Archeria (6 spp.), native to Tasmania and New Zealand. Powell et al. (1996) considered Archeria related to Epacris, but this relationship is not supported by the matK analysis or other molecular analyses of Crayn et al. (1998) and Crayn and Quinn (2000). Relationships of Archerieae to the remaining styphelioids are discussed in detail in Crayn and Quinn (1998). Prionotes and Archeria share the character of flowers with bracteoles in the middle of the pedicel. This characteristic is also common in Vaccinioideae, which is consistent with its being plesiomorphic in Styphelioideae.

c. Oligarrheneae

Although not included in the morphological analysis presented here, Oligarrhena and Needhamiella form a dade in the matK analysis that is sister to Richeeae, Epacrideae, Cosmelieae, and Styphelieae. This dade is supported by the potential synapomorphies of a two-locular ovary and a glabrous corolla tube (Crayn et al., 1998). Traditionally, Oligarrhena and Needhamiella have been considered closely related to the Styphelia group (see Crayn et al., 1998 for discussion), but this relationship is not supported by the matK analysis presented here or by other molecular analyses of the styphelioids (Crayn et al., 1998).

d. Cosmelieae

This tribe comprises three genera, Andersonia, Cosmelia, and Sprengelia, and is strongly supported as a dade in the matK analysis presented here and in other molecular analyses (Crayn & Quinn, 2000; Crayn et al., 1998). Results of previous morphological studies (Powell et al., 1996) suggested that likely synapomorphies for Cosmelieae are the sheathing leaves that do not leave a scar and the unilacunar nodal anatomy. In the matK analysis (Fig. 4), Cosmelieae are sister to Styphelieae. This result differs from the rbcL analysis of Crayn et al. (1998), in which Cosmelieae are sister to the Epacrideae + Styphelieae. However, parsimony jackknife support for the rbcL branching pattern (Crayn et al., 1998) is lacking.

e. Richeeae

In contrast to Cosmelieae, Richeeae possess sheathing leaves that leave a distinct annular scar and possess nodes that are tri- or multilacunar (Crayn et al., 1998; Powell et al., 1996). In the analysis of Powell et al. (1996), the sheathing leaf base was a synapomorphy linking Cosmelieae and Richeeae, but in Cosmelieae the stem cortex detaches with the abscissing leaf, leaving an unscarred stem, whereas in the Richeae the cortex remains. These differences may indicate that the states are not homologous in the two tribes.

Richeeae are strongly supported in the matK analysis presented here, as well as in previous analyses using rbcL (Crayn et al., 1998). Three genera are recognized within Richeeae: Dracophyllum, Richea, and Sphenotoma. In the matK tree (Fig. 4), Richeeae are sister to Cosmeliaeae

+ Styphelieae, but this relationship lacks parsimony jackknife support. The rbcL data (Crayn et al., 1998) indicate Richeeae as branching earlier in the history of Styphelioideae than the matK data presented in this study, but again without parsimony jackknife support. Because of this lack of support Richeeae are indicated as sedis mutabilis in the classification (Fig. 1).

f. Epacrideae

Epacrideae comprise five genera: Budawangia, Epacris, Lysinema, Rupicola, and Woollsia. In the matK analysis (Fig. 4) they are represented only by Epacris, Lysinema, and Rupicola. This is a well-supported dade in the rbcL analysis (Fig. 5) as well as in the more comprehensive studies by Crayn et al. (1998). Generic relationships within this group have been studied by Powell et al. (1996) and Crayn et al. (1998) but require further investigation because preliminary results indicate that the recognition of Budawangia and Rupicola would result in a paraphyletic Epacris (Crayn et al., 1998).

g. Styphelieae

The Styphelieae, which are characterized by fleshy fruit, have the broadest distribution of the styphelioid tribes, being found in all parts of the range of the subfamily except for South America. It is also the largest tribe, with 20 currently recognized genera and at least 320 species (Powell et al., 1987; Weiller, 1996a, 1996b). Although some genera--Acrotriche, Astroloma, Brachyloma, Leucopogon, Monotoca, and Styphelia--show east-west Australian disjunctions, several of these genera are likely not monophyletic (Powell et al., 1997). Relationships within Styphelieae are complex. Recent morphological (Powell et al., 1997) and molecular studies (Taaffe et al., 2001) indicate that Leucopogon as traditionally defined is polyphyletic, but details of the relationships have yet to be resolved (Taaffe et al., 2001). As a dade, Styphelieae are strongly supported as monophyletic (Figs. 4-6). Previous morphological studies have indicated that Styphelieae have the following synapomorphies: fruit drupaceous, ovary 1-11 locular (each locule containing a single ovule), and the corolla mostly hairy inside (Taaffe et al., 2001). The fleshy fruits of Styphelieae are homoplasious, as this character has evolved several times within Ericaceae (e.g., Arbutoideae, Empetreae, and Vaccinieae).

9. Vaccinioideae

Two homoplasious characters diagnose Vaccinioideae: the presence of disintegration tissue on the back of the anthers (char. #63; Fig. 24) and a base chromosome number of 12 (char. #89; Fig. 25). However, in the combined molecular (Fig. 6) and the combined morphological and molecular analyses (Fig. 7), Vaccinioideae are strongly supported. This circumscription of Vaccinioideae differs from that proposed by Watson et al. (1967) and Stevens (1971), because it does not include Arbutoideae, Enkianthus, Cassiope, or Harrimanella. A previous analysis using matK and a larger sample of taxa within the traditional "Vaccinioideae" indicates the same pattern of relationships that was found in this study (Kron et al., 1999a).

a. Oxydendreae

Oxydendrum is sister to the remaining Vaccinioideae and is distinctive because its terminal inflorescence produces fruits in the same year the shoot is initiated (Lems, 1962). Oxydendrum possesses distinctive floral anatomy--all traces to the floral organs leave the elongated floral axis separately (Palser, 1952)--and the elongate-ovoid capsules are unique to the genus. Its leaves have paracytic stomata, which are of limited occurrence in Ericaceae but may be found in most Gaultherieae. Finally, the basal placentae and stout, non-glandular, multicellular hairs are distinctive. The remaining genera of Vaccinioideae belong to one of four major clades: Lyonieae, Vaccinieae, Andromedeae, or Gaultherieae. Together, these four tribes may form a dade, which can be diagnosed by its inflorescences axillary at initiation (char. #29; Fig. 12), developing on shoots of the previous season (char. #30), and bearing flowers with articulated pedicels (char. #37; Fig. 26).

b. Lyonieae

Lyonieae are clearly supported as the sister group to a large clade comprising Vaccinieae, Andromedeae, and Gaultherieae. The Lyonieae are diagnosed by bands of fibers in the phloem (char. #7; Fig. 27), a lignified leaf epidermis (char. #21; Fig. 22), and S-shaped filaments (char. #53, which have been lost in some species of Pieris and have evolved in parallel in some Styphelioideae and Ericoideae). The phylogenetic relationships within this dade are discussed in detail in Kron and Judd (1997) and Kron et al. (1999b). The monophyly of the Gaultherieae + Andromedeae + Vaccinieae clade is not apparent morphologically, although the clade has a bootstrap value of 98% in the combined analysis (Fig. 7). Most of the members of this dade, however, possess paracytic stomata (char. #28), although anomocytic stomata are present in Andromeda and Zenobia (Fig. 28).

c. Vaccinieae

The Vaccinieae are easily recognized because they possess a distinctive combination of synapomorphic features: anther tubules (char. #62, also found in parallel in a few Gaultherieae), an inferior ovary (char. #72; Fig. 29), and indehiscent, brightly colored fruits (chars. #78 and 82), usually berries, but drupes in Gaylussacia. It is worth noting that fleshy, colorful fruits, either drupes or berries, have also evolved in some Styphelioideae (i.e., Styphelieae), some Gaultherieae, Empetreae, and Arbutoideae (Fig. 30), likely in connection with a shift from wind to bird dispersal. Generic relationships within Vaccinieae are problematic (Kron et al., 1999a; Luteyn, 1996) and are currently being investigated by Kron and Luteyn.

d. Andromedeae

Traditionally this tribe was broadly circumscribed, including genera here placed in Lyonieae, Gaultherieae, and Oxydendreae (Stevens, 1971). As traditionally delimited, the group clearly is paraphyletic (Fig. 7; see also Kron et al., 1999b; Stevens, 1995). In this classification Andromedeae is restricted to Andromeda and Zenobia and may be characterized by a lack of multicellular hairs on the leaves (disregarding the reduced non-glandular hairs associated with the marginal teeth of Zenobia; see Kron et al., 1999b). The leaves of Andromeda are entire and thus lack these tooth-associated hairs (and any other multicellular hairs). Despite the lack of distinctive morphological synapomorphies, the monophyly of this small dade is strongly supported in the molecular (Figs. 4-6) and combined analyses (Fig. 7).

e. Gaultherieae

Although the Gaultherieae clade is present in all of the equally parsimonious trees discovered in the combined analysis (Fig. 7), its monophyly is not (as yet) well supported. It is probably most closely related to the Andromedeae, and it is likely significant that 4-appendaged anthers (char. #61; Fig. 3 1) and a base chromosome number of 11 (char. #89) characterize Gaultherieae and also occur in Zenobia. A strongly supported subclade comprising Diplycosia. Tepuia, Gauliheria, and Pernettya is diagnosed by the presence of methyl salicylate (char. #88). This fragrant compound, which gives the plants a distinct wintergreen odor, has been lost in several species, especially within Gaultheria. Generic relationships are problematic within this subclade, and phylogenetic relationships within this group are currently being investigated by Powell and Kron (2001).

VI. Conclusions

It is noteworthy that some major clades within Ericaceae--Enkianthus, Monotropoideae, Pyroleae, Monotropeae, Arbutoideae, Ericeae (except Daboecia), Empetreae, Rhodoreae (except Diplarche) , Styphelioideae, Lyonieae, and Vaccinieae--are well supported morphologically, and historically most of these have received taxonomic recognition at various taxonomic ranks. In contrast, other clades--Pterosporeae, Ericoideae, Bejarieae, Phyllodoceae, Andromedeae, and Gaultherieae--which also have strong support in the morphology + matK + rbcL tree, are morphologically heterogeneous, possess only a few, usually homoplasious synapomorphies, and usually have not been accorded taxonomic recognition, at least as circumscribed herein. The early anther inversion clade is a special case. It has not been recognized previously, even though it can be delimited by a uniquely derived character. This likely results from the cryptic nature of this developmental feature. Other cryptic characters, such as the lignified epidermal cells and the bands of fibers in the secondary phloem, also have been somewhat overlooked in the classification of the family, and a group diagnosed by these characters-the Lyonieae-has only recently received formal taxonomic recognition (Kron & Judd, 1997; Kron et al., 1999b), although these characters had been noted earlier by Stevens (1970a, 1971). Likewise, the phenotypic distinctiveness of Harrimanella and Cassiope, many of which are anatomical, have only been addressed by Stevens (1970b). The same goes for Enkianthus, which until Stevens (1971) had been retained within a broadly circumscribed Andromedeae (see also Anderberg, 1994a).

An obvious conclusion is that the phylogeny of Ericaceae has often been only imperfectly reflected in traditional classifications. This is true not only because previous classifications usually lacked a phylogenetic perspective but also because undue emphasis was placed on certain visually striking morphological characteristics, especially on certain aspects of reproductive morphology. Some of these characters, including inflorescence position and "winter" development, bracteole position, indumentum of the corolla and filaments, anther texture, presence and positioning of staminal appendages (spurs, awns, tubules, etc.), and fruit dehiscence and texture (leading to various fruit types), are extremely homoplasious. In Empetreae, Bejarieae, some Phyllodoceae (Kalmia buxifolia, Elliottia), Pyroleae (Fig. 18), and some Monotropeae (Pityopus, Pleuricospora, Monotropa), separate petals (or tepals) have evolved several times from sympetalous ancestors. Likewise, it appears that stamen appendages, either spurs or awn s, have been repeatedly lost within Ericaceae (Fig. 15). It is also apparent that indehiscent, fleshy fruits have evolved multiple times (Fig. 30), and even the pattern of capsule dehiscence seems quite homoplasious (Fig. 16). These and other, similar patterns may relate to changes in selective forces resulting from shifts in pollinators or dispersal agents, and they are currently under detailed investigation (Kron et al., in prep.). Certain vegetative features, such as leaf persistence and curvature (chars. #13 and 17), are also very homoplasious and are under obvious selective forces. Deciduous leaves have repeatedly evolved from an evergreen ancestral condition (Fig. 32), and reversals back to evergreen leaves are also likely (Kron & Judd, 1990). Traditional classifications that incorporated detailed anatomical and developmental data, such as that of Stevens (1971), therefore, match our classification (see Appendix 3) better than do those that stress easily observed features (such as those typically report ed in floras). Admittedly, nearly all morphological features show some homoplasy.

We conclude that the phylogeny of Ericaceae, or that of any other large and diverse group, is best approached through a detailed phylogenetic analysis, based on several genes and a detailed survey of phenotypic characters. Patterns of morphological change can then be traced onto the combined-data trees, allowing not only clarification of clade diagnosis but also a better understanding of patterns of character change.

The new classification of Ericaceae presented here is intended to provide a framework for future studies that address various aspects of the biology of these fascinating plants. In addition, it is hoped that the identification key (Appendix 4) and morphological characterizations provided with each named group will facilitate the incorporation of the new classification into future floras.
IX. Appendix 1

Species Sampled in the Molecular Systematic Study of Ericaceae

                                            Family or
Species                                     subfamily

Outgroups

Actinidia chinensis Planchon                Actinidiaceae
Clethra alnifolia L.                        Clethraceae
Cyrilla racemiflora L.                      Cyrillaceae
Diapensia lapponica L.                      Diapensiaceae
Diospyros virginiana L.                     Ebenaceae
Symplocos paniculata Miq.                   Symplocaceae

Ingroup (=Ericaceae)

Agapetes buxifolia Nutt. ex Hook. f.        Vaccinioideae
Agapetes hosseana Diels                     Vaccinioideae
Agapetes scortechinii (King &               Vaccinioideae
 Gamble) Sleumer
Agapetes serpens (Wight) Sleumer            Vaccinioideae
Agarista populifolia (Lam.) Judd            Vaccinioideae
Agarista salicifolia                        Vaccinioideae
 (Comm. ex Lam.) G. Don
Allotropa virgata Torrey & A.               Monotropoideae
 Gray ex A. Gray
Andersonia sprengelioides R. Br.            Styphelioideae
Andromeda glaucophylia Link                 Vaccinioideae
Andromeda polifolia L.                      Vaccinioideae
Arbutus canariensis Duhamel                 Arbutoideae
Arbutus menziesii Pursh                     Arbutoideae
Archeria comberi Melville                   Styphclioideae
Arctostaphylos alpina (L.) Sprengel         Arbutoideae
Arctostaphylos tomentosa Lindley            Arbutoideae
Arctostaphylos uva-ursi (L.) Sprengel       Arbutoideae
Astroloma humifusum (Cav.) R. Br.           Styphelioideae
Bejaria racemosa Vent.                      Ericoideae
Bejaria resinosa Mutis ex L.f.              Ericoideac
Brachyloma daphnoides (Sm.) Benth.          Styphclioideae
Bryanthus gmelini D. Don                    Ericoideae
Calluna vulgaris (L.) Hull                  Ericoideae
Cassiope mertensiana (Bong.) G. Don         Cassiopoideae
Cavendishia complectens Hemsl.              Vaccinioideae
Cavendishia tarapotana (Meissner) Benth.    Vaccinioideae
 & Hook.
Ceratiola ericoides Michaux                 Ericoideae
Chamaedaphne calyculata (L.) Moench         Vaccinioideae
Chimaphila maculata (L.) Pursh              Monotropoideae
Chimaphila menziesii Sprengel               Monotropoideae
Chimaphila umbellata (L.) Barton            Monotropoideae
Comarostaphylis arbutoides Lindley          Arbutoideae
Corema conradi Torrey ex Loud.              Ericoideac
Cosmelia rubra R. Br.                       Styphclioideae
Costera endertii J. J. Smith                Vaccinioideae
Craibiodendron yunnanense W. Smith          Vaccinioideae
Cyathodes colensoi Hook.                    Styphelioideae
Daboecia cantabrica (Hudson) K. Koch        Ericoideae
Dimorphanthera dekockii J. J. Smith         Vaccinioideae
Diplarche multiflora Hook. f. & Thomson     Ericoideae
Diplycosia acuminala Becc.                  Vaccinioideae
Disterigma trimerum Wilbur & Luteyn         Vaccinioideae
Dracophyllum longifolium (J.R. Forster)     Styphelioideae
 Roem. & Schultes
Elliottia bracteata (Maxim.) Hook. f.       Ericoideac
Elliottia racemosa Muhl. ex Elliott         Ericoideac
Empetrum atropurpureum Fernald & Wiegand    Ericoideac
Empetrum nigrum L.                          Ericoideac
Empetrum rubrum Vahl                        Ericoideae
Enkianthus campanulatus G. Nicholson        Enkianthiodeae
Epacris impressa Labill.                    Styphelioideae
Epigaea repens L.                           Ericoideae
Erica australis L.                          Ericoideae
Erica sicula Guss.                          Ericoideae
Erica spiculifolia Salisb.                  Ericoideae
Erica tetralix L.                           Ericoideae
Eubotrys racemosa L.                        Vaccinioideae
Gaultheria eriophylla (Pers.) Sleumer       Vaccinioideae
Gaultheria miqueliana Takeda                Vaccinioideae
Gaultheria shallon Pursh                    Vaccinioideae
Gaylussacia frondosa (L.) Torrey & A. Gray  Vaccinioideae
Harrimanella hypnoides Cov.                 Harrimanelloideae
Kalmia angustifolia L.                      Ericoideae
Kalmia buxifolia (Berg) Gift, Kron &        Ericoideae
 Stevens
Kalmia latifolia L.                         Ericoideae
Kalmia polifolia Wangenh.                   Ericoideae
Kalmia procumbens (L.) Gift,  Kron &        Ericoideae
 Stevens
Kalmiopsis leachiana (Henderson) Rehder     Ericoideae
Lebetanthus myrsinites Macl.                Styphelioideae
Ledothamnus guyanensis Meissner             Ericoideae
Leucopogon fraseri A.M. Cunn.               Styphelioideae
Leucopogon muticus R. Br.                   Styphelioideae
Leucothoe fontanesiana (Steudel) Sleumer    Vaccinioideae
Lyonia ferruginea (Walter) Nutt.            Vaccinioideae
Lyonia ligustrina (L.) DC.                  Vaccinioideae
Lyonia lucida (Lam.) K. Koch                Vaccinioideae
Lyonia ovalifolia (Wall) Drude              Vaccinioideae
Lysinema ciliatum R. Br.                    Styphelioideae
Macleania bullata Yeo                       Vaccinioideae
Macleania rotundifolia Sodiro & Hoerold     Vaccinioideae
Macleania rupestris (Kunth)A. C. Smith      Vaccinioideae
Menziesia ciliicalyx Maxim.                 Ericoideae
Menziesia pilosa Pers.                      Ericoideae
Monotoca scoparia (W. Smith) R. Br.         Styphelioideae
Monotropa hypopithys L.                     Monotropoideae
Needhamiella pumilio (R. Br.) L. Watson     Styphelioideae
Oligarrhena micrantha R. Br.                Styphelioideae
Orthilia secunda (L.) House                 Monotropoideae
Oxydendrum arboreum (L.) DC.                Vaccinioideae
Paphia meiniana (F.Muell.) Schltr.          Vaccinioideae
Pentachondra pumila                         Styphelioideae
 (Forster & G. Forster) R. Br.
Pernettya tasmanica Hook. f.                Vaccinioideae
Phyllodoce caerulea Bab.                    Ericoideae
Phyllodoce empetriformis D. Don             Ericoideae
Phyllodoce nipponica Makino                 Ericoideae
Pieris floribunda (Pursh) Benth. & Hook.    Vaccinioideae
Pieris formosa (Wallich) D. Don             Vaccinioideae
Pieris nana (Maxim.) Makino                 Vaccinioideae
Pieris phillyraeifolia (Hook.) DC.          Vaccinioideae
Prionotes cerinthoides (Labill.) R. Br.     Styphelioideae
Psammisia columnbiensis Hoerold             Vaccinioideae
Psammisia dolichopoda A. C. Smith           Vaccinioideae
Psammisia sclerantha A. C. Smith            Vaccinioideae
Pyrola picta Smith                          Monotropoideae
Pyrola rotundifolia L.                      Monotropoideae
Rhododendron albiflorum Hook.               Ericoideae
Rhododendron canescens (Michaux) Sweet      Ericoideae
Rhododendron edgeworthii Hook.              Ericoideae
Rhododendron falconeri Hook. f.             Ericoideae
Rhododendron farrerae Tate                  Ericoideae
Rhododendron grande Wight                   Ericoideae
Rhododendron groenlandicum (Oeder)          Ericoideae
 Kron & Judd
Rhododendron hippophaeoides Balf.           Ericoideae
 f. Forrest
Rhododendron hongkongense Hutch.            Ericoideae
Rhododendron kaempferi Planchon             Ericoideae
Rhododendron kiusiamum Makino               Ericoideae
Rhododendron maximum L.                     Ericoideae
Rhododendron molle (Blume) G. Don           Ericoideae
Rhododendron ovatum (Lindley)               Ericoideae
 Maxim.
Rhododendron pendulum Hook. f.              Ericoideae
Rhododendron semibarbatum Maxim.            Ericoideae
Rhododendron stamineum Franchet             Ericoideae
Rhododendron tomentosum (Stokes)            Ericoideae
 Harmaja
Rhododendron lsusiophyllum Sugim.           Ericoideae
Rhododendron virgatum Hook. f.              Ericoideae
Rhodothammus chamaecistus Rchb.             Ericoideae
Richea pandanifolia Hook. f.                Styphelioideae
Rupicola sprengeloides Maiden &             Stypehlioideae
 Betche
Satyria warszewiczii Klotzsch               Vaccinioideae
Sphenotoma dracophylloides Sonder           Satyphelioideae
Sphyrospermum buxifolium Popepping          Vaccinioideae
 & Endl.
Sphyrospermum cordifolium Benth.            Vaccinioideae
Sprengelia incarnata W. Smith               Styphelioideae
Symphysia racemosa (Vahl) Stearn            Vaccinioideae
Tepuia cardonae A. C. Smith                 Vaccinioideae
Themistoclesia costaricensis Luteyn         Vaccinioideae
 & Wilbur
Therorhodion camtschaticum (Pallas)
 Small                                      Ericoideae
Vaccinium cercidifolium J. J. Smith         Vaccinioideae
Vaccinium crenatum
 (D. Don ex Dunal) Sleumer                  Vaccinioideae
Vaccinium macrocarpon Aiton                 Vaccinioideae
Vaccinium meridionale Sw.                   Vaccinioideae
Vaccinium myrsinites Lam.                   Vaccinioideae
Vaccinium myrtillus L.                      Vaccinioideae
Vaccinium poasamum J. D. Smith              Vaccinioideae
Vaccinium vitis-idaea L.                    Vaccinioideae
Zenobia pulverulenta (Bartram)
 Pollard                                    Vaccinioideae


Species                                     Voucher and source

Outgroups

Actinidia chinensis Planchon                Kron 2117, NCU
Clethra alnifolia L.                        Kron 1884, NCU
Cyrilla racemiflora L.                      Kron s.n., NCU
Diapensia lapponica L.                      Hills 89018, NCU
Diospyros virginiana L.                     Kron 3004, NCU
Symplocos paniculata Miq.                   Kron 3005, NCU

Ingroup (=Ericaceae)

Agapetes buxifolia Nutt. ex Hook. f.        RBGK 1969-16943
Agapetes hosseana Diels                     RBGE 1967-2592
Agapetes scortechinii (King &               RBGE 1968-0672
 Gamble) Sleumer
Agapetes serpens (Wight) Sleumer            RBGK 1969-17805
Agarista populifolia (Lam.) Judd            Judd DNA190, WFU
Agarista salicifolia                        RBGE 1973-0764A
 (Comm. ex Lam.) G. Don
Allotropa virgata Torrey & A.               Kron DNA200, WFU
 Gray ex A. Gray
Andersonia sprengelioides R. Br.            UNSW 202801
Andromeda glaucophylia Link                 Powell s.n., WFU
Andromeda polifolia L.                      RBGK 1976-6099
Arbutus canariensis Duhamel                 RBGE 1975-2562
Arbutus menziesii Pursh                     RBGE 1941-0182
Archeria comberi Melville                   UNSW 22567
Arctostaphylos alpina (L.) Sprengel         Nordenstam BN9254, S
Arctostaphylos tomentosa Lindley            RBGE 1963-7636
Arctostaphylos uva-ursi (L.) Sprengel       AA 361-68
Astroloma humifusum (Cav.) R. Br.           UNSW 22502
Bejaria racemosa Vent.                      Kron 2070, NCU
Bejaria resinosa Mutis ex L.f.              Luteyn 15067, NYBG
Brachyloma daphnoides (Sm.) Benth.          UNSW 22543
Bryanthus gmelini D. Don                    Stevens s.n., WFU
Calluna vulgaris (L.) Hull                  RBGE 721433
Cassiope mertensiana (Bong.) G. Don         AA 75-83
Cavendishia complectens Hemsl.              Luteyn s.n., NYBG
Cavendishia tarapotana (Meissner) Benth.    Luteyn s.n., NYBG
 & Hook.
Ceratiola ericoides Michaux                 Kron 2069, NCU
Chamaedaphne calyculata (L.) Moench         RBGE 1975-2181
Chimaphila maculata (L.) Pursh              Kron DNA164, WFU
Chimaphila menziesii Sprengel               Kron DNA164, WFU
Chimaphila umbellata (L.) Barton            Kron DNA164, WFU
Comarostaphylis arbutoides Lindley          Luteyn 14815, NYBG
Corema conradi Torrey ex Loud.              Stevens s.n., A
Cosmelia rubra R. Br.                       UNSW 23107
Costera endertii J. J. Smith                RBGE 1980-1343
Craibiodendron yunnanense W. Smith          RBGE 1982-5379
Cyathodes colensoi Hook.                    RBGE 1969-4055
Daboecia cantabrica (Hudson) K. Koch        RBGE 1975-1770
Dimorphanthera dekockii J. J. Smith         RBGK 1975-1556
Diplarche multiflora Hook. f. & Thomson     Suzuki et al., GH
Diplycosia acuminala Becc.                  RBGE 1982-0862
Disterigma trimerum Wilbur & Luteyn         Luteyn 14809, NYBG
Dracophyllum longifolium (J.R. Forster)     RBGE 1976-1673
 Roem. & Schultes
Elliottia bracteata (Maxim.) Hook. f.       RBGK 1979-5019
Elliottia racemosa Muhl. ex Elliott         RBGE 1967-2632
Empetrum atropurpureum Fernald & Wiegand    RBGK 1988-433
Empetrum nigrum L.                          Hills 89024, NCU
Empetrum rubrum Vahl                        RBGK 1976-77
Enkianthus campanulatus G. Nicholson        AA 14528-C
Epacris impressa Labill.                    RBGE 1968-0556
Epigaea repens L.                           Kron DNA162, WFU
Erica australis L.                          RBGE 781912
Erica sicula Guss.                          RBGK 1985-2612
Erica spiculifolia Salisb.                  RBGK 1969-52067
Erica tetralix L.                           AA 195-79
Eubotrys racemosa L.                        Kron s.n., NCU
Gaultheria eriophylla (Pers.) Sleumer       RBGE 1976-3043
Gaultheria miqueliana Takeda                AA 1636-77
Gaultheria shallon Pursh                    Kron DNAl85, WFU
Gaylussacia frondosa (L.) Torrey & A. Gray  Kron 1889, WFU
Harrimanella hypnoides Cov.                 Nordenstarn 9252, S
Kalmia angustifolia L.                      Kron DNA 156, WFU
Kalmia buxifolia (Berg) Gift, Kron &        Kron 2067, NCU
 Stevens
Kalmia latifolia L.                         Kron 3020, WFU
Kalmia polifolia Wangenh.                   Kron DNAl59, WFU
Kalmia procumbens (L.) Gift,  Kron &        AA s.n., S
 Stevens
Kalmiopsis leachiana (Henderson) Rehder     Kron DNA 177, WFU
Lebetanthus myrsinites Macl.                Pisano 5273, GH
Ledothamnus guyanensis Meissner             Picon & Williams 29l0, WFU
Leucopogon fraseri A.M. Cunn.               RBGE 1934-1059
Leucopogon muticus R. Br.                   Powell 4866, UNSW
Leucothoe fontanesiana (Steudel) Sleumer    Kron 1876, WFU
Lyonia ferruginea (Walter) Nutt.            Kron 2068, WFU
Lyonia ligustrina (L.) DC.                  Kron 1892, WFU
Lyonia lucida (Lam.) K. Koch                Kron 1891, WFU
Lyonia ovalifolia (Wall) Drude              RBGE 1975-0214
Lysinema ciliatum R. Br.                    UNSW 22193
Macleania bullata Yeo                       Luteyn s.n., NYBG
Macleania rotundifolia Sodiro & Hoerold     Dieterman, ABG
Macleania rupestris (Kunth)A. C. Smith      Luteyn s.n., NYBG
Menziesia ciliicalyx Maxim.                 RBGE 1969-5350
Menziesia pilosa Pers.                      Kron DNA157, WFU
Monotoca scoparia (W. Smith) R. Br.         UNSW 22188
Monotropa hypopithys L.                     Kron DNA98, WFU
Needhamiella pumilio (R. Br.) L. Watson     UNSW 22976
Oligarrhena micrantha R. Br.                UNSW 22956
Orthilia secunda (L.) House                 Kron DNA182, WFU
Oxydendrum arboreum (L.) DC.                Kron DNA161, WFU
Paphia meiniana (F.Muell.) Schltr.          Gadek 18:3, JCU-C
Pentachondra pumila                         RBGE 1968-1038
 (Forster & G. Forster) R. Br.
Pernettya tasmanica Hook. f.                RBGK 1977-5050
Phyllodoce caerulea Bab.                    RBGE 1940-1013
Phyllodoce empetriformis D. Don             RBGK 1978-2120
Phyllodoce nipponica Makino                 AA s.n.
Pieris floribunda (Pursh) Benth. & Hook.    AA 21301
Pieris formosa (Wallich) D. Don             RBGE 1947-1034
Pieris nana (Maxim.) Makino                 RBGK 1979-4813
Pieris phillyraeifolia (Hook.) DC.          Judd DNA191, WFU
Prionotes cerinthoides (Labill.) R. Br.     RBGK 1984-4432
Psammisia columnbiensis Hoerold             Luteyn s.n., NYBG
Psammisia dolichopoda A. C. Smith           Luteyn s.n., NYBG
Psammisia sclerantha A. C. Smith            Luteyn s.n., NYBG
Pyrola picta Smith                          Kron DNA183, WFU
Pyrola rotundifolia L.                      Kron 1906, NCU
Rhododendron albiflorum Hook.               RSF 77/254
Rhododendron canescens (Michaux) Sweet      Kron s.n., WFU
Rhododendron edgeworthii Hook.              RBGE 1969-8546
Rhododendron falconeri Hook. f.             RBGE 1971-5025
Rhododendron farrerae Tate                  RSF 78/037
Rhododendron grande Wight                   RBGE 1969-8606
Rhododendron groenlandicum (Oeder)          Hills 89022, NCU
 Kron & Judd
Rhododendron hippophaeoides Balf.
 f. Forrest                                 RBGE 1932-1022
Rhododendron hongkongense Hutch.            RSF 77/779
Rhododendron kaempferi Planchon             RBGE 1976-1898
Rhododendron kiusiamum Makino               RBGE 1919-1029
Rhododendron maximum L.                     RBGE 1973-401
Rhododendron molle (Blume) G. Don           RBGE 1981-0379
Rhododendron ovatum (Lindley)               RSF 76/050
 Maxim.
Rhododendron pendulum Hook. f.              RSF 76/141
Rhododendron semibarbatum Maxim.            RBGE 1977-1391
Rhododendron stamineum Franchet             RSF 70/369
Rhododendron tomentosum (Stokes)            RBGK 1979-1730
 Harmaja
Rhododendron lsusiophyllum Sugim.           RBGK 1985-4647
Rhododendron virgatum Hook. f.              RSF 66/672
Rhodothammus chamaecistus Rchb.             RBGK 1989-459
Richea pandanifolia Hook. f.                UNSW 22534
Rupicola sprengeloides Maiden &             UNSW 22545
 Betche
Satyria warszewiczii Klotzsch               RBGE 1978-1009
Sphenotoma dracophylloides Sonder           CBG 7900865
Sphyrospermum buxifolium Popepping          VanderKloet, ACAD
 & Endl.
Sphyrospermum cordifolium Benth.            Luteyn s.n., NYBG
Sprengelia incarnata W. Smith               UNSW 22539
Symphysia racemosa (Vahl) Stearn            VanderKloet, ACAD
Tepuia cardonae A. C. Smith                 Anderson 13870, MICH
Themistoclesia costaricensis Luteyn         Luteyn 14996, NYBG
 & Wilbur
Therorhodion camtschaticum (Pallas)
 Small                                      RBGe 1969-9487B
Vaccinium cercidifolium J. J. Smith         RBGE 1982-0845
Vaccinium crenatum
 (D. Don ex Dunal) Sleumer                  Luteyn s.n., NYBG
Vaccinium macrocarpon Aiton                 Hills 89019, NCU
Vaccinium meridionale Sw.                   VanderKloet, ACAD
Vaccinium myrsinites Lam.                   Kron DNA189, WFU
Vaccinium myrtillus L.                      Anderberg s.n., S
Vaccinium poasamum J. D. Smith              VanderKloet, ACAD
Vaccinium vitis-idaea L.                    RBGe 1977-3274A
Zenobia pulverulenta (Bartram)
 Pollard                                    AA 21087

                                              GenBank or EMBL accession
                                                       number
Species                                         18s           rbcL

Outgroups

Actinidia chinensis Planchon                 AF419792        L01882
Clethra alnifolia L.                         AF419793        L12609
Cyrilla racemiflora L.                        U43294         L01900
Diapensia lapponica L.                       AF419794        L12612
Diospyros virginiana L.                       U43295         L12613
Symplocos paniculata Miq.                     U43297         L12624

Ingroup (=Ericaceae)

Agapetes buxifolia Nutt. ex Hook. f.            NA          AF419811
Agapetes hosseana Diels                         NA             NA
Agapetes scortechinii (King &                   NA             NA
 Gamble) Sleumer
Agapetes serpens (Wight) Sleumer                NA             NA
Agarista populifolia (Lam.) Judd                NA          AF124589
Agarista salicifolia                            NA          AF124588
 (Comm. ex Lam.) G. Don
Allotropa virgata Torrey & A.                   NA             NA
 Gray ex A. Gray
Andersonia sprengelioides R. Br.                NA             NA
Andromeda glaucophylia Link                     NA          AF419812
Andromeda polifolia L.                          NA          AF124572
Arbutus canariensis Duhamel                  AF419795        L12597
Arbutus menziesii Pursh                         NA          AF419813
Archeria comberi Melville                       NA             NA
Arctostaphylos alpina (L.) Sprengel             NA          AF419814
Arctostaphylos tomentosa Lindley                NA          AF419815
Arctostaphylos uva-ursi (L.) Sprengel           NA           L12598
Astroloma humifusum (Cav.) R. Br.               NA           U80433
Bejaria racemosa Vent.                       AF419796        L12600
Bejaria resinosa Mutis ex L.f.                  NA             NA
Brachyloma daphnoides (Sm.) Benth.              NA             NA
Bryanthus gmelini D. Don                        NA          AF419816
Calluna vulgaris (L.) Hull                   AF419797        L12601
Cassiope mertensiana (Bong.) G. Don          AF419798        L12603
Cavendishia complectens Hemsl.                  NA             NA
Cavendishia tarapotana (Meissner) Benth.        NA          AF419817
 & Hook.
Ceratiola ericoides Michaux                  AF419799        L12605
Chamaedaphne calyculata (L.) Moench          AF419800        L12606
Chimaphila maculata (L.) Pursh                  NA             NA
Chimaphila menziesii Sprengel                   NA             NA
Chimaphila umbellata (L.) Barton                NA          AF419818
Comarostaphylis arbutoides Lindley              NA          AF419819
Corema conradi Torrey ex Loud.                  NA          AF419820
Cosmelia rubra R. Br.                        AF419801        U80420
Costera endertii J. J. Smith                    NA             NA
Craibiodendron yunnanense W. Smith              NA          AF124589
Cyathodes colensoi Hook.                        NA           L12610
Daboecia cantabrica (Hudson) K. Koch            NA           L12611
Dimorphanthera dekockii J. J. Smith             NA             NA
Diplarche multiflora Hook. f. & Thomson         NA          AF419821
Diplycosia acuminala Becc.                      NA          AF124586
Disterigma trimerum Wilbur & Luteyn             NA             NA
Dracophyllum longifolium (J.R. Forster)      AF419809        L12614
 Roem. & Schultes
Elliottia bracteata (Maxim.) Hook. f.           NA           U49285
Elliottia racemosa Muhl. ex Elliott             NA           L12615
Empetrum atropurpureum Fernald & Wiegand        NA             NA
Empetrum nigrum L.                              NA         AF4 19822
Empetrum rubrum Vahl                            NA             NA
Enkianthus campanulatus G. Nicholson         AF419802        L12616
Epacris impressa Labill.                     AF419803        L01915
Epigaea repens L.                               NA           U49284
Erica australis L.                              NA           L12617
Erica sicula Guss.                              NA          AF419823
Erica spiculifolia Salisb.                      NA          AF419824
Erica tetralix L.                               NA          AF419825
Eubotrys racemosa L.                            NA           U83915
Gaultheria eriophylla (Pers.) Sleumer           AF           L12618
Gaultheria miqueliana Takeda                    NA          AF124590
Gaultheria shallon Pursh                        NA          AF124574
Gaylussacia frondosa (L.) Torrey & A. Gray      NA          AF419836
Harrimanella hypnoides Cov.                     NA           U82766
Kalmia angustifolia L.                          NA          AF419826
Kalmia buxifolia (Berg) Gift, Kron &            NA           L12619
 Stevens
Kalmia latifolia L.                             NA           U49294
Kalmia polifolia Wangenh.                       NA           U49289
Kalmia procumbens (L.) Gift,  Kron &            NA           U49288
 Stevens
Kalmiopsis leachiana (Henderson) Rehder         NA           U49290
Lebetanthus myrsinites Macl.                    NA           U81797
Ledothamnus guyanensis Meissner                 NA          AF419827
Leucopogon fraseri A.M. Cunn.                   NA           L12620
Leucopogon muticus R. Br.                       NA             NA
Leucothoe fontanesiana (Steudel) Sleumer        NA          AF124585
Lyonia ferruginea (Walter) Nutt.                NA          AF124584
Lyonia ligustrina (L.) DC.                      NA          AF124578
Lyonia lucida (Lam.) K. Koch                    NA           U82764
Lyonia ovalifolia (Wall) Drude                  NA          AF124580
Lysinema ciliatum R. Br.                        NA             NA
Macleania bullata Yeo                           NA             NA
Macleania rotundifolia Sodiro & Hoerold         NA             NA
Macleania rupestris (Kunth)A. C. Smith          NA             NA
Menziesia ciliicalyx Maxim.                     NA          AF419828
Menziesia pilosa Pers.                          NA           U49293
Monotoca scoparia (W. Smith) R. Br.             NA             NA
Monotropa hypopithys L.                         NA             NA
Needhamiella pumilio (R. Br.) L. Watson         NA             NA
Oligarrhena micrantha R. Br.                    NA             NA
Orthilia secunda (L.) House                     NA          AF419838
Oxydendrum arboreum (L.) DC.                    NA          AF124583
Paphia meiniana (F.Muell.) Schltr.              NA             NA
Pentachondra pumila                          AF419805        L12621
 (Forster & G. Forster) R. Br.
Pernettya tasmanica Hook. f.                    NA           U82765
Phyllodoce caerulea Bab.                        NA          AF419829
Phyllodoce empetriformis D. Don                 NA           U49291
Phyllodoce nipponica Makino                     NA           U49292
Pieris floribunda (Pursh) Benth. & Hook.        NA          AF124577
Pieris formosa (Wallich) D. Don                 NA          AF124581
Pieris nana (Maxim.) Makino                     NA          AF124582
Pieris phillyraeifolia (Hook.) DC.              NA          AF124573
Prionotes cerinthoides (Labill.) R. Br.      AF419806        U79743
Psammisia columnbiensis Hoerold                 NA             NA
Psammisia dolichopoda A. C. Smith               NA             NA
Psammisia sclerantha A. C. Smith                NA             NA
Pyrola picta Smith                              NA             NA
Pyrola rotundifolia L.                       AF419810        L12622
Rhododendron albiflorum Hook.                   NA             NA
Rhododendron canescens (Michaux) Sweet          NA          AF419830
Rhododendron edgeworthii Hook.                  NA             NA
Rhododendron falconeri Hook. f.                 NA             NA
Rhododendron farrerae Tate                      NA             NA
Rhododendron grande Wight                       NA             NA
Rhododendron groenlandicum (Oeder)              NA          AF419831
 Kron & Judd
Rhododendron hippophaeoides Balf.
 f. Forrest                                  AF419807        L01949
Rhododendron hongkongense Hutch.                NA             NA
Rhododendron kaempferi Planchon                 NA          AF419832
Rhododendron kiusiamum Makino                   NA             NA
Rhododendron maximum L.                         NA          AF419833
Rhododendron molle (Blume) G. Don               NA             NA
Rhododendron ovatum (Lindley)                   NA             NA
 Maxim.
Rhododendron pendulum Hook. f.                  NA             NA
Rhododendron semibarbatum Maxim.                NA             NA
Rhododendron stamineum Franchet                 NA             NA
Rhododendron tomentosum (Stokes)                NA           U49286
 Harmaja
Rhododendron lsusiophyllum Sugim.               NA             NA
Rhododendron virgatum Hook. f.                  NA             NA
Rhodothammus chamaecistus Rchb.                 NA           U49287
Richea pandanifolia Hook. f.                    NA             NA
Rupicola sprengeloides Maiden &                 NA             NA
 Betche
Satyria warszewiczii Klotzsch                   NA          AF124579
Sphenotoma dracophylloides Sonder               NA             NA
Sphyrospermum buxifolium Popepping              NA             NA
 & Endl.
Sphyrospermum cordifolium Benth.                NA             NA
Sprengelia incarnata W. Smith                   NA           U80421
Symphysia racemosa (Vahl) Stearn                NA             NA
Tepuia cardonae A. C. Smith                     NA          AF124575
Themistoclesia costaricensis Luteyn             NA             NA
 & Wilbur
Therorhodion camtschaticum (Pallas)
 Small                                          NA          AF419834
Vaccinium cercidifolium J. J. Smith             NA             NA
Vaccinium crenatum
 (D. Don ex Dunal) Sleumer                      NA             NA
Vaccinium macrocarpon Aiton                  AF419808        L12625
Vaccinium meridionale Sw.                       NA          AF124576
Vaccinium myrsinites Lam.                       NA          AF419835
Vaccinium myrtillus L.                          NA          Ay145447
Vaccinium poasamum J. D. Smith                  NA             NA
Vaccinium vitis-idaea L.                        NA          AF419837
Zenobia pulverulenta (Bartram)
 Pollard                                        NA           L12626

                                            GenBank or
                                               EMBL
                                             accession
                                              number
Species                                       matK

Outgroups

Actinidia chinensis Planchon                 U61324
Clethra alnifolia L.                        AF440407
Cyrilla racemiflora L.                      AF440408
Diapensia lapponica L.                      AF440409
Diospyros virginiana L.                     AY145446
Symplocos paniculata Miq.                   AF440433

Ingroup (=Ericaceae)

Agapetes buxifolia Nutt. ex Hook. f.        AF015629
Agapetes hosseana Diels                      U89752
Agapetes scortechinii (King &                U89755
 Gamble) Sleumer
Agapetes serpens (Wight) Sleumer             U89754
Agarista populifolia (Lam.) Judd             U61306
Agarista salicifolia                         U61313
 (Comm. ex Lam.) G. Don
Allotropa virgata Torrey & A.               AF440410
 Gray ex A. Gray
Andersonia sprengelioides R. Br.            AF015631
Andromeda glaucophylia Link                    NA
Andromeda polifolia L.                      AF124569
Arbutus canariensis Duhamel                  U61345
Arbutus menziesii Pursh                        NA
Archeria comberi Melville                   AF015632
Arctostaphylos alpina (L.) Sprengel            NA
Arctostaphylos tomentosa Lindley               NA
Arctostaphylos uva-ursi (L.) Sprengel       AF440411
Astroloma humifusum (Cav.) R. Br.              NA
Bejaria racemosa Vent.                       U61327
Bejaria resinosa Mutis ex L.f.              AF440412
Brachyloma daphnoides (Sm.) Benth.          AF015633
Bryanthus gmelini D. Don                    AF440413
Calluna vulgaris (L.) Hull                   U61326
Cassiope mertensiana (Bong.) G. Don          U61346
Cavendishia complectens Hemsl.               U89762
Cavendishia tarapotana (Meissner) Benth.       NA
 & Hook.
Ceratiola ericoides Michaux                  U61334
Chamaedaphne calyculata (L.) Moench         AF015630
Chimaphila maculata (L.) Pursh              AF440414
Chimaphila menziesii Sprengel               AF440415
Chimaphila umbellata (L.) Barton            AF440416
Comarostaphylis arbutoides Lindley             NA
Corema conradi Torrey ex Loud.              AF440417
Cosmelia rubra R. Br.                       AF015634
Costera endertii J. J. Smith                 U89760
Craibiodendron yunnanense W. Smith           U613707
Cyathodes colensoi Hook.                       NA
Daboecia cantabrica (Hudson) K. Koch         U61349
Dimorphanthera dekockii J. J. Smith          U89756
Diplarche multiflora Hook. f. & Thomson     AF440418
Diplycosia acuminala Becc.                  AF124563
Disterigma trimerum Wilbur & Luteyn         AF001924
Dracophyllum longifolium (J.R. Forster)     AF015635
 Roem. & Schultes
Elliottia bracteata (Maxim.) Hook. f.        U61339
Elliottia racemosa Muhl. ex Elliott            NA
Empetrum atropurpureum Fernald & Wiegand     U61355
Empetrum nigrum L.                             NA
Empetrum rubrum Vahl                         U61342
Enkianthus campanulatus G. Nicholson         U61344
Epacris impressa Labill.                    AF015636
Epigaea repens L.                            U61319
Erica australis L.                           U61329
Erica sicula Guss.                           U61341
Erica spiculifolia Salisb.                   U61337
Erica tetralix L.                            U61340
Eubotrys racemosa L.                        AF124564
Gaultheria eriophylla (Pers.) Sleumer        U61317
Gaultheria miqueliana Takeda                AF124567
Gaultheria shallon Pursh                    AF124565
Gaylussacia frondosa (L.) Torrey & A. Gray     NA
Harrimanella hypnoides Cov.                  U61315
Kalmia angustifolia L.                       U61348
Kalmia buxifolia (Berg) Gift, Kron &         U61347
 Stevens
Kalmia latifolia L.                            NA
Kalmia polifolia Wangenh.                      NA
Kalmia procumbens (L.) Gift,  Kron &         U61352
 Stevens
Kalmiopsis leachiana (Henderson) Rehder      U61323
Lebetanthus myrsinites Macl.                AF539983
Ledothamnus guyanensis Meissner             AF440419
Leucopogon fraseri A.M. Cunn.               AF015637
Leucopogon muticus R. Br.                   AF015638
Leucothoe fontanesiana (Steudel) Sleumer    AF124570
Lyonia ferruginea (Walter) Nutt.             U61312
Lyonia ligustrina (L.) DC.                   U61311
Lyonia lucida (Lam.) K. Koch                 U61308
Lyonia ovalifolia (Wall) Drude               U61305
Lysinema ciliatum R. Br.                    AF015639
Macleania bullata Yeo                        U89758
Macleania rotundifolia Sodiro & Hoerold      U89757
Macleania rupestris (Kunth)A. C. Smith      AF440420
Menziesia ciliicalyx Maxim.                  U61331
Menziesia pilosa Pers.                       U61351
Monotoca scoparia (W. Smith) R. Br.         AF015640
Monotropa hypopithys L.                     AF440421
Needhamiella pumilio (R. Br.) L. Watson     AF539984
Oligarrhena micrantha R. Br.                AF539985
Orthilia secunda (L.) House                 AF440422
Oxydendrum arboreum (L.) DC.                AF124562
Paphia meiniana (F.Muell.) Schltr.           U89748
Pentachondra pumila                         AF015641
 (Forster & G. Forster) R. Br.
Pernettya tasmanica Hook. f.                AF124568
Phyllodoce caerulea Bab.                     U61318
Phyllodoce empetriformis D. Don              U61333
Phyllodoce nipponica Makino                  U61325
Pieris floribunda (Pursh) Benth. & Hook.     U61304
Pieris formosa (Wallich) D. Don              U61303
Pieris nana (Maxim.) Makino                  U61310
Pieris phillyraeifolia (Hook.) DC.           U61309
Prionotes cerinthoides (Labill.) R. Br.     AF015642
Psammisia columnbiensis Hoerold             AF440423
Psammisia dolichopoda A. C. Smith           AF440424
Psammisia sclerantha A. C. Smith            AF440425
Pyrola picta Smith                          AF440426
Pyrola rotundifolia L.                       U61328
Rhododendron albiflorum Hook.                U61358
Rhododendron canescens (Michaux) Sweet         NA
Rhododendron edgeworthii Hook.               U61354
Rhododendron falconeri Hook. f.              U61343
Rhododendron farrerae Tate                  AF440427
Rhododendron grande Wight                    U61336
Rhododendron groenlandicum (Oeder)             NA
 Kron & Judd
Rhododendron hippophaeoides Balf.
 f. Forrest                                  U61353
Rhododendron hongkongense Hutch.             U61338
Rhododendron kaempferi Planchon             AF440428
Rhododendron kiusiamum Makino                U61332
Rhododendron maximum L.                        NA
Rhododendron molle (Blume) G. Don            U61356
Rhododendron ovatum (Lindley)                  NA
 Maxim.
Rhododendron pendulum Hook. f.               U61356
Rhododendron semibarbatum Maxim.             U61330
Rhododendron stamineum Franchet             AF440429
Rhododendron tomentosum (Stokes)            AF440430
 Harmaja
Rhododendron lsusiophyllum Sugim.           AF440431
Rhododendron virgatum Hook. f.               U61335
Rhodothammus chamaecistus Rchb.              U61357
Richea pandanifolia Hook. f.                AF539986
Rupicola sprengeloides Maiden &              AF01543
 Betche
Satyria warszewiczii Klotzsch                U61314
Sphenotoma dracophylloides Sonder           AF015644
Sphyrospermum buxifolium Popepping           U89751
 & Endl.
Sphyrospermum cordifolium Benth.             U89749
Sprengelia incarnata W. Smith               AF015645
Symphysia racemosa (Vahl) Stearn             U89750
Tepuia cardonae A. C. Smith                 AF124566
Themistoclesia costaricensis Luteyn         AF382788
 & Wilbur
Therorhodion camtschaticum (Pallas)
 Small                                       U61322
Vaccinium cercidifolium J. J. Smith          U89753
Vaccinium crenatum
 (D. Don ex Dunal) Sleumer                   U89761
Vaccinium macrocarpon Aiton                  U61316
Vaccinium meridionale Sw.                    U89759
Vaccinium myrsinites Lam.                      NA
Vaccinium myrtillus L.                         NA
Vaccinium poasamum J. D. Smith               U89763
Vaccinium vitis-idaea L.                    AF382819
Zenobia pulverulenta (Bartram)
 Pollard                                    AF124571

NA = sequence not obtained for that species

AA = Arnold Arboretum, Cambridge, Massachusetts

ABG = Atlanta Botanical Garden, Atlanta, Georgia

ACAD = Acadia University, Nova Scotia, Canada

CBG = Canberra Botanical Garden, Australian National University,
Canberra

GH = Gray Herbarium, Harvard University, Cambridge, Massachusetts

NCU = University of North Carolina, Chapel Hill

MICH = University of Michigan Herbarium, Ann Arbor

NYBG = New York Botanical Garden, Bronx

RBGE = Royal Botanic Garden, Edinburgh, Scotland

RBGK = Royal Botanic Garden, Kew, England

S = Swedish Museum of Natural History, Stockholm

UNSW = University of New South Wales, Sydney; Australia;

WFU = Wake Forest University, Winston-Salem, North Carolina
X. Appendix 2: Morphological Data Character numbers correspond to the
list of characters and states in Table I.


                                         1     11111     11112
                              12345  67890     12345     67890

Enkianthus campanulatus       00000  00000     01000     00000
Daboecia cantabrica           00000  00111     00000     01000
Erica spiculifolia            00000  00111     20100     01000
Calluna vulgaris              00000  00211     10100     0100?
Cassiope mertensiana          00000  00211     10100     01010
Erica sicula                  00000  00111     20100     01000
Erica tetralix                00000  00111     20100     01000
Harrimanella hypnoides        00000  00111     00100     01010
Empetrum nigrum               00000  00111     20100     01010
Ceratiola ericoides           00000  00111     20100     01010
Corema conradi                00000  00111     20100     01010
Chimaphila umbellata          01000  0?000     01000     01010
Moneses uniflora              11000  0?100     01000     01010
Orthilia secunda              11000  0?100     01000     01010
Pyrola rotundifolia           11000  0?100     01000     01010
Allotropa virgata             01101  1?1?0     00001     0?010
Hemitomes congestum           11101  1?1?0     00001     0????
Monotropa uniflora            01111  1???0     00001     0?010
Monotropsis odorata           01111  ??1?0     00001     0?010
Pityopus californica          01101  1?1?0     00001     0?010
Pleuricospora fimbriolata     01100  1?1?0     20001     0?010
Pterospora andromeda          01100  1?110     00001     0???0
Sarcodes sanguinea            01100  1?1?0     00001     0?010
Arbutus menziesii             00000  00100     00000     01110
Arctostaphylos uva-ursi       00000  00100     00000     01010
Comarostaphylis diversifolia  00000  00100     00000     01100
Satyria warszewiczii          00000  00100     00001     01100
Vaccinium meridionale         00000  00100     00000     01000
Vaccinium macrocarpon         00000  00100     00000     01000
Bryanthus gmelini             00000  00001     00100     01000
Diplarche multiflora          00000  00110     00000     01010
Elliottia bracteata           00000  0?000     00000     00010
Elliottia racemosa            00000  0?000     00000     00100
Epigaea repens                00000  00001     00000     01000
Kalmia angustifolia           00000  00001     20000     01000
Kalmiopsis leachiana          00000  00000     0{01}000  01000
Ledothamnus guyanensis        00000  00011     10100     01000
Kalmia buxifolia              00000  00110     {01}0010  01000
Kalmia procumbens             00000  00011     10000     01000
Phyllodoce caerulea           000?0  00011     00100     000{01}0
Phyllodoce empetriformis      000?0  00011     00100     00000
Rhodothamnus chamaecistus     000?0  000{01}0  00000     01000
Agarista populifolia          00000  01201     00000     11000
Agarista salicifolia          00000  01001     00000     11000
Andromeda polifolia           00000  00001     00000     01000
Chamaedaphne calyculata       00000  00100     00000     11000
Craibiodendron yunnanense     00000  {01}1100  00000     0110{01}
Diplycosia acuminata          00000  00100     00000     01001
Gaultheria miqueliana         00000  00000     00000     01000
Gaultheria shallon            00000  00000     00000     01000
Leucothoe fontanesiana        00000  00200     00000     01000
Leucothoe racemosa            00000  00100     00000     00000
Lyonia ferruginea             00000  01100     00000     01100
Lyonia lucida                 00000  01100     00000     01100
Oxydendrum arboreum           00000  00200     00000     00000
Pernettya tasmanica           00000  00000     00000     00000
Pieris nana                   00000  01100     20000     00000
Pieris phillyraeifolia        00000  01000     0{01}000  01000
Tepuia cardonae               00000  00100     00000     01000
Zenobia pulverulenta          00000  01000     00000     00000
Bejaria racemosa              00000  00101     00000     01000
Menziesia pilosa              00000  00101     00000     00000
Rhododendron canescens        00000  00101     00000     00100
Rhododendron groenlandicum    00000  00001     00000     01100
Rhododendron minus            00000  00100     00000     01100
Rhododendron kaempferi        00000  00101     00000     01000
Rhododendron maximum          00000  00101     00000     01100
Therorhodion camtschaticum    00000  00101     00000     00000
Rhododendron tsusiophyllum    00000  00101     00000     01000
Sprengelia incarnata          00000  10100     00012     ?1000
Dracophyllum longifolium      10000  10000     00012     ?1000
Epacris impressa              00000  1?10?     00001     ?1000
Lebetanthus myrsinites        00000  0?100     00000     01000
Pentachondra pumila           00000  10100     00002     ?1000
Prionotes cerinthoides        00000  10100     00000     01001
Gaylussacia frondosa          00000  00100     00000     00000
Vaccinium myrsinites          00000  00100     00000     01000
Arbutus canariensis           00000  00100     00000     01110
Arctostaphylos tomentosa      00000  00100     00000     {01}1110
Arctostaphylos alpina         00000  00100     00000     00010


                              22223   22223      33333
                              12345   67890      12345

Enkianthus campanulatus       00000   00001      00000
Daboecia cantabrica           00000   0?001      00000
Erica spiculifolia            00000   10001      0001?
Calluna vulgaris              01???   ?1001      0001{012}
Cassiope mertensiana          01???   ?0011      00002
Erica sicula                  01???   ?0001      00001
Erica tetralix                00000   10001      00011
Harrimanella hypnoides        01???   ?1001      00000
Empetrum nigrum               01???   ?0001      01011
Ceratiola ericoides           01???   ?0001      01011
Corema conradi                01???   ?0001      01010
Chimaphila umbellata          01???   ?0001      00000
Moneses uniflora              01???   ?0001      00000
Orthilia secunda              01???   ?0001      00000
Pyrola rotundifolia           01???   ?0001      00000
Allotropa virgata             01???   ???00      00002
Hemitomes congestum           ?????   ??00?      ?0012
Monotropa uniflora            01???   ???01      0001?
Monotropsis odorata           01???   ???01      10022
Pityopus californica          01???   ???0?      ?0000
Pleuricospora fimbriolata     00000   0??0?      00{01}10
Pterospora andromeda          00000   0?000      ?0000
Sarcodes sanguinea            00000   0??0?      000{02}0
Arbutus menziesii             00000   00000      00002
Arctostaphylos uva-ursi       00000   {012}0000  10002
Comarostaphylis diversifolia  00000   00000      00002
Satyria warszewiczii          00000   00110      00002
Vaccinium meridionale         00000   00110      00002
Vaccinium macrocarpon         00000   00110      00002
Bryanthus gmelini             00000   1000?      ?0012
Diplarche multiflora          00000   0000?      00002
Elliottia bracteata           01???   ?0001      ?0102
Elliottia racemosa            01???   ?0101      ?0112
Epigaea repens                00000   00201      11002
Kalmia angustifolia           00000   00001      10002
Kalmiopsis leachiana          00100   10001      10002
Ledothamnus guyanensis        00000   00011      ?0002
Kalmia buxifolia              00000   10001      10002
Kalmia procumbens             00000   10001      10002
Phyllodoce caerulea           00100   {01}0000   10002
Phyllodoce empetriformis      00100   10000      00002
Rhodothamnus chamaecistus     00000   00000      ?0002
Agarista populifolia          10000   00010      0000{12}
Agarista salicifolia          10000   00010      00002
Andromeda polifolia           11???   ?0000      00002
Chamaedaphne calyculata       00000   00{01}00   10002
Craibiodendron yunnanense     10000   10010      00002
Diplycosia acuminata          00000   00110      00002
Gaultheria miqueliana         00000   00110      00002
Gaultheria shallon            00000   00110      00002
Leucothoe fontanesiana        00000   00110      10002
Leucothoe racemosa            00000   00110      10002
Lyonia ferruguinea            10000   10010      00002
Lyonia lucida                 10000   10010      00002
Oxydendrum arboreum           00000   00101      00002
Pernettya tasmanica           00000   00110      00001
Pieris nana                   10000   100{01}0   10002
Pieris phillyraeifolia        10000   10010      10002
Tepuia cardonae               00000   001{01}1   00002
Zenobia pulverulenta          00000   00010      00002
Bejaria racemosa              00000   01200      00022
Menziesia pilosa              00001   00000      00012
Rhododendron canescens        00000   00000      00102
Rhododendron groenlandicum    00010   {01}0000   00002
Rhododendron minus            00010   10000      00102
Rhododendron kaempferi        00001   00000      00102
Rhododendron maximum          00000   {01}0000   00102
Therorhodion camtschaticum    00000   00000      00102
Rhododendron tsusiophyllum    00001   00000      00102
Sprengelia incarnata          11???   ?1201      10001
Dracophyllum longifolium      11???   ?0101      10002
Epacris impressa              11???   ?0011      10001
Lebetanthus myrsinites        10000   00010      00001
Pentachondra pumila           1????   ?0001      00002
Prionotes cerinthoides        10000   00011      00001
Gaylussacia frondosa          00100   00110      00002
Vaccinium myrsinites          00000   00110      00002
Arbutus canariensis           00000   00000      ?0002
Arctostaphylos tomentosa      00000   {01}000    10002
Arctostaphylos alpina         00000   00001      00000
Matrix for 80 Taxa of Ericaceae.

A list of voucher specimens can be obtained from the first author.


33334     44444     44445     55555     55556     66666  66667  77777
67890     12345     67890     12345     67890     12345  67890  12345

?0000     {12}0100  01000     00000     10000     00000  00000  00000
?0000     01100     01001     00002     2001?     ?00?0  10100  00100
?0001     00101     01100     00002     2011?     ?00?0  10100  00100
10000     00001     01100     00102     20001     000?0  10100  00100
10000     01000     01000     00002     2000{01}  00000  10100  00000
10000     00?00     01100     00102     2001?     ?00?0  10100  00000
10000     00?00     01100     00102     20001     000?0  10100  00000
?0000     00000     01000     00011     20101     00000  10100  00000
?0110     0??11     01200     10002     2011?     ?00?0  10101  00000
?0110     0??11     01200     10002     2011?     ?00?0  10101  00002
?0110     0??11     01200     10002     2011?     ?00?0  10101  00000
?0000     10111     01000     00010     0001?     ?1000  00000  10000
?0000     11111     01000     00002     0001?     ?1000  00100  00000
?0000     10111     01000     00002     1001?     ?0000  01000  00000
?1000     11111     01000     00012     1011?     ?0000  00100  10000
10010     101?1     ?????     00002     1011?     ?0000  1?000  00000
000{01}?  10101     01110     00000     0011?     ?0000  10000  00102
?1010     10010     1??1?     00000     3011?     ?0001  10000  00000
11000     10000     11100     00002     1?11?     ?0000  1?000  00101
?0000     10011     01110     00000     ?011?     ?0001  10000  00102
?0010     10011     01100     00002     0011?     ?0000  00000  00?02
?1000     10000     01000     00002     10001     000?0  00000  00000
?0000     10000     01100     00002     10101     000?0  ?0000  00000
10000     21000     01010     00010     10000     00000  1?100  00010
10000     21000     01010     00010     10000     00000  1?100  00002
10000     21100     01010     00010     10000     00000  1?100  00010
11000     20100     01000     00000     2001?     ?1000  1?100  01000
11000     20100     01000     00000     20001     01000  1?100  01000
11000     20100     01000     00000     2011?     ?1000  10100  01000
11000     11111     01000     00001     2011?     ?00?0  1?100  00200
01000     20101     01000     01001     2011?     ?00?0  1?110  00000
11000     10111     01000     00001     2011?     ?00?0  1?100  10000
11000     ?011?     ?1?00     00001     2011?     ?00?0  1?110  10000
11100     {12}1001  01010     00000     2011?     ?00?0  1?110  10000
11000     11001     0101{01}  00000     2011?     ?00?0  1?100  00000
11000     10001     01011     00000     2011?     ?00?0  1?110  000?0
1?000     {12}1111  01000     1000?     2011?     ?00?0  1?100  00210
11000     11001     01000     10001     2011?     ?00?0  11100  001?1
10001     11001     01000     10001     2011?     ?00?0  11110  00101
11000     11000     01001     00000     2011?     ?00?0  10100  00000
11?00     11000     01000     0000{01}  2001?     ?00?0  1?100  00000
11?00     11001     01000     00000     ?011?     ?00?0  10110  00000
11000     20100     01000     00100     2001?     ?0100  10100  00000
11000     20100     01000     00100     2001?     ?0100  10100  00000
11000     20000     01010     00000     20000     00000  10100  00000
01000     20100     01000     00002     2001?     ?1000  10100  00000
11000     20100     01000     00101     20{01}1?  ?0100  10100  00000
01000     30100     01000     00002     2001?     ?10?0  10100  00000
11000     30100     01000     00001     20000     10100  10100  00000
11100     30000     01011     00000     20000     10100  10100  00000
11000     20100     01000     00001     20000     10100  10100  00000
11000     20100     01000     00002     20000     00100  10100  00000
11000     20100     01001     001{01}1  20002     00110  10100  00000
11001     20100     01001     00101     20002     00110  10100  00000
10000     20100     01000     00000     2001?     ?0100  10100  00000
11000     20100     01010     00000     20000     10000  10100  00000
11001     20100     01000     00001     20001     00100  10100  00000
11001     20100     01000     0?102     20001     00100  10100  00000
01000     20100     01010     00000     2001?     ?1000  10100  00000
11000     20100     01000     00011     20000     10100  10100  00000
10000     20111     01000     00000     2011?     ?0000  10100  00000
10100     20101     01010     0000{01}  2011?     ?0000  00110  00000
10100     20?01     01011     10000     2011?     ?0000  10100  00000
10100     20101     01000     00000     2011?     ?0000  10110  00000
10100     20101     00001     00000     2011?     ?0000  10110  00000
10100     20101     00000     10000     2011?     ?0000  10110  00000
10100     20101     00011     00000     2011?     ?0000  10110  00000
10000     01101     00010     00000     2011?     ?0000  1?110  00000
10100     20101     01001     10000     2011?     ?0000  10110  00001
00000     20101     01100     11102     2101?     ?0000  1?100  00000
10100     20101     01100     11002     ?111?     ?0000  1?100  00000
00100     20000     01100     11002     2101?     ?0000  1?100  000?0
10100     20101     01100     10002     2111?     ?0000  1?100  00000
00100     21101     01110     11002     2111?     ?0000  1?100  00100
10100     20100     01100     10002     2111?     ?0000  1?100  00010
11000     20000     00000     00002     2001?     ?1000  1?100  01000
11000     20100     00000     00000     2001?     ?1000  1?100  01000
10000     21000     00010     00010     10100     00000  1?100  00010
10{01}00  21000     00010     00010     10{01}00  00000  1?100  00100
?1{01}00  21000     00010     00010     10000     00000  1?100  00100


33334     77778     88888  88889     9
67890     67890     12345  67890     1

?0000     00000     00000  00000     0
?0000     00010     00000  00010     0
?0001     00000     00100  00040     0
10000     00010     00100  00030     0
10000     00000     00100  00020     0
10000     00000     00100  ??010     0
10000     00000     00100  00010     0
?0000     00000     00100  01030     0
?0110     O1l??     11100  01020     ?
?0110     O1l??     11100  ?1020     ?
?0110     011??     10100  ?1020     ?
?0000     00000     00101  ??020     0
?0000     10010     00101  ??020     0
?0000     00000     00101  ??050     1
?1000     00000     00101  ??060     1
10010     10000     00102  ??020     0
000{01}?  201??     0?102  ??0??     ?
?1010     10000     00102  ??0?0     0
11000     201??     00102  ??0?0     ?
?0000     201??     0?102  ??0?0     ?
?0010     201??     0?102  ??020     ?
?1000     10000     00111  ??030     0
?0000     10020     00001  ??030     0
10000     001?0     11100  1?020     ?
10000     011?0     11100  10020     ?
10000     011?0     11100  11020     ?
11000     001?0     01100  0?010     ?
11000     001?0     01100  0?011     ?
11000     001?0     01100  ??010     ?
11000     00010     00100  ??0?0     0
01000     00010     00100  ??0?0     0
11000     00010     0011?  ??000     0
11000     00010     0011?  ??000     0
11100     00010     0010?  ??0?0     0
11000     00010     0011?  ??010     0
11000     000{01}0  00100  ??01?     0
1?000     00010     00100  ??010     0
11000     000{01}0  0000?  ??010     0
10001     000{01}0  00000  ??010     0
11000     00010     00100  ??010     0
11?00     0001?     00100  ??010     0
11?00     000{01}0  00100  ??010     0
11000     00000     00100  0?010     0
11000     00000     00000  0?0?0     0
11000     00000     00100  00010     0
01000     00000     00100  01000     0
11000     00000     00100  ??0?0     0
01000     00000     00100  ??140     0
11000     00000     00100  00100     0
11100     00000     00100  00000     0
11000     00000     00100  00000     0
11000     00000     00100  0?000     0
11000     00001     00110  00010     0
11001     00001     00100  00010     0
10000     00000     00000  01010     0
11000     001??     01100  ??100     ?
11001     00000     00100  000?0     0
11001     00000     00100  0?0?0     0
01000     001??     01100  ??1?0     ?
11000     00000     00100  ?0000     0
10000     00010     00100  000?0     0
10100     00010     00000  01020     0
10100     00010     00010  00020     0
10100     00010     00010  01020     0
10100     00010     00000  0{01}020  0
10100     00010     00000  01020     0
10100     00010     00010  01020     0
10000     00010     00000  01010     0
10100     00010     00100  0?020     0
00000     00000     0000?  ??010     0
10100     00000     0010?  ??020     0
00100     00000     0010?  ??020     0
10100     00000     0000?  ??0?0     0
00100     O11??     2110?  ??0?0     ?
10100     00000     ?000?  ??0?0     0
11000     011?0     21100  00011     ?
11000     001?0     01100  00011     ?
10000     {01}01?0  11100  10020     ?
10{01}00  011?0     20100  1?0?0     ?
?1{01}00  01??0     21100  1?020     ?


XI. Appendix 3: Formal Descriptions of Ericaceae, with Latin Diagnoses.

Characters in boldface type are likely synapomorphic.

ERICACEAE

Ericaceae Durande, Notions Elem. Bot. 270. 1782 (as Ericae), nom. cons.

Andromedaceae DC. ex Schnizl., Iconogr. Fam. Regni Veg. 2:[iv], ad t. 161 c. 1834-1870.

Arbutaceae Bromhead, Mag. Nat. Hist., n.s., 4: 337, 338. 1840 (as Arbuteae).

Arctostaphylaceae J. Agardh, Theoria Syst. Pl.: 106. 1858 (as Arctostaphyleae).

Azaleaceae Vest, Anleit. Stud. Bot.: 272, 294. 1818 (as Azaleoideae).

Diplarchaceae Klotzsch, Monatsber. Konigl. Preuss. Akad. Wiss. Berlin 1859.: 15. 1860.

Empetraceae Bercht. & J. Presl, Prir. Rostlin: 251. 1820 (as Empetreae), nom. cons.

Epacridaceae R. Br., Prodr.: 535. 1810, nom. cons.

Hypopityaceae Link, Handbuch 2: 403. 1831 (as Hypopithydes).

Kalmiaceae Durande, Notions Elem Bot.: 271. 1782 (as Kalmiae).

Ledaceae J. F. Gmel., Aug. Gesch. Pflanzengifte, ed. 2: 404. 1803.

Menziesiaceae Klotzsch, Linnaea 24: 11. 1851.

Monotropaceae Nutt., Gen. N. Amer. Pl. 1: 272, 1818 (as Monotropeae).

Oxycoccaceae A. Kern., Pflanzenleben 2: 713, 714. 1891.

Prionotaceae Hutch., Evol. Phylog. FL. Pl.: 306. 1969.

Pyrolaceae Lindl., Syn. Brit. Fl.: 175. 1829 (as Pyroleae), nom. cons.

Rhododendraceae Juss., Gen. Pl.: 158. 1789 (as Rhododendra).

Rhodoraceae Vent., Tabl. Regne Veg. 2: 449. 1799.

Salaxidacene J. Agardh, Theoria Syst. Pl.: 104. 1858 (as Salaxidene).

Stypheliaceae Horan., Prim. Lin. Syst. Nat.: 72. 1834.

Vacciniaceae Adans., Fam. Pl. 2: 160. 1763 (as Vaccinia), nom. cons.

Evergreen or deciduous shrubs, lianas, trees, or herbs (and these sometimes lacking chlorophyll), sometimes epiphytic. Leaves alternate, decussate, or whorled, sometimes distally clustered on shoots, entire to serrate, and often with margins strongly revolute and leaves needle-like (i.e., ericoid); stipules lacking; i.e., leaf epidermal cells lignified or not, and hypodermal cells often present; stomata usually anomocytic or paracytic; vascular bundles frequently associated with a fiber sheath; nodes usually unilacunar. Buds protected by 2 to numerous scales or naked. Indumentum a mixture of unicellular and multicellular hairs, or sometimes only unicellular-pubescent, the multicellular hairs extremely various, glandular or non-glandular, unbranched and short to elongate, variously branched, or peltate. Inflorescences terminal or axillary, usually indeterminate and often racemose, with variously developed bracts and usually a pair of bracteoles, these sometimes recaulescent, replacing calyx lobes; flowers pend ulous to erectly held, articulated with pedicel or not. Flowers showy to inconspicuous, perfect to imperfect, usually 4- or 5-merous, actinomorphic to zygomorphic. Sepals (1-)4-5(-9), usually connate, persistent or deciduous, occasionally colorful and fleshy; petals (3-)4-5(-9), usually connate and typically campanulate, cylindrical, or urceolate, occasionally rotate, deciduous or persistent. Stamens (2-)5-10(-16), free from the corolla or adnate, sometimes connate, included or exserted, the filaments straight to variously curved, unicellular-pubescent or glabrous; anthers tetrasporangiate or bisporangiate, inverting (late, or more commonly early) in development, with 2 or 4 apparently terminal or dorsal appendages (awns, spurs) or these sometimes on the filaments, opening by pores or short to long slits, sometimes through narrowed tubules, an endothecium usually lacking. Pollen usually tricolpate to tricolporate, in tetrahedral tetrads, but these sometimes reduced, or occasionally in monads, sometimes associ ated with viscin threads. Nectary present or absent. Ovary (2-)4-5(-14) carpellate, superior to inferior, with usually axile to intruded parietal placentation, rarely apical or basal; ovules 1 to many per carpel, anatropous to nearly campylotropous, unitegmic, tenuinucellate, the embryo sac usually of the Polygonum type; style short to long, hollow, sometimes expanded apically; stigma truncate to capitate or slightly to strongly lobed, sometimes cup shaped, funnel shaped, or flabellate to pinnatifid. Fruit loculicidal to septicidal capsules, berries, drupes (with 1 or several pits), or occasionally dry and indehiscent; seeds small, testa usually single layered, with cells isodiametric to strongly elongated, sometimes winged or tailed, sometimes mucilaginous on wetting; embryo fusiform or spatulate, white or less commonly green, with 2 cotyledons, to extremely reduced and undifferentiated; endosperm cellular, well developed, with haustoria at both ends; germination epigeal.

ENKIANTHOIDEAE

Enkianthoideae Kron, Judd & Anderb., stat. nov. Type genus: Enkianthus Lour. Enkiantheae Stevens, Bot. J. Linn. Soc. 64: 35. 1971.

Shrubs. Leaves evergreen or deciduous, alternate or pseudoverticillate, flat, non-ericoid, serrate or entire, convolute in bud. Leaf epidermal cells not lignified. Indumentum of distinctive, elongate, non-glandular hairs. Winter buds perulate. Inflorescence racemose or umbellate; bracteoles lacking; calyx not articulated with pedicel. Flowers 5-merous, actinomorphic. Calyx lobes deciduous; corolla sympetalous, campanulate, tubular or urceolate. Stamens 10, included, the filaments [+ or -] straight, unicellular pubescent; anthers tetrasporangiate, inverted late in development, papillose or smooth, with a pair of awns, with fibrous endothecium, opening with longitudinal slits. Pollen in monads, without viscin threads. Ovary 5-locular, with axile placentation, superior; style long, impressed; stigma not expanded. Fruit a loculicidal capsule; seeds thin walled with a few layered testa, with winglike ridges, or smooth, with vascular bundle in raphe; embryo sac on nucellar pedestal with earlike projections; embryo with 2 cotyledons. [Enkianthus Lour., Fig. 33)

MONOTROPOIDEAE

Monotropoideae Arn., Encycl. Brit., ed. 7,5:118. 1832 (as Monotropeae). Type genus: Monotropa L.

Pyroloideae Kostel., Allg. Med.-Pharm. Fl. 3: 1026. Apr.-Dec. 1834 (as Pyroleae).

Evergreen herbs and subshrubs, or echlorophyllous, mycotrophic herbs. Leaves alternate, spirally arranged, non-ericoid, and convolute in bud. Pith homogeneous. Leaf epidermal cells not lignified, fibers not associated with midrib. Indumentum of unicellular or multicellular hairs, or none. Inflorescence terminal, racemose or solitary, bracts present, bracteoles absent, calyx articulated or continuous with the pedicel. Flowers (4-)5(-6)-merous, actinomorphic. Calyx and corolla gamopetalous or polypetalous and sometimes not well differentiated, rotate, campanulate, urceolate, or tubular. Stamens 10, anthers inverting usually just before anthesis, with or without appendages at the anther-filament junction, endothecium present but sometimes poorly developed. Pollen shed in monads, tetrads, and/or polyads. Ovary (4-)5(-8), with axile to parietal placentation, with many ovules per locule, superior, style usually impressed, stigma sometimes expanded and lobed. Fruit a loculicidal capsule, berry, or irregularly dehisc ent, seeds with or without tails, testa cells slightly to moderately elongated and unthickened, or the cells isodiametric and moderately to massively thickened on the inner surface; embryo small, with 2 cotyledons, or minute and not differentiated.

PYROLEAE

Pyroleae Dumort., Anal. Fam. Pl. 47: 1829. Type genus: Pyrola L.

Herbs or subshrubs, evergreen, with spiral, serrate, non-ericoid leaves, convolute in bud. Pith homogeneous. Leaf epidermal cells not lignified, fibers not associated with midrib. Indumentum of unicellular hairs, or none. Inflorescence terminal, racemose, bracts present, bracteoles none; calyx not articulated with pedicel. Flowers 5-merous, actinomorphic, the calyx lobes rather small, [+ or -] persistent and the corolla polypetalous, rotate, glabrous. Stamens 10, the filaments [+ or -] dilated basally, straight, usually smooth; anther dehiscing by terminal pores, inverting just before anthesis, appendages none, surface usually smooth, endothecium present, resorbtion tissue present or absent. Pollen in monads, tetrads, or polyads. Ovary 5-locular, with axile placentation and many ovules per locule, superior; style impressed, short to quite long; stigma usually peltate, sometimes also lobed. Fruit a loculicidal capsule; seeds with tails at both ends, cells moderately elongated and unthickened; embryo small, wit h 2 cotyledons; n = 13, 19, 23. (Chimaphila Pursh, Fig. 34; Moneses Salisb., Fig. 35; Orthilia Raf., Fig. 36; Pyrola L., Fig. 37]

MONOTROPEAE

Monotropeae Dumort., Anal. Fam. Pl. 47: 1829. Type genus: Monotropa L. Pleuricosporeae A. Gray, Proc. Amer. Acad. Arts 7: 370. 1868.

Echlorophyllous mycotrophic herbs usually blackening on drying, stems annual, with spiral, entire, sessile, non-differentiated leaves convolute in bud. Pith homogeneous. Leaf epidermal cells not lignified, fibers not associated with midrib. Indumentum none, rarely with multicellular, obscurely glandular, hairs. Inflorescence terminal, racemose or flower single, terminal; bracts present, bracteoles none, basal or apical; calyx articulated with pedicel or not. Flowers (4-)5(-6)-merous, actinomorphic. Calyx lobes small, [+-] persistent; corolla sympetalous or polypetalous, campanulate or tubular, or perianth uniseriate, [+-] tubular, parts free, basally saccate, glabrous or with hairs adaxially. Stamens 10, the filaments not dilated basally, straight, glabrous or with short hairs; anther dehiscing by terminal pores or [+-] elongated slits, inverting just before anthesis, appendages uncommon, small, smooth, paired, at anther--filament junction, or anthers hippocrepiform, dehiscing by slits, appendages none, surfa ce usually smooth, endothecium at most poorly developed, pollen monadinous. Ovary (4-)5(-8)-locular, with axile to parietal placentation and many ovules per locule, superior; style usually impressed, little longer than ovary; stigma barely expanded, rarely lobed. Fruit a loculicidal capsule, berry or irregularly dehiscent; seeds with isodiametric cells moderately to massively thickened on inner surfaces, or tailed, with moderately elongated and unthickened cells; embryo minute, undifferentiated, with <30 cells; n = 8, 13, 26, 32 (x = 8, 13?). [Allotropa A. Gray, Fig. 38; Cheilotheca Hook. f.; Hemitomes A. Gray; Hypopitys Crantz; Monotropa L., Fig. 39; Monotropastrum Andres; Monotropsis Schwein.; Pityopus Small; Pleuricospora A. Gray] Note: Interpretation of the morphological nature of the parts of the flower in genera like Monotropa is difficult.

PTEROSPOREAE

Pterosporeae Baillon, Hist. Pl. 11: 161, 206. 1891. Type genus: Pterospora Nutt.

Echlorophyllous mycotrophic herbs, stems annual, with spiral entire sessile non-differentiated leaves convolute in bud. Pith homogeneous. Leaf epidermal cells not lignified, fibers not associated with midrib. Indumentum of multiseriate-stalked glandular hairs. Inflorescence terminal, racemose; bracts present, bracteoles none; calyx articulated with pedicel. Flowers 5-merous, actinomorphic. Calyx lobes, [+-] persistent; corolla sympetalous, urceolate, glabrous. Stamens 10, the filaments not dilated basally, straight, smooth; anther dehiscing by slits widening considerably apically, inverting some time before anthesis, appendages long, strongly papillate, paired, at anther-filament junction, surface papillate, endothecium present, pollen monadinous. Ovary 5-locular, with axile and partly intruded parietal placentation, many ovules per locule, superior; style impressed; stigma barely expanded. Fruit a loculicidal capsule; seeds with a tail at one end, cells slightly elongated and unthickened; embryo minute, undi fferentiated, with 30-40 cells; n = 8. [Pterospora Nutt.; Sarcodes Torrey, Fig. 40]

ARBUTOIDEAE

Arbutoideae Nied., Bot. Jahrb. Syst. 11: 135. 1889. Type genus: Arbutus L.

Arbuteae Meisn., Pl. Vasc. Gen.: Tab. Diagn. 243, Comm. 154. 1839.

Trees or shrubs, usually evergreen. Leaves usually alternate, non-ericoid, entire or serrate, convolute in bud. Leaf epidermal cells not lignified. Indumentum of stalked glandular hairs and sometimes elongate, non-glandular hairs. Inflorescence terminal, racemose or paniculate; bracts present, bracteoles paired, [+-] basal; calyx not articulated with pedicel. Flowers (4-)5-merous, actinomorphic. Calyx lobes small, persistent; corolla sympetalous, urceolate, with long unicellular hairs on the inside, the lobes small. Stamens (8-)l0, included, the filaments [+ or -] strongly dilated basally, [+ or -] straight, unicellular-pubescent or smooth; anthers tetrasporangiate, inverting only just before anthesis, slightly roughened, with a pair of dorso-apical spurs, lacking a fibrous endothecium, lacking disintegration tissue, dehiscing by terminal pores or slitlike pores. Pollen in tetrads, without viscin threads. Ovary 4-10-locular, with axile placentation and only 1 to few ovules per locule, superior, sometimes with fleshy papillae; style continuous with the ovary; stigma truncate. Fruit [+ or -] fleshy with bony or fibrous endocarp; i.e., a drupe or drupe-like berry; seeds with testa cells [+ or -] elongated; embryo with 2 cotyledons. [Arbutus L., Figs. 41-42; Arctostaphylos Adans., Figs. 43-45; Comarostaphylos Zucc., Fig. 46; Ornithostaphylos Small, Fig. 47]

CASSIOPOIDEAE

Cassiopoideae Kron & Judd, stat. nov. Type genus: Cassiope D. Don.

Cassiopeae Stevens, Bot. J. Linn. Soc. 64: 37. 1971.

Low shrubs, stem lacking pericyclic sheath of fibers, with Calluna-type pith. Leaves evergreen, decussate, small, ericoid, auriculate, revolute in bud. Leaf epidermal cells not lignified. Indumentum of fasciculate branched and small glandular hairs. Inflorescence axillary, one-flowered, the flower with 4-6 basal bracteoles; calyx articulated with pedicel. Flowers (4-)5-merous, actinomorphic. Calyx lobes [+ or -] moderate sized, [+ or -] deciduous; corolla sympetalous, [+ or -] campanulate, the lobes moderately large. Stamens (8-)10, included, the filaments [+ or -] straight, unicellular-pubescent or smooth; anthers inverting early in development, roughened, with a pair of dorso-apical spurs at (or just above) junction with filament, lacking a fibrous endothecium, with or without a white deposit of disintegration tissue dorsally, dehiscing by terminal, slitlike pores. Pollen in tetrads, without viscin threads. Ovary (4-)5-locular, with axile placentation, superior; style impressed; stigma truncate. Fruit a loc ulicidal capsule; seeds with elongate testa cells; embryo with 2 cotyledons. [Cassiope D. Don, Fig. 48]

ERICOIDEAE

Ericoideae Link, Handbuch 1: 602. 1829 (as Ericeae). Type genus: Erica L.

Empetroideae Nutt. ex Sweet, Hort. Brit.: 491. 1826 (as Empetreae).

Ledoideae Alef., Linnaea 28: 3, 7. 1856 (as Ledeae).

Rhododendroideae (Juss.) Sweet, Brit. Fl. Gard. 3: 263. 1828 (as Rhodoraceae).

Rhodoroideae Kostel., Allg. Med.-Pharm. Fl. 3: 1015. 1834 (as Rhodoraceae).

Shrubs to trees. Leaves evergreen or deciduous, alternate, decussate, 3- or 4(6)-whorled or pseudowhorled, ericoid or non-ericoid, revolute or convolute in bud, margin entire to serrulate or serrulate-ciliate. Leaf epidermal cells not lignified. Indumentum of unbranched, variously branched, peltate, or scalelike, glandular to non-glandular hairs, and sometimes papillae. Inflorescence terminal or axillary, racemose, spicate, corymbose, umbellate, fasciculate, capitate, or of solitary flowers; bracts and bracteoles extremely various; flower articulated or continuous with pedicel. Flowers usually 4-7(-9)-merous, actinomorphic or slightly zygomorphic, sometimes variably reduced and wind pollinated, with a distinct calyx and corolla or perianth parts [+ or -] undifferentiated, pendulous to erectly held. Calyx lobes (or sepals) usually 1-7, or lacking, variable in size, persistent or deciduous; corolla choripetalous or more commonly sympetalous, and variously shaped, persistent or deciduous, with the usually 1-7 lo bes small to large. Stamens 2-15, included or sometimes exserted, the filaments straight to curved or S-shaped, distinct or connate, unicellular-pubescent or glabrous (and smooth); anthers tetrasporangiate, inverting early in development, variably roughened or smooth, with or without a pair of dorsal appendages (spurs), lacking disintegration tissue, usually lacking a fibrous endothecium, dehiscing by terminal pores or longitudinal slits. Pollen in tetrads, with viscin threads present or absent. Ovary 1-14-locular, with axile placentation, superior; style gradually tapered or impressed; stigma truncate, capitate, peltate, cup shaped, funnel shaped, or strongly lobed, considerably enlarged in wind pollinated species. Fruit a septicidal to loculicidal capsule, indehiscent pod, or drupe; seeds with variable testa cells; embryo with 2 cotyledons.

ERICEAE

Ericeae DC. ex Duby, Bot. Gall. 1:316. 1829 (as Ericaceae). Type genus: Erica L.

Calluneae Klotzsch, Monatsber. Konigl. Preuss. Akad. Wiss. Berlin 1857: 3. 1857.

Daboecieae Cox, Amer. Midl. Naturalist 39: 238. 1948.

Eleutherostemoneae Klotzsch, Linnaea 12: 217. 1838 (as Eleutherostemones).

Eleutherostemoninae Meisn., Pl. Vasc. Gen.: Tab. Diago. 244. 1839 (as Eleutherostemones).

Ericinae D. Don, Edinburgh New Philos. J. 17: 152. 1834 (as Ericeae normales).

Salaxideae Drude in Engl. & Prantl, Nat. Pflanzenfam. IV, 1: 32. 1889.

Salaxidinae Engl., Gen. Pl.: 751. 1839 (as Salaxideae).

Shrubs to trees. Leaves evergreen, 3- or 4(6)-whorled, rarely decussate or alternate, ericoid, revolute in bud. Leaf epidermal cells not lignified. Indumentum of unbranched or branched, glandular to non-glandular hairs. Inflorescence racemose to umbellate, at end of branch; bract always present, on the main axis or partially recaulescent to fully recaulescent forming the abaxial lobe of calyx; bracteoles lacking, solitary, or paired, usually fully recaulescent as the lateral lobes of calyx, rarely more than 2. Flowers usually 4-merous, actinomorphic. Calyx lobes l-4(-5), or lacking, variable in size, persistent; corolla sympetalous, urceolate, cylindric, or campanulate, the lobes [+ or -] small, usually persistent. Stamens (3-)8(-l0), sometimes exserted, the filaments straight to S-shaped, distinct or connate, unicellular-pubescent or smooth; anthers variably roughened, with or without a pair of dorsal appendages, dehiscing by terminal pores. Pollen without viscin threads. Ovary (l-)4(-8)-locular, with axile placentation, superior; style gradually tapered or impressed; stigma truncate, capitate, peltate, cup shaped, funnel shaped, or strongly 4-lobed, considerably enlarged in wind pollinated species. Fruit a loculicidal capsule, indehiscent pod, or drupe; seeds with variable testa cells; embryo with 2 cotyledons. [Calluna Salisb., Fig. 49; Daboecia D. Don, Fig. 50; Erica L., Figs. 51-53]

EMPETREAE

Empetreae D. Don, Edinburgh New Philos. J. 1827: 59. 1827. Type genus: Empetrum L. Coremateae Pax in Engl. & Prantl, Nat. Pflanzenfam. III, 5:126. 1891.

Shrubs, or shrublets, sometimes mat forming. Leaves evergreen, alternate or almost verticiliate, ericoid, revolute in bud. Leaf epidermal cells not lignified. Inflorescence axillary, with solitary flowers, few-flowered clusters, or in terminal heads; bracts and bracteoles various; perianth not articulated with the pedicel. Flowers actinomorphic, reduced, choripetalous, wind pollinated, unisexual or bisexual. Sepals and petals 3-6, inconspicuous, more or less undifferentiated. Stamens 2 or 3, exserted, the filaments straight and smooth, anthers [+ or -] smooth, lacking appendages, without fibrous endothecium, dehiscing by longitudinal slits. Pollen without viscin threads. Ovary 2 to many locular, with axile placentation, superior, style short, impressed, stigma lobes deeply divided. Fruit a dry or fleshy drupe with 2 to many pyrenes; seeds with [+ or -] isodiametric testa cells; embryo with 2 cotyledons. [Ceratiola Michx;, Figs. 54-55; Corema D. Don, Empetrum L., Fig. 56]

BEJARIEAE

Bejarieae Copeland, Amer. Midl. Naturalist 30: 547. 1943. Type genus: Bejaria Mutis ex L.

Low shrubs. Leaves evergreen, alternate or 3-4 whorled or decussate, ericoid to flat, revolute in bud. Leaf epidermal cells not lignified. Indumentum of glandular and elongate, or nonglandular hairs. Inflorescence terminal or axillary, racemose to paniculate, or a solitary flower; bracts present, bracteoles paired, [+ or -] basal to just below calyx; calyx articulated or continuous with pedicel. Flowers 4-7(-9)-merous, actinomorphic. Calyx lobes variable in size, persistent or withering-persistent; corolla [+ or -] choripetalous, rotate, campanulate, salverform, fusiform or infundibular. Stamens (5-)6-14(-l8), the filaments straight, smooth pubescent; anthers smooth or roughened, lacking appendages, dehiscing by short to elongate slits. Pollen with viscin threads. Ovary 4-7-locular, with axile placentation, superior; style articulated with the ovary; stigma truncate to capitate. Fruit a septicidal capsule, warty in Ledothamnus, seeds with [+ or -] isodiametric to elongate testa cells; embryo with 2 cotyledons . [Bejaria Mutis ex L., Fig. 57; Bryanthus D. Don; Ledothamnus Meisn., Fig. 58]

PHYLLODOCEAE

Phyllodeceae Drude, Engl. & Prantl, Nat. Pflanzenfam. 4, 1: 31, 38. 1889. Type genus: Phyllodoce Salisb.

Cladothamneae Copeland, Amer. Midl. Naturalist 30: 548. 1943.

Epigaeae Britton & Brown, Illus. Fl. N. U. S., ed. 2, 2: 676. 1913.

Leiophylleae Dippel, Handb. Laubholzk. 1: 435. 1889.

Shrubs. Leaves evergreen, alternate and spiral, decussate, or whorled, ericoid or non-ericoid, entire to serrulate, convolute or revolute in bud. Leaf epidermal cells not lignified.

Indumentum of glandular hairs and elongate non-glandular hairs, and sometimes papillae. Inflorescence terminal, rarely axillary, spicate, racemose, or corymbose, sometimes reduced; bracts paired and [+ or -] basal, bracteoles usually present; calyx articulated with pedicel. Flowers 4-6-merous, actinomorphic. Calyx lobes variable in size, withering persistent or persistent; corolla sympetalous or choripetalous, rotate to urceolate, campanulate, the lobes small to large. Stamens 5-10, included or exserted, the filaments straight to curved, unicellular-pubescent or smooth; anthers smooth or roughened, lacking appendages, dehiscing by slits or terminal pores. Pollen usually with viscin threads. Ovary 2-6-locular, with axile placentation; style impressed or not; stigma truncate, sometimes slightly expanded. Fruit a septicidal capsule, sometimes partly loculicidal as well; seeds with testa cells [+ or -] isodiametric to elongated; embryo with 2 cotyledons. [Elliottia Muhl., Fig. 59; Epigaea L., Kalmia L., including Loiseleuria L. and Leiophyllum (Pers.) Hedwig f. (Kron & King, 1996; Gift & Stevens, 1997); Kalmia procumbens (L.) Gift, Kron, & Stevens, comb. nov.; Kalmia buxifolia (Bergius) Gift, Kron, & Stevens, comb, nov.; Figs. 60-61; Kalmiopsis Rehder, Fig. 62; Phyllodoce Salisb., Fig. 63; Rhodothamnus Reichb., Fig. 64]

RHODOREAE

Rhodoreae DC. ex Duby, Bot. Gall. 1: 318. 1828 (as Rhodoraceae). Type genus Rhodora L. (= Rhododendron L.)

Diplarcheae Airy Shaw, Kew Bull. 17: 509. 1964.

Ledeae Rchb. f. in Rchb., Icon. Fl. Germ. Helv. 17: 78. 1854-1855.

Ledinae Benth. in DC., Prodr. 7: 729. 1839 (as Ledeae).

Menziesieae K. Koch, Hort. Dendrol. 2: 340. 1854 (as Menziesiaceae).

Rhododendrinae Benth. in DC., Prodr. 7: 712. 1839 (as Rhododendreae).

Shrubs or small trees. Leaves evergreen or deciduous, alternate, non-ericoid, entire to serrulate-ciliate, convolute or revolute in bud. Leaf epidermal cells not lignified. Indumentum very variable, sometimes glandular, peltate, or highly branched. Inflorescence terminal or rarely axillary, usually corymbose or umbellate; bracts present, usually as perulae (except Therorhodion where bracts are green), bracteoles paired, usually [+ or -] basal; calyx articulated with pedicel or not. Flowers usually 4- or 5-merous, slightly zygomorphic, but occasionally secondarily actinomorphic. Calyx lobes variably developed, persistent or deciduous; corolla usually sympetalous, variously shaped, often with spots or blotches, the lobes usually moderate to large; stamens 5-15, included or exserted, the filaments straight to curved, unicellular-pubescent or smooth; anthers smooth, lacking spurs, dehiscing by terminal pores to elongate slits. Pollen with viscin threads. Ovary 3-14-locular, with axile placentation, superior; styl e impressed or not; stigma truncate or expanded. Fruit an ovoid to cylindric septicidal capsule; seeds with testa cells usually elongated; embryo with 2 cotyledons. [Diplarche Hook. f.; Menziesia Sm., Fig. 65; Rhododendron L., Figs. 66-68; Therorhodion Small, Fig. 69]

HARRIMANELLOIDEAE

Harrimanelloideae Kron & Judd, subfam. nov. Type genus: Harrimanella Cov.

Latin diagnosis: Harrimanelloideae foliis alternis et ericoidiis, pilis solum unicellularis, floribus 5-mero, solitariis et terminatibus, bracteis et bracteolis nullis, stylo brevi et crasso differt.

Mat-forming shrubs, stem lacking pericyclic sheath of fibers, with homogeneous pith. Leaves evergreen, alternate, small, acicular, ericoid-like, petiolate, revolute in bud. Leaf epidermal cells not lignified. Indumentum of only unicellular hairs. Inflorescence terminal, flowers solitary; bracts and bracteoles lacking; calyx articulated with pedicel. Flowers 5-merous, actinomorphic. Calyx lobes moderate sized, [+ or -] deciduous; corolla sympetalous, [+ or -] campanulate, the lobes large. Stamens 10, included, the filaments [+ or -] straight, smooth; anthers tetrasporangiate, inverting early in development, [+ or -] smooth, with a pair of dorso-apical spurs at junction with filament, lacking a fibrous endothecium, without disintegration tissue dorsally, dehiscing by terminal, slitlike pores. Pollen in tetrads, without viscin threads. Ovary 5-locular, with axile placentation, superior; style slightly impressed, short and stout; stigma truncate. Fruit a loculicidal capsule; seeds with moderately elongate testa c ells; embryo with 2 cotyledons. [Harrimanella Cov.; Fig. 70]

STYPHELIOIDEAE

Styphelioideae Sweet, Fl. Australas.: ad t. 47. 1828 (as Stypheline).

Epacridoideae Link, Handbuch 1: 601. 1829 (as Epacrideae). Type genus: Styphelia R. Br.

Shrubs, small trees or climbers. Leaves evergreen, alternate, usually entire; veins often [+ or -] palmate, parallel, or reduced to one. Leaf epidermal cells lignified; bracts and bracteoles scarious. Inflorescence variable, axillary or terminal. Flowers 4-5-merous, actinomorphic. Calyx persistent; corolla persistent, sympetalous, usually tubular or campanulate, the lobes small to large. Stamens 2, 4 or 5, usually epipetalous; anthers hisporangiate, inverting early in development, mostly monothecal, dehiscing by longitudinal slits, lacking a fibrous endothecium and disintegration tissue. Pollen in monads, tetrads, or shed as reduced tetrads with one or more cells aborting, without viscin threads. Ovary 1-11-locular, with axile or apical placentation, superior; style impressed or not; stigma capitate or lobed. Fruit drupaceous or a loculicidal capsule; seeds with variable testa; embryo with 2 cotyledons.

PRIONOTEAE

Prionoteae Drude, in Engl. & Prantl, Nat. Pflanzenfam. IV( 1): 72. 1889. Type genus: Prionotes R. Br.

Straggling shrubs or climbers. Leaves alternate, glabrous, margins with a few blunt teeth, a single stalked, glandular hair located in each sinus, leaf base narrow, veins pinnate-reticulate, having abaxial fiber cap contacting neither epidermis. Inflorescence axillary at the end of long branches, flowers solitary; bracts small, grading upward to 1-2 larger bracteoles. Flowers 5-merous. Calyx lobes ciliolate; corolla sympetalous, cylindrical, somewhat constricted at the throat, the lobes small, blunt, recurved. Stamens 5, free, the filaments long, straight; anthers dithecal, located at the throat, lacking appendages. Pollen in tetrads. Ovary 5-locular, with axile placentation, several ovules in each cell, superior; style long, filiform, impressed. Fruit loculicidal. [Lebetanthus Endl.; Prionotes R. Br.]

ARCHERIEAE

Archerieae Crayn & Quinn, Aust. Syst. Bot. 11: 23, 1998. Type genus: Archeria Hook. f.

Shrubs. Leaves alternate, glabrous, entire or serrulate, 1 to several longitudinal veins having abaxial fiber cap contacting neither epidermis. Flowers solitary or in short terminal racemes; bracts present, usually caducous; bracteoles small and paired. Flowers 5-merous. Calyx lobes ciliolate, persistent; corolla sympetalous, broadly cylindrical to campanulate, the lobes imbricate in bud, spreading or recurved. Stamens 5, epipetalous, the filaments short and straight; anthers monothecal, lacking appendages. Pollen in tetrads. Ovary 5-locular, deeply lobed, with axile to basal placentation, ovules numerous; style deeply impressed. Fruit a loculicidal capsule. No obvious morphological synapomorphies are known for Archerieae. [Archeria Hook. f., Fig. 71]

OLIGARRHENEAE

Oligarrheneae Crayn & Quinn, trib. nov. Type genus: Oligarrhena R. Br.

Latin diagnosis: Venae in transsectione folii fasciculis fibrarum epidermides ambo non contingentibus; fibres 4- vel 5-meri staminibus 2 vel 4; ovarium biloculare ovulis in quoque loculo solitariis ad apicem affixis; stylus haud impressus; fructus drupaceus.

Shrubs. Leaves minute, appressed, alternate, entire, with narrow leaf base and 1 to several longitudinal veins with abaxial fiber cap contacting neither epidermis. Flowers in terminal compound racemes or blastotelic spikes; bracts present or absent; bracteoles 2. Flowers 4 or 5-merous. Calyx lobes ciliolate, persistent; corolla sympetalous, cylindrical, the lobes imbricate/valvate in bud, straight or spreading. Stamens 2 or 4, epipetalous; anthers monothecal, lacking appendages. Pollen in monads or shed as reduced tetrads with one or more cells aborting. Ovary 2-locular, with apical placentation, a single ovule in each cell; style very short, not impressed. Fruit a drupe. [Needhamiella Watson; Oligarrhena R. Br.]

RICHEEAE

Richeeae Crayn & Quinn, trib. nov. Type genus: Richea R. Br.

Latin diagnosis: Folia basibus caulem cingentibus cicatricibus annularibus remanentibus; venae in transsectione folii fasciculis fibrarum epidermides ambo contingentibus; nodi multilacunares.

Small trees with unbranched stems or much-branched shrubs. Leaves alternate, glabrous, with a few blunt teeth; leaf base sheathing the stem and falling with the leaf to leave annular scars; the veins parallel, having both adaxial and abaxial fiber caps that extend as a flange to contact each epidermis or the lignified hypodermis; nodes multilacunar. Flowers solitary or in spikes, racemes or panicles; bracts persistent or deciduous; bracteoles 2, deciduous. Flowers 5-merous. Calyx lobes ciliate, persistent; corolla sympetalous, cylindrical, the lobes either fused to form a calyptra that splits transversely and falls at anthesis to leave a persistent basal ring, or lobes free and the corolla tube sometimes constricted at the throat. Stamens 5, free, or epipetalous, if free the filaments short, straight; anthers monothecal, lacking appendages. Pollen in tetrads. Ovary 5-locular, with axile placentation, several ovules in each cell; stigma small, capitate; style impressed. Fruit a loculicidal capsule. [Dracophyll um Labill., Fig. 72; Richea R. Br., Fig. 73; Sphenotoma Sweet]

EPACRIDEAE

Epacrideae Dumort., Anal. Fam. P1. 28. 1829 (as Epacreae). Type genus: Epacris Cav.

Epacridinae Kitt. in A. Rich., Nouv. Elem. Bot. ed. 3., Germ. transl.: 824. 1840 (as Epacreae).

Shrubs. Leaves alternate, glabrous, sessile or shortly petiolate, the veins palmate or reduced to one, with abaxial fiber cap contacting neither epidermis. Flowers solitary, axillary; bracts numerous, persistent, bracteoles more than 2. Flowers 5-merous. Calyx lobes ciliate, persistent; corolla sympetalous, cylindrical, the lobes imbricate in the bud, then spreading. Stamens 5, epipetalous or sometimes free; anthers monothecal, lacking appendages. Pollen in tetrads. Ovary 5-locular, with several ovules in each cell; placentation axile; style impressed. Fruit a loculicidal capsule. [Epacris Cav. (including Rupicola Maiden & Betche and Budawangia Telford), Fig. 74; Lysinema R. Br.; Woollsia F. Muell., Fig. 75]

COSMELIEAE

Cosmelieae Crayn & Quinn, trib. nov. Type genus: Cosmelia R. Br.

Latin diagnosis: Folia basibus caulem cingentibus sed folia cum cortice exuta cicatricibus itaque deficientibus; venae in transsectione folii fasciculis fibrarum epidermidem adaxialem basin versus contingentibus.

Shrubs. Leaves alternate, glabrous; the leaf base sheathing the stem, the sheath and cortex falling with the leaf to leave a stem free of scars, the veins parallel with abaxial fiber cap that contacts the adaxial epidermis toward the leaf base; adaxial leaf stomata tetracytic; nodes unilacunar. Flowers solitary, axillary or terminal, often crowded into heads; bracts not scarious, persistent, I to many; bracteoles deciduous, more than 2. Flowers 5-merous. Calyx lobes persistent; corolla sympetalous, though sometimes petals almost free, cylindrical, sometimes conspicuously constricted at the throat, or spreading; the lobes imbricate to almost valvate in the bud. Stamens 5, epipetalous, or sometimes free and then filaments flat, strongly flexuose; anthers dithecal or monothecal, lacking appendages. Pollen in tetrads. Ovary 5-locular, with several ovules in each cell, placentation axile; style impressed. Fruit a loculicidal capsule. [Andersonia R. Br.; Cosmelia R. Br., Sprengelia Sm., Figs. 76-77]

STYPHELIEAE

Styphelieae Bartl., Ord. Nat. Pl. 158, 1830. Type genus: Styphelia R. Br. Stenanthereae Dumort., Anal. Fam. Pl.: 28. 1829.

Stypheliinae Kitt. in A. Rich., Nouv. Elem. Bot., ed. 3, Germ. transl.: 824. 1840 (as Styphelieae).

Shrubs. Leaves alternate, entire, with leaf base narrow and I to several longitudinal veins having abaxial fiber cap that contacts the abaxial epidermis. Flowers in terminal or reduced axillary racemes; bracts several grading upward in size, or one, mostly small and persistent, sometimes leaflike and caducous; bracteoles 2. Flowers 5-merous. Calyx lobes ciliate, persistent; corolla sympetalous, cylindrical, the lobes straight, spreading or recurved, mostly valvate in bud, occasionally imbricate. Stamens 5, epipetalous; anthers monothecal, with or without appendages. Pollen in monads, sometimes in tetrads or shed as reduced tetrads with one or more cells aborting. Ovary 2-11-locular, each with a single ovule, placentation apical, style not impressed. Fruit drupaceous and brightly colored, sometimes splitting to release endocarp, which sometimes splits into separate pyrenes. [Acrotriche R. Br., Fig. 78; Androstoma Hook. f.; Astroloma R. Br.; Brachyloma Sond.; Coleanthera Shchegl.; Conostephium Lindl.; Croninia J. M. Powell; Cyathodes Labill., Figs. 79-80; Cyathopsis Brongn.& Gris; Decatoca F. Muell.; Leptecophylla C.M. Weiller; Leucopogon R. Br.; Lissanthe R. Br., Fig. 81; Melichrus R. Br.; Monotoca R. Br., Fig. 82; Pentachondra R. Br., Fig. 83; Planocarpa C. M. Weiller; Styphelia R. Br., Fig 84; Trochocarpa R. Br., Fig. 85.] Note: The monophyly of most of the above genera has not been rigorously assessed.

VACCINIOIDEAE

Vaccinioideae Am., Encycl. Brit., ed. 7,5: 118. 1832 (as Vaccinieae). Type genus: Vaccinium L.

Shrubs, trees, or lianas, often epiphytic. Leaves evergreen or deciduous, alternate, nonericoid, margin entire to serrulate or serrate, convolute or less commonly revolute in bud. Leaf epidermal cells lignified or not. Indumentum of elongate and unbranched, or peltate, and glandular to non-glandular hairs, and occasionally papillae. Inflorescence terminal or axillary, racemose, paniculate, or fasciculate, or of solitary flowers; bracts present, bracteoles usually paired, variable in position; flower articulated or continuous with pedicel. Flowers usually 4- or 5-merous, actinomorphic, pendulous. Calyx lobes usually 4 or 5, variable in size, usually persistent, sometimes fleshy; corolla usually sympetalous, urceolate to cylindrical or campanulate, deciduous, with the usually 4 or 5 lobes small to less commonly large. Stamens usually 5-10, usually included, the filaments straight to S-shaped, distinct or connate, unicellular-pubescent or glabrous (smooth or roughened), rarely papillose; anthers tetrasporangiate , inverting early in development, (+ or -) roughened, with or without a pair of dorsal appendages (spurs), or these occurring instead on the distal portion of the filament, or with 2 or 4 projections (awns) arising from the dorso-apical portion of the anther, with or without disintegration tissue, lacking a fibrous endothecium, dehiscing by terminal pores, and these often associated with tubules. Pollen in tetrads, lacking viscin threads. Ovary usually 4- or 5-locular, or 10-locular due to secondary division of each locule, with axile placentation, superior to inferior; style usually impressed; stigma truncate. Fruit a loculicidal capsule, berry, or drupe; seeds with variable testa cells; embryo with 2 cotyledons.

OXYDENDREAE

Oxydendreae Cox, Amer. Midl. Naturalist 40: 505. 1948. Type genus: Oxydendrum DC.

Trees. Leaves deciduous, non-ericoid, entire to serrulate or serrate, convolute in bud. Leaf epidermal cells not lignified. Indumentum of stout, non-glandular hairs. Inflorescence terminal, produced on the shoots of the current season, paniculate, of few to numerous arching-declinate racemes or secondary panicles; bracts present, bracteoles paired, near midpoint to apex of pedicel; calyx not articulated with pedicel. Flowers 5-merous, actinomorphic, with distinctive anatomy. Calyx lobes small, persistent; corolla sympetalous, [+ or -] urceolate, the lobes small; stamens 10, the filaments straight, unicellular-pubescent, lacking spurs; anthers slightly roughened, with a white deposit of disintegration tissue dorsally, dehiscing by terminal, slitlike pores. Ovary 5-locular, with axile placentation, superior; style strongly impressed; stigma truncate. Fruit a loculicidal capsule; seeds with elongate testa cells; embryo with 2 cotyledons. [Oxydendrum DC., Fig. 86]

LYONIEAE

Lyonieae Kron & Judd, Amer. J. Bot. 86: 1298. 1999. Type genus: Lyonia Nutt.

Shrubs, rarely lianas. Leaves evergreen or deciduous, non-ericoid, entire, serrulate, or serrate, convolute or less commonly revolute in bud. Secondary phloem with bands of fibers. Leaf epidermal cells [+ or -] lignified. Indumentum of stalked glandular and/or elongate and eglandular hairs, or ferrugineous, peltate scales. Inflorescence usually axillary, rarely terminal, racemose, paniculate, or fasciculate; bracts present, bracteoles paired, variable in position; calyx usually articulated with pedicel. Flowers usually 5-merous, but rarely 4- to 8-merous, actinomorphic. Calyx lobes usually small, persistent; corolla sympetalous, usually urceolate to tubular, the lobes small. Stamens usually 10, the filaments usually S-shaped, unicellular pubescent, roughened, or smooth, sometimes with paired spurs on the anther-filament junction or upper portion of filament; anthers slightly to clearly roughened, with a white deposit of disintegration tissue dorsally, dehiscing by terminal pores. Ovary usually 5-locular, with axile placentation, superior; style impressed; stigma [+ or -] truncate. Fruit a loculicidal capsule; seeds usually with elongate testa cells; embryo with 2 cotyledons. [Agarista D. Don, Figs. 87-88; Craibiodendron W. W. Sm.; Lyonia Nutt., Figs. 89-90; Pieris D. Don, Fig. 91]

ANDROMEDEAE

Andromedeae Klotzsch, Pfl. Abbild. Beschr. 1838. Type genus: Andromeda L. Andromedinae D. Don, Edinburgh New Philos. J. 17: 157. 1834 (as Andromedeae).

Shrubs. Leaves evergreen, non-ericoid, entire or serrulate, convolute or revolute in bud. Secondary phloem with bands of fibers in Zenobia, these lacking in Andromeda. Leaf epidermal cells [+ or -] lignified in Andromeda, but unlignified in Zenobia. Indumentum of multicellular hairs lacking except for globose, eglandular hairs associated with serrations of Zenobia. Inflorescence axillary or terminal, fasciculate; bracts present, bracteoles paired, basal; calyx articulated with pedicel. Flowers 5-merous, actinomorphic. Calyx lobes small, persistent; corolla sympetalous, urceolate or campanulate, the lobes small. Stamens 10, the filaments straight, unicellular-pubescent or roughened, with 2 or 4 awns arising from the dorso-apical portion of the anther; anthers slightly to clearly roughened, with a white deposit of disintegration tissue dorsally in Zenobia but lacking in Andromeda, dehiscing by terminal pores. Ovary 5-locular, with axile placentation, superior; style impressed; stigma truncate. Fruit a loculicid al capsule; seeds with elongate or [+ or -] isodiametric testa cells; embryo with 2 cotyledons. [Andromeda L., Fig. 92; Zenobia D. Don, Fig. 93]

GAULTHERIEAE

Gaultherieae Nied., Bot. Jahrb. Syst. 11:145. 1889. Type genus: Gaultheria L.

Shrubs, rarely lianas. Leaves evergreen or deciduous, entire, serrulate, or serrate, non-ericoid, convolute in bud. Leaf epidermal cells not lignified. Indumentum of multicellular, multi senate glandular and non-glandular hairs. Inflorescence usually axillary, rarely terminal, racemose, paniculate, or fasciculate, occasionally reduced to a solitary flower; bracts present, bracteoles usually paired, variable in position; calyx articulated with pedicel. Flowers usually 5-merous, actinomorphic. Calyx lobes small to enlarged, often fleshy and colorful in fruit, persistent; corolla sympetalous, usually urceolate to tubular, the lobes small. Stamens usually 10, the filaments [+ or -] straight, unicellular-pubescent, roughened, or smooth, lacking projections, or with 4 minute to conspicuous awns arising from the dorso-apical portion of the anther; anthers slightly to definitely roughened, occasionally with tubules, with or without a white deposit of disintegration tissue dorsally, dehiscing by terminal pores. Ovary usually 5-locular, with axile placentation, superior, rarely half inferior; style impressed; stigma truncate. Fruit a loculicidal capsule or indehiscent berry, often associated with fleshy calyx; seeds usually with [+ or -]isodiametric testa cells; embryo with 2 cotyledons. [Chamaedaphne Moench, Fig. 94; Diplycosia Blume; Eubotrys Nutt., Fig. 95; Gaultheria L. s.l., Figs. 96-98; Leucothoe D. Don, Figs. 99-100; Tepuia Camp, Fig. 101]

VACCINIEAE

Vaccinieae Rchb., Fl. Germ. Excurs. 1(3): 203. 1831. Type genus: Vaccinium L. Thibaudieae Benth. & Hook. f., Gen. P1. 2: 564. 1876.

Vacciniiae Klotzsch, Monatsber. Konigl. Preuss. Akad. Wiss. Berlin 1857: 4. 1857 (as Vaccinieae).

Shrubs, small trees, or lianas, often epiphytic. Leaves evergreen or deciduous, non-ericoid, entire or serrate, convolute in bud. Leaf epidermal cells not lignified. Indumentum of short- to long-stalked glandular or non-glandular hairs. Inflorescence usually axillary, rarely terminal, paniculate, racemose or fasciculate, or solitary axillary flowers; bracts present, bracteoles usually paired, variable in position; calyx articulated with the pedicel or not. Flowers usually 4- or 5-merous, rarely 3 or 6-7-merous, actinomorphic. Calyx lobes small to large, persistent, the calyx winged or not; corolla almost always sympetalous, usually urceolate or tubular, sometimes winged, the lobes small to large. Stamens 5-l0(-14), the filaments straight, sometimes connate, unicellular pubescent, roughened, or smooth, sometimes spurred (with either one spur or, more commonly, a pair of spurs) on the anther-filament junction; anthers slightly to clearly roughened, usually lacking disintegration tissue, usually with tubules, de hiscing by pores or slitlike pores at the end of the tubules. Ovary (3-)4-5(-7)-locular, or 10-locular due to secondary division of each locule, with axile placentation, inferior; style impressed; stigma [+ or -]truncate. Fruit fleshy, a berry or 10-pitted drupe; seeds with short to elongate testa cells; embryo with 2 cotyledons. [Anthopteropsis A. C. Sm.; Anthqpterus Hook.; Cavendishia Lindl.; Ceratostema Juss., Fig. 102; Costera J. J. Sm.; Demosthenesia A. C. Sm.; Didonica Luteyn & Wilbur; Dimorphanthera F. Muell., Fig. 103; Diogenesia Sleumer; Disterigma (Klotzsch) Nied.; Gaylussacia Kunth, Fig. 104; Gonocalyx Planch. & Lindi., Fig. 105; Lateropora A. C. Sm.; Macleania Hook., Fig. 106; Mycerinus A. C. Sm.; Notopora Hook. f.; Oreanthes Benth.; Orthaea Klotzsch; Paphia Seem., Fig. 107; Pellegrinia Sleumer; Plutarchia A. C. Sm.; Polyclita A. C. Sm.; Psammisia Klotzsch; Rusbya Britton; Satyria Klotzsch, Fig. 108; Semiramisia Klotzsch; Siphonandra Klotzsch; Sphyrospermum Poepp. & Endl.; Themistoclesia Klotzsch; Thibaudia Ruiz & Pavon; Utleya Wilbur & Luteyn; Vaccinium L., Figs. 109-113. Note: The monophyly of most of the above genera has not been rigorously assessed.
XII. Appendix 4: Key to Named Groups


1. Plants with [+ or -] herbaceous habit and           Monotropoideae, 2
   reduced embryo
  2. Plants with chlorophyll                                    Pyroleae
  2. Plants lacking chlorophyll; i.e., strongly                        3
     mycotrophic
      3. Stamen appendages present; multicellular
         hairs on leaves present; embryo
         undeveloped, with 30-40 cells                      Pterosporeae
      3. Stamen appendages absent; multicellular
         hairs on leaves usually absent; embryo
         extremely undeveloped, with fewer than
         30 cells                                            Monotropeae
1. Plants shrubs, trees, or lianas; embryo with
    differentiated cotyledons                                          4
   4. Pollen shed in monads; seeds with vascular
       bundle in raphe; anther with a fibrous
       endothecium, opening by longitudinal slits             Enkianthus
                                                        (Enkianthoideae)
   4. Pollen shed in tetrads; seeds lacking a
       vascular bundle in raphe; anthers usually
       lacking a fibrous endothecium (except
       Bejaria), opening by pores or slits                             5
       5. Flowers wind pollinated, inconspicuous,
          imperfect, with perianth of distinct
          tepals, the stigma flabellate to
          pinnatifid; leaves always ericoid                  Ericoideae,
                                                               Empetreae
       5. Flowers usually [+ or -] showy and
          insect or bird pollinated,
          usually perfect, with perianth
          of a calyx and usually sympetalous
          corolla, the stigma truncate to [+ or -]                     6
          capitate, rarely cup shaped or funnel
          shaped; leaves ericoid to broad and flat

          6. Anthers bisporangiate, monothecal;
             leaves usually [+ or -] parallel
             veined; stamens equaling the number
             of corolla lobes                          Styphelioideae, 7
             7. Fruit a loculicidal capsule                            8
                 8. Leaf bases broadly sheathing
                    the stem                                           9
                   9. Leaf scars annular                        Richeeae
                   9. Leaf scars absent; stems smooth         Cosmelieae
                8. Leaf bases not sheathing (but
                    may be stem clasping)                             10
                   10. Leaf margins serrate/
                       serrulate, anthers dehiscing
                       by two slits                           Prionoteae
                   10. Leaf margins entire, anthers
                        dehiscing by a single slit                    11
                       11. Flowers pedicellate above
                           the upper bracts                   Archerieae
                       11. Flowers not pedicellate
                           above the upper bracts             Epacrideae
              7. Fruit a drupe                                        12
                 12. Ovary 2 locular, corolla
                     glabrous inside                       Oligarrheneae
                 12. Ovary 1-10 locular, corolla
                      hairy inside or, if glabrous,
                      then ovary 1 locular                   Styphelieae
6. Anthers tetrasporangiate, dithecal; leaves
   pinnate to palmately veined; stamens usually
   twice the number of corolla lobes, occasionally
   reduced and equaling the corolla lobes                             13
   13. Ovary inferior; anthers with tubules               Vaccinioideae,
                                                              Vaccinieae
   13. Ovary superior; anthers usually lacking
       tubules                                                        14
       14. Anthers inverting just before anthesis;
           fruit with an endocarp, indehiscent;
           1 to a few ovules per locule                      Arbutoideae
       14. Anthers inverting early in development
           of the flower; fruit lacking an endocarp,
           usually dehiscent; [+ or -] several
           to numerous ovules per locule                              15
           15. Corolla persistent; leaves ericoid            Ericoideae,
                                                           Ericeae, p.p.
           15. Corolla deciduous; leaves ericoid
               to broad and flat                                      16
               16. Fruit a [+ or -] septicidal
                   capsule                                            17
                   17. Stamens epipetalous 2/3
                       length of filament                      Diplarche
    17. Stamens not adnate to corolla                                 18
        18. Endothecium present and fibrous;
            flowers more than 5-merous                           Bejaria
                                                             (Bejarieae)
        18. Endothecium lacking, or present and
            lacking a fibrous layer; flowers 4- to
            5-merous                                                  19
            19. Seeds papillose; pedicel lacking                Daboecia
                bracteoles; filaments smooth                (ericoideae,
                                                                Ericeae)
            19. Seeds not papillose; pedicel with
                2 bracteoles; filaments roughened
                or unicellular-pubescent                              20
                20. Leaves ericoid; viscin threads
                    usually absent                                    21
                    21. Petals distinct                      Ericoideae,
                                                               Bejarieae
                    21. Petals connate                        Phyllodoce
                                                            (Ericoideae,
                                                           Phyllodoceae)
                20. Leaves not ericoid, although
                    sometimes with revolute
                    margins; often with viscin
                    threads                                           22
                    22. Pedicel usually not
                        articulated with flower;
                        flowers usually
                        zygomorphic and with spots
                        or a blotch; bracts
                        usually scarious; ovary
                        [+ or -] ovoid to
                        cylindric; perulae usually           Ericoideae,
                        present                           Rhodoreae p.p.
                    22. Pedicel articulated with
                        pedicel; flowers usually
                        actinomorphic, lacking
                        spots or a blotch; bracts
                        usually not scarious;
                        ovary [+ or -] globose to
                        subglobose; perulae                  Ericoideae,
                        lacking                        Phyllodoceae p.p.
16. Fruit usually a loculicidal capsule,
    occasionally a berry                                              23
    23. Leaves ericoid; bud scales lacking;
        anthers lacking disintegration tissue;
        fibers lacking in petiole and leaf
        midrib                                                        24
        24. Pith Calluna-type; leaves decussate;
            pedicel with 2 bracteoles;
            multicellular, branched or
            fasciculate hairs present                      Cassiopoideae
        24. Pith homogeneous; leaves alternate;
            pedicel lackin g bracteoles;
            multicellular hairs lacking                Harrimanelloideae
    23. Leaves not ericoid, although they may have
        a revolute margin; bud scales present;
        anthers usually with disintegration
        tissue; fibers associated with vascular
        tissue in petiole and leaf midrib                             25
        25. Secondary phloem with bands of fibers;
            stamens lacking appendages or with 2
            spurs at anther-filament junction or
            on upper portion of filament; leaf
            epidermis [+ or -] lignified;                 Vaccinioideae,
            filaments usually S-shaped                          Lyonieae
        25. Secondary phloem lacking bands of
            fibers (except Zenobia, which has 4
            awns on anthers); stamens lacking
            appendages or with 2 or 4 awns
            arising [+ or -] from anther apex;
            leaf epidermis not lignified (except
            Andromeda, which has 2 awns on
            anthers); filaments straight                              26
26. Leaves lacking multicellular
    hairs, except for [+ or -]
    globose, non-glandular hairs
    associated with serrations in                         Vaccinioideae,
    Zenobia; stomata anomocytic                              Andromedeae
26. Leaves with scattered gland-
    headed, multi-cellular hairs;
    stomata usually paracytic                                         27
    27. Pedicel not articulated;
        inflorescence terminal;
        flowers borne on shoots of
        the current season; anthers
        not appendaged; fruits                            Vaccinioideae,
        [+ or -] elongate-ovoid                              Oxydendreae
    27. Pedicel articulated with
        flower; inflorescences
        usually axillary; flowers
        usually on shoots of the
        previous season; anthers
        often with 4 "apical"
        appendages (awns); fruits
        [+ or -] globose to                               Vaccinioideae,
        subglobose                                          Gaultherieae
Table 1

Morphological characters and character states used in the phylogenetic
analysis of Ericaceae

 1. No basal rosette (0); basal rosette present (1)

 2. Plants woody (0); plants herbaceous (1)

 3. Chlorophyll present (plants green)(0); chlorophyll apparently
    lacking (plants white-reddish)(1)

 4. Shoots emerge from soil erect (0); shoots emerge from soil
    nodding (1)

 5. Plants do not blacken upon drying (0); plants blacken upon
    drying (1)

 6. Trace unbranched in petiole/leaf base (0); trace branched in
    petiole/leaf base (1)

 7. Secondary phloem without bands of fibers (0); secondary phloem
    with bands of fibers (1) (a)

 8. Pith heterogeneous (0); pith homegeneous (1); pith Calluna-
    type (2)

 9. Bud scales present (0); bud scales lacking (1)

10. Leaves convolute in bud (0); leaves revolute in bud (1)

11. Leaves alternate (0); leaves decussate (1); leaves whorled
    (2)

12. Leaves variously scattered along stem (0); leaves distinctly
    pseudoverticillate and appearing to be whorled at ends of
    branches (1) (b)

13. Leaves more of less flat (0); leaves ericoid, with an
    abaxial groove (1) (c)

14. Leaves petiolate to narrowed at base and sessile (0);
    leaves broadly sheating at base (1)

15. Leaf venation pinnate (brochidodromous, reticulodromous,
    etc.)(0); leaf venation more or less palmate (with 2 to
    several basal veins prominent and convering)(1); leaf
    venation appearing more or less parallel (2)

16. Vein reticulum more or less open, the secondary and tertiary
    viens not equally prominent (0); vein reticulum dense, the
    secondary and tertiary veins more or less equally prominent
    (1)

17. Leaves deciduous (0); Leaves persistent (1)

18. Leaf midrib (as seen in cross-section) unifacial (0); leaf
    midrib bifacial (1)

19. Fibers associated with leaf midrib (0); fibers lacking or
    poorly developed (1) (d)

20. Fiber strands absent from leaf mesophyll (0); fiber strands
    present in leaf mesophyll (1)

21. Leaf epidermis not lignified (0); leaf epidermis at least
    slightly lignified (1)

22. Multicellular hairs on leaves present (0); multicellular
    haris on leaves lacking (1)

23. Multicellular haris not as following (0); multicellular hair
    subsessile, with glandular head with gold secretion layer (1)

24. Multicellular hairs simple to branched or eglandular peltate
    (0); multicellular hairs in the form of peltate scales (with
    secretory head of the Rhododendron type)(1)

25. Multicellular hairs various but not forming a flattened, more
    or less traingular scale (0); multicellular hairs forming
    flattened, more or less narrowly triangular scales (1)

26. Stalk of multicellular hair multiseriate (0); stalk of
    multicellular hair biseriate (1); stalk of multicellular hair
    uniseriate (2)

27. Stomata limited to abaxial leaf surface (0); stomata also on
    adaxial surface (1)

28. Stomata anomocyctic (0); stomata paracytic (1); stomata
    tetracytic (2)

29. Inflorescence axillary at initiation (0); inflorescence
    terminal at initiation (1)

30. Flowers developing on shoots of the previous season (0);
    flowers developing on shoots of current season (1)

31. Inflorescence in bud (0); inflorescence exposed during
    winter (1)

32. Flowers perfect (0); flowers imperfect, more or less
    dioecious (1)

33. Flowers actionomorphic (0); flowers zygomorphic (1)

34. Flowers 5-merous (0); 4-merous or 2-merous (1); more
    that 5-merous (2)

35. Pedicel lacking bracteoles (0); pedicel with more than 2
    bracteoles (1); pedicel with 2 bracteoles (2)

36. Bracteoles apical (0); bracteoles not apical (1)

37. Pedicel not articulated with flower (0); pedicel articulated
    with flower (1)

38. Bracts and bracteoles not searious (0); bracts and bracteoles
    scarious (1)

39. Perianth differentiated into a calyx and corolla (0);
    perianth of tepals, or missing clayx and/or corolla (1)

40. Calyx of imbricate lobes (0); calyx of valvate lobes (1)

41. Calyx deciduous (0); withering persistent (1); persistent not
    withering (2); colorful and fleshy (persistent) (3)

42. Adaxial calyx stomata lacking (0); adaxial calyx stomata present
    (1)

43. Abaxial calyx stomata lacking (0); abaxial calyx stomata present
    (1)

44. Sympetalous (0); choripetalous (1)

45. Corolla pendulous (0); corolla held erectly to more or less
    horizontally (1)

46. Base of petals not or only slightly saccate (0); base of petals
    conspicuously saccate (1)

47. Corolla with spots or blotch (0); spots or blotch lacking (1)

48. Corolla deciduous (0); persistent (1); tepals persistent (2)

49. Corolla lacking unicellular hairs on inside surface (0); corolla
    with unicellular hairs on inside surface (1)

50. Corolla lacking multicellular hairs on outer surface (0); corolla
    with multicellular hairs on outer surface (1)

51. Stamens 2x corolla lobes (0); stamens less than 2x corolla lobes
    (1)

52. Stamens free from the corolla (0); epipetalous (1)

53. Filaments straight (0); filaments S-shaped (1)

54. Filaments gradually tapering toward base (0); filaments
    conspicuously asymmetrically dilated at or below middle (1)

55. Filaments with unicellular hairs (0); filaments papillose or
    minutely roughened (1); filaments smooth (2)

56. Anthers noninverting (0); anthers inverting late in development
   (1); anthers inverting early in development (2)

57. Anthers tetrasporangiate, dithecal (0); anthers bisporangiate,
    monothecal (1)

58. Anthers more or less papillose (0); anthers more or less smooth
    (1)

59. Stamen appendages present (0); stamen appendages absent (1)

60. Appendage on anther (0; appendage on anther-filament junction (1);
    appendage on filament (2)

61. Anther appendages 2 in number (0); anther appendages 4 in number
   (1)

62. Tubules on anthers lacking (0); tubules on anthers present (1)

63. Disintegration tissue on back of anthers lacking (0);
    disintegration tissue present (1)

64. Disintegration tissue not extending onto the spurs (appendages) or
    spurs absent (0); disintegration tissue extending onto the spurs
    (1)

65. Anthers not hippocrepiform (0); anthers hippocrepiform (1)

66. Endothecium present and with a fibrous layer (0); endothecium
    absent or present and without a fibrous layer (1) (e)

67. Exothecium lacking (0); exothecium present (cells of anther
    epidermis thickened)(l) (e)

68. Pollen shed as monads (0); pollen shed as tetrads or as reduced
    tetrads with only one functional grain (1)

69. Viscin threads lacking (0); viscin threads associated with pollen
    (1)

70. Stigma unlobed or with simple lobes (0); stigma lobes flabellate
    to pinnatifid (1)

71. Stigma not surrounded by a collar (0); stigma surrounded by a
    collar (1)

72. Ovary superior (0); ovary inferior (1)

73. Style impressed into ovary (0); style gradually tapering in ovary
    (1); style articulated with ovary but neither impressed nor
    tapering (1)

74. Ovary more or less smooth (0); ovary papillose (1)

75. Number of carpels equals number of petals (0); number of carpels
    less than number of petals but more than one (1); number of
    carpels more than the number of petals (2)

76. Ovary with several locules, the placentation axile (0); ovary
    with several locules, the placentation partly intruded parietal
    (1); ovary with a single locule, the placentation distinctly
    parietal throughout (2)

77. Ovary with few to numerous ovules per locule (0); ovary with only
    one ovule per locule (1)

78. Fruit dehiscent (0); fruit indehiscent (1)

79. Fruit a loculicidal capsule (or fruit noncapsular) (0); fruit a
    septicidal capsule (1); capsule irregularly circumcissile (2)

80. Capsule with carpel midvein lacking thickened sutures (0);
    capsule with carpel midvein with thickened sutures (1)

81. Fruit lacking bony endocarp(s) (0); fruit drupaceous, i.e., with
    a single pyrene (1); fruit drupaceous with several pyrenes (2)

82. Fruit more or less brown (0); fruit brightly colored or appearing
    black (very dark red) (1)

83. Fruit more or less ovoid to cylindric (0); fruit globose to
    subglobose (t) (f)

84. Seeds without tails (0); seed with tails (expanded portion of
    testa at hilum and chalazal ends) (1)

85. Embryo well developed, with 2 cotyledons (0); embryo undeveloped,
    with 30-40 cells (1); embryo extremely undeveloped, with ca. 10
    cells (2)

86. Ellagic acid lacking (0); ellagic acid present (1)

87. Gossypetin absent (0); gossypetin present (1)

88. Methyl salicylate absent (0); methyl salicylate present (1)

89. Basic chromosome number x = 11(0); x 12(1); x = 13 (2); x = 8(3);
    x = 18(4); x 19(5); x =9(6); x 14(7)

90. Locules not divided (0); locules incompletely or completely
    divided by outgrowth of ovary wall (1)

91. Capsules with valves not cobwebby on edges (or fruit noncapsular)
    (0); loculicidal capsules with valves cobwebby on edges (1)

(a) This character was recorded from older stems.

(b) Recorded as the length of innovation with leaves attached.

(c) Recorded from cross-sections of leaves made 2/5 of lamina length up
from the base of the lamina.

(d) Recorded from hand sections, fibers on the abaxial side of the
midrib vascular bundle. Observed by making a transverse section of the
anther at the apparent apex.

(f) Based on length:width ratio.


VII. Acknowledgments

This study was supported by National Science Foundation grants BSR-8821264 (K.A.K.), DEB-9407350 (K.A.K.), INT-9602886 (K.A.K.) and DEB-9628841 (J.L.L.), Swedish Natural Science Research Council (NFR) grant B-BU 08950 (A.A.A.), Australian Research Council grant A19330712 (C.J.Q.), A19700885 (C.J.Q. and P.A.G.), a grant from the Technology Branch of the Department of Industry, Science, and Technology, Australia (P.A.G.), an Australian Postgraduate Award (D.M.C.), and the Wake Forest University Research and Publication Fund. Material was kindly provided by: W. Anderson; M. Chase, Royal Botanic Gardens, Kew; D. Chamberlain and R. Hyam, Royal Botanic Gardens, Edinburgh; A. Denton; F. Hrusa; D. Nickrent; G. Picon, S. Vander Kloet. Thanks also to Kent Perkins for assistance in processing specimen loans to FLAS.

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(1.) Department of Biology Wake Forest University Winston-Salem, NC 27109-732S, U.S.A. (author for correspondence)

K.A. KRON (1), W.S. JUDD (2), P.F. STEVENS (3), D.M. CRAYN (4), A.A. ANDERBERG (5), P.A. GADEK (6), C.J. QUINN (4), AND J.L. LUTEYN (7)

(2.) Department of Botany University of Florida Gainesville, FL 32611-8526, U.S.A.

(3.) Harvard University Herbaria 22 Divinity Avenue Cambridge, MA 02138, U.S.A. Current address: Missouri Botanical Garden Saint Louis, MO 63166-0299, U.S.A.

(4.) National Herbarium of New South Wales Royal Botanic Gardens Sydney Mrs. Macquaries Road Sydney 2000, Australia

(5.) Swedish Museum of Natural History Stockholm S-10404, Sweden

(6.) School of Tropical Biology James Cook University Cairns 4870, Queensland, Australia

(7.) The New York Botanical Garden Bronx, NY 10458-5126, U.S.A.
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Author:Kron, K.A.; Judd, W.S.; Stevens, P.F.; Crayn, D.M.; Anderberg, A.A.; Gadek, P.A.; Quinn, C.J.; Lutey
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