Phylogenetic interrelationships in the order

Primulales, with special emphasis on the

family circumscriptions
Arne A. Anderberg and Bertil StQhl

Abstract: Cladistic parsimony analyses, based on morphological data, have been undertaken with the
purpose of identifying major monophyletic groups and phylogenetic interrelationships within the
Can. J. Bot. Downloaded from www.nrcresearchpress.com by UNIV CHICAGO on 02/16/13

Primulales. Actinidia (Actinidiaceae, Ericales) and three genera from two families of the Ebenales
(Diospyros of the Ebenaceae and Marzilkara and Monotheca of the Sapotaceae) were used as outgroups
in the analyses. The results indicate that the Primulaceae, Theophrastaceae, and Myrsinaceae (excluding
Maesa) represent three major monophyletic groups. The Myrsinaceae were found to be paraphyletic,
with the majority of taxa forming a monophyletic group but with the genus Maesa constituting the sister
group of the Primulaceae. It is proposed that Maesa should be raised to the rank of family to obtain
strictly monophyletic groups in the Primulales. The genera Aegiceras and Coris, for which family
affinities have been controversial, are well nested within the families Myrsinaceae and Primulaceae,
respectively.
Key words: Primulales, Theophrastaceae, Myrsinaceae, Primulaceae, morphology, taxonomy,
phylogeny, cladistics, classification.

Resume : Les auteurs ont effectuC des analyses cladistiques en parcimonie, basCes sur des donnCes
For personal use only.

morphologiques, afin d'identifier les principaux groupes monophylCtiques et les interrelations

phylogCnCtiques chez les Primulales. Ils ont utilisC comme groupe externe pour les analyses, le genre
Actinidia (Actinidiaceae, Ericales) et trois genres appartenant i deux familles d'EbCnales (le genre
Diospyros des EbCnaceae, ainsi que les genres Manilkara et Monorheca des Sapotaceae). Les rCsultats
indiquent que les Primulaceae, les ThCophrastaceae et les Myrsinaceae (excluant les Maesa) representent
trois groupes monophylCtiques majeurs. On constate que les Myrsinaceae sont paraphylCtiques, la
majoritt des taxons formant un groupe monophylCtique, mais le genre Maesa constituant un groupe
frtre des Primulaceae. Les auteurs proposent que le genre Maesa devrait Ctre ClevC au rang de famille
dans l'ordre afin d'obtenir des groupes strictement monophylCtiques chez les Primulales. Les genres
Aegiceras et Coris, pour lesquels les affinitts familiales ont CtC controversCes, sont bien placCs dans les
familles Myrsinaceae et Primulaceae, respectivement.
Mots clis : Primulales, ThCophrastaceae, Myrsinaceae, Primulaceae, morphologie, taxonomic,
phylogCnie, cladistique, classification.
[Traduit par la rtdaction]

Introduction flowers, and tenuinucellate ovules, but unlike the Asteridae,

The Primulales are a small group of flowering plants consist-
ing of only three families, the Myrsinaceae R.Br., the Theo-
phrastaceae Link, and the Primulaceae Vent. Recent
estimates (Mabberley 1990; Brummitt 1992; S t h l 1993)
suggest that the ~ ~ r s i n a c e consist
ae of 37 genera and about
1250 species, the Theophrastaceae 6 genera and 90 species,
and the Primulaceae about 20 genera and 800 species. The
Primulales share several floral features with the members of
the subclass Asteridae, e.g., sympetalous, haplostemonous
Received September 12, 1994.
A.A. Anderberg. Department of Phanerogamic Botany,
Swedish Museum of Natural History, P.O. Box 50007, S-104
05 Stockholm, Sweden.
B. St%hl.Department of Botany, University of Stockholm,
S-106 91 Stockholm, Sweden.
Can. J. Bot. 73: 1699- 1730 (1995). Printed in Canada 1 Imprime au Canada
have free central placentae and bitegmic ovules. Therefore
the Primulales are traditionally placed in the subclass Dil-
leniidae (e.g., Cronquist 1981; Takhtajan 1987).
The Myrsinaceae include mainly shrubs and small to
medium-sized trees and have a worldwide tropical and sub-
tropical distribution. In the past, myrsinaceous plants also
inhabited areas that nowadays are temperate (e.g., Conwenz
1886; Chandler 1964). Characteristic of the Myrsinaceae are
the schizogeneous secretory cavities that appear as lines or
dots on vegetative and reproductive parts. Also characteristic
of many genera are the ovules that are distinctly immersed
and more or less concealed by the placental tissue. All genera
of the Myrsinaceae, with the exception of Maesa, have one-
seeded, drupaceous fruits.
Pax (1889) recognized four subfamilies in the Myrsinaceae:
the Theophrastoideae comprising the two tribes Monothe-
ceae and Theophrasteae; the Aegiceratoideae and the

Maesoideae, both monogeneric (Aegiceras and Maesa); and
the Myrsinoideae, including the remaining part of the family,
and further subdivided into four tribes, Myrsineae, Cono-
morpheae, Ardisieae, and Hymenandreae, chiefly on differ-
ences in corolla aestivation.
Mez (1902, 1903) raised the Theophrastoideae to the rank
of family and excluded Monotheca, which was transferred to
the Sapotaceae in agreement with Radlkofer (1889). The
Myrsinaceae were subdivided into two subfamilies, the Mae-
soideae and the Myrsinoideae, the latter comprising the two
tribes Ardisieae, with many ovules arranged in more than
one series, and Myrsineae, with few ovules arranged in a sin-
gle series. ~ e ~ i c e r was
a s included as a member-of the tribe
Ardisieae of the Myrsinoideae.
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The Theophrastaceae are today considered to be a sepa-
rate family (Mez 1903; Cronquist 1981; Takhtajan 1987;
St%l 1990a; Thorne 1992) and were considered by some
(e.g., Melchior 1964) to be the most primitive family of the
order. The Theophrastaceae comprise shrubs and small trees
and have rather robust, staminodial flowers and berrylike
fruits. In several respects they resemble the Sapotaceae, e.g.,
in their firm and thick corolla with well-develo~ed tube.
extrorsely dehiscent anthers, and staminodes fixed in the
corolla lobe sinuses. The Theophrastaceae are today con-
fined to tropical America, but fossil leaves were reported
from central European tertiary deposits (Schindehiitte 1907).
The members of the third family, Primulaceae, were,
For personal use only.
according to Cronquist (1981), separable from those of the
other two by their herbaceous or exceptionally half-shrubby
habit and dry, many-seeded capsular fruits. In spite of the
obvious differences there are also many similarities between
the Myrsinaceae and the ~rimulaceae-,such as the schizo-
geneous cavities found in the Myrsinaceae and in several
Lysimuchia species. Handel-Mazzetti (1928) saw this as a
linking character between the two families and proposed that
Lysimachia should be placed at the beginning of the Primula-
ceae system because of its apparent affinities with the Myr-
sinaceae. As further support of his view, Handel-Mazzetti
(1928) pointed out the floral similarity of some Lysimuchia
(sect. Apodanthera) with members of the Myrsinaceae and
stressed the Maesa-habit shown by Lysimachia insignis
(subg. Idiophyton Hand.-Mazz.) with respect to its loose
lateral racemes protruding along the stem. A similar view
regarding Lysimachia was also expressed by Ray (1956),
who stated that the shrubby representatives of this genus
approached the Myrsinaceae in their floral morphology. In a
recent paper, Judd et al. (1994) pointed out that a phylo-
genetic study of the Primulales was needed because they sus-
pected that the Myrsinaceae is a paraphyletic group that has
given rise to the Primulaceae.
The Primulaceae are usually subdivided into five tribes:
Primuleae (or Androsaceae), Samoleae, Lysimachieae,
Cyclamineae, and Corideae (Pax 1889; Pax and Knuth
1905). The majority of species are placed in either of the two
tribes, Primuleae with imbricate or quinquncial corolla aesti-
vation, or the Lysimachieae with twisted corolla aestivation.
The Primuleae consist of genera such as Primula, Andro-
sace, Douglasia, Dodecatheon, and Soldanella. The Lysima-
chieae include the genera Glaux, Trientalis, Anagallis, Aste-
rolinon, Pelletiera, and the large and heterogeneous
Lysimuchia. The Cyclamineae usually comprise Cyclamen
only, although Pax (1889) included also Dodecatheon in that
Can. J. Bot. Vol. 7 3 , 1995
tribe. The Samoleae and the Corideae are both monogeneric,
containing Samolus and Coris, respectively. ROsvik (1968)
added another monogeneric tribe, the Ardisiandreae, for the
genus Ardisiandra Hook., which he considered to be so dif-
ferent that it could no longer be included in the Primuleae.
The Primulaceae are mainly distributed in temperate and
arctic areas of the northern Hemisphere and only the genus
Samolus has its main species diversity concentrated in the
southern Hemisphere.
Aberrant taxa
Three genera of the Primulales have features that make them
unique compared with the main stock of genera. Maesa
Forssk. (Myrsinaceae) with its many-seeded capsules and
perigynous flowers was placed in a separate tribe by Hooker
(18766) and has been treated by others as a separate tribe or
subfamily (tribe Maeseae of de Candolle 1844; subfamily
Maesoideae of Pax 1889; Mez 1902; Thorne 1983, 1992;
Takhtajan 1987).
Aegiceras Gaertn. is adapted to life in mangrove vegeta-
tion, which is expressed by its elongated fruit with the seed
germinating before leaving the fruit (Rendle 1925). Aegi-
ceras is the only genus of the Primulales known to have exal-
buminous seeds, but this is probably an autapomorphy linked
to its specialized germination. Aegiceras has been raised by
some to family rank, Aegicerataceae (Thorne 1983; Takhta-
jan 1987; Dahlgren 1989), but more often it has been
included in the Myrsinaceae (Hooker 1876b; Mez 1902;
Cronquist 1981; Thorne 1992).
The third enigmatic taxon is the primulaceous genus
Coris, which is a shrublet with zygomorphic flowers.
Hooker (1876~)placed Coris in the monogeneric tribe
Corideae, a view that has also been followed by others (Pax
1889; Pax and Knuth 1905; ROsvik 1968). Takhtajan (1987)
placed Coris in a separate subfamily, the Coridoideae, and
still others have been of the opinion that Coris does not
belong in the Primulaceae at all. Agardh (1858) treated Coris
as a separate family called the Corideae (often cited as
Coridaceae Agardh by later authors, e.g., Takhtajan 1987;
Dahlgren 1989), and Airy Shaw (1951) suggested that it
belonged more properly in the Lythraceae (cf. Sattler 1962).
Today, most workers seem to agree that Coris is nothing but
a derived member of the Primulaceae (Hutchinson 1969;
Cronquist 1981; Takhtajan 1987; Thorne 1992; Chant 1993).
Although the taxonomic treatments of Maesa, Aegiceras,
and Coris differed over the years, there is little controversy
about their position in the order Primulales. For the purpose
of the present investigation we also accept that these genera
belong there and that they consequently should be included
in cladistic analyses of this group.
Objectives
The current classification of the Primulales is founded on
rather few characters and to some extent it seems artificial.
The salient features used to diagnose the Primulaceae for
example, seem to be autapomorphies of that group, and it
therefore seemed possible that they were a derived ingroup
in either of the other two families.
The present investigation was initiated to evaluate the cur-
rent classification of the order Primulales and the phyla-
genetic status of the three families included therein. Our
 Anderberg and St8hl
main objective was to identify, and to diagnose, major mono-
phyletic groups, and to find their sister groups. Another
objective was to elucidate the systematic position of the
genera Aegiceras, Maesa, and Coris in relation to the other
genera of the Primulales, under the assumption that they
belong in the order. The analyses will allow us to continue
our investigations of each major monophyletic group within
the framework of a corroborated phylogenetic hypothesis.
This paper, which is the first of two, has its aims set at
formulating a hypothesis of phylogenetic interrelationships
inferred from morphological, palynological, and embryolog-
ical data. In a forthcoming paper we will test this hypothesis
by comparing it with results from our recently initiated
investigation of cpDNA sequence data.
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Morphology
The Primulales display a considerable variation in most
characters, from growth habit to reproductive structures and
embryology. Since we later discuss aspects of character evo-
lution in the Primulales, we found it necessary to give a brief
description of the characters that were used for the cladistic
analyses, many of which are characters that were used by
earlier authors for their classifications of the group. An
abbreviated list of characters used in the analyses is
presented in Appendix 1.
For personal use only.
Growth habit
The taxa belonging to the Primulales are anything from trees
to minute annual herbs, but many transitional forms
are known, leading from perennial herbs and cushion plants
to subshrubs and dwarf shrubs, with an increasing degree of
lignification of the stems. The vast majority of the Primula-
ceae are herbaceous, whereas shrubs or trees prevail in the
other two families. In the Myrsinaceae and Theophrastaceae,
pachycaulous growth forms are known from several genera,
i .e., Clavija, Theophrasta, Tapeinosperma, Badula, Loheria,
and Correlliana (Stihl 1987, 1991; Stone and Whitmore
1970; D'Arcy 1973; Coode 1981; Stone 1991a), and
although the few suffrutescent members of the Myrsinaceae,
i.e., Ardisia subg. Bladhia (Yang and Dwyer 1989) and
Labisia, often have unbranched aerial parts, their habit can-
not be considered homologous to the ~achycauloushabit of
the woody taxa.
The difference in growth habit has always been acknowl-
edged in taxonomic treatments of the order to tell the families
u
apart, but the most extreme view regarding the phylogenetic
importance of lignification is that of Hutchinson (1969), who
believed that the frutescent taxa were not closelv related to
the herbaceous ones at all and that the floral similarities
between the families were due to parallel evolution. Hutchin-
son (1969) consequently recognized an order Myrsinales as
distinct from the Primulales, but this opinion has gained no
support from later authors. Melchior (1964) had earlier
hypothesized that an herbaceous growth habit in the Primula-
ceae is a derived trait in relation to the lignified growth habit
of the two other families.
Leaves
The leaf arrangement is typically alternate, but some taxa of
the Primulaceae have opposite or whorled leaves, particu-
larly in species of the tribe Lysimachieae. In Anagallis, a
majority of the species have alternate leaves, and it could
therefore be assumed that the apparently opposite leaves in
sect. Anagallis are actually alternate. However, the phyllo-
taxis of Anagallis arvensis (sect. Anagallis) was investigated
by Kwiatkowska (1992), who showed that the leaves in this
species are truly decussate. In Trientalis the leaves form a
false whorl but are in fact alternate and held in a pseudoverti-
cillate position. Most taxa of the tribe Primuleae are scapose
with inflorescences borne on short stems crowned by a
rosette of leaves. In Androsace most species have a basal
rosette of leaves, but there are some species (e.g., Androsace
erecta Maxim.) with long internodes and clearly alternate
leaves, which could indicate that the leaves are alternate in
rosettes of this genus. In most Theophrastaceae the leaves are
arranged in two or a few pseudoverticels towards the tips of
branches, or in pachycaulous members, the main stem.
Exceptions are several species of Jacquinia, in which the
leaves are distributed more or less regularly along the young
branches. Pseudoverticillate leaves occur also in some Myr-
sinaceae, e.g., Tapeinosperma. The leaves of Manilkara
(Sapotaceae) are often more or less clustered at the tips of the
branches (Pennington 1990) but not arranged in dense pseu-
doverticels as in the Theophrastaceae.
The leaves are usually short-petiolate or have the blades
gradually tapering towards the base. In some genera of the
Primulaceae, e. g ., Ardisiandra, Cortusa, Cyclamen, Sol-
danella, and several Primula species, the petioles are
distinctly demarcated from the lamina. In some Theophrasta-
ceae and Myrsinaceae, e.g., Clavija and Labisia, the petioles
are sometimes rather long, but the lamina is then more or less
decurrent on the petiole.
Many Primulaceae - Primuleae have dentate or serrate
leaf margins (e.g., Cortusa, Primula, Ardisiandra), but in
the Lysimachieae species have entire leaf margins. In Cycla-
men, the leaves in the "lower" taxa are somewhat lobed but
with completely entire margins. Cartilaginous marginal leaf
teeth are found only in some more derived taxa of Cyclamen,
but these are not homologous with the incised leaf teeth
found in other genera (Anderberg 1994). In Theophrasta-
ceae, leaf margins are either serrate to serrate-spinulose, or
entire, and in the Myrsinaceae, entire-margined leaves seem
to prevail, although most genera usually have one or a few
species with crenate, serrate, or occasionally spinulose leaf
margins. Among the larger genera of the Myrsinaceae,
Maesa may contitute an exception since a majority of its spe-
cies are characterized by variously serrate- or crenulate-
margined leaves (Mez 1902).
Another aspect of the leaves is their vernation (ptyxis).
Many Primula species have revolute leaf vernation, whereas
others have the leaves involute in bud, and this has been used
as a key character in the classification of the genus. All other
genera of the Primulaceae, except Cortusa and Dionysia, as
well as the other families of the order, seem to have involute
or conduplicate leaves. The true state in small-leaved plants
is often hard to assess, as in Monotheca, for example, in
which the young leaves are plane but at a very early stage of
development slightly involute towards the base. The same is
true for Jacquinia, most species of which have small leaves
like those of Monotheca.
Leaf anatomy
A diagnostic feature of the Myrsinaceae is their schizogeneous
 1702 Can. J. Bot. Vol. 7 3 , 1995

Fig. 1. Myrsinaceae. (A) Embelia phillipinensis, flower. (B) Wallenia laurifolia, flower. (C) Heberderzia
bahamensis, flower. (D) Parathesis cubana, flower. (E) Myrsirze afn'cana, flower. ( F ) Oncosternurn sp.,
flower. (G) Aegiceras, calyx surrounding style. (H) Aegiceras, corolla. (I) Aegiceras, anther with
compartments. (J) Ardisia elliptica, anther with compartments. (K) Ardisia crenata, flowers and buds.
(L) Ardisia crenata, pseudoumbellate inflorescence in fruit. (M) Ardisia crenata, detail of petal surface
showing capitate trichomes. (A) Fenix 12597 (S) ( X 12). (B) von Tiirckheim 2682 (S) ( x 12). (C) Santesson
26665 (S) (x5). (D) Ekman 10466 (S) ( ~ 8 ) (E) . Goldblatt 3114 (S) ( x 12). (F) Lowry & Randrianasolo 4420
(S) ( ~ 8 ) (G-I)
. Stihl 800 (GB) ( x 5 , x 5 , x 1I). (J) Degener 12404 (S) (X 8). (K-M) Unvouchered,
cultivated in Stockholm ( ~ 5 x, 2 , x8).
Can. J. Bot. Downloaded from www.nrcresearchpress.com by UNIV CHICAGO on 02/16/13
For personal use only.
 Anderberg and St%hl
secretory cavities that are present in virtually all taxa, albeit
sparse in some species. In the Primulaceae, several species
of Lysimachia also have conspicuous secretory cavities in the
leaves and often also in the corolla or other floral parts, e.g.,
Lysirnachia thyrsiflora, Lysimachia atropulpurea, Lysi-
machia vulgaris, and Lysimachia punctata. In species lack-
ing secretory cavities in the aerial parts, homologous
structures are often found in the roots, e.g., Lysirnachia nurn-
rnularia (Ray 1956). Such secretory cavities are also found
in Sarnolus,and they are very in Coris, particu-
larly in the calyx. Metcalf and Chalk (1950) stated that
schizogeneous secretory cavities also occur in Anagallis
p.p., Centunculus, and Androsace p.p., but we have not been
able to verify these observations. The Theophrastaceae do
Can. J. Bot. Downloaded from www.nrcresearchpress.com by UNIV CHICAGO on 02/16/13
not have schizogeneous secretory cavities, although this was
implied by Cronquist (1981).
Many genera of the Myrsinaceae have epidermis cells
with gelatinized inner walls (Grosse 1908; Metcalf and
Chalk 1950; Pipoly 1987), although the condition is often
lacking in some species. In some genera, e.g., Maesa and
Aegiceras, this feature is lacking altogether.
Extraxylary bundles of unbranched fibres running more
or less parallel to the secondary veins are found in leaves of
virtually all Threophrastaceae, a feature that S t a l (1987,
1989, 1990a, 1991) considered to be one of three synapo-
morphies of the family. Extraxylary foliar fibres also occur
widely in the Sapotaceae (Holle 1892; Rao 1961) but in this
For personal use only.
family are usually branched and not arranged in layers or
strands. Corelliana (D' Arcy 1973), which was included in
Cybianthus by Agostini (1980) and Pipoly (1987), is an
intriguing group since it is similar to Clavija of the Theo-
phrastaceae in growth habit, and like Clavija, also has
subepidermal strands of foliar sclerenchyma (Mez 1902;
Grosse 1908). However, a close examination of the scleren-
chymatic strands in the leaves of Corelliana reveals that they
consist of cells with numerous pits and consequently should
be regarded as sclereids rather than fibres. Short and more
or less irregularly shaped foliar sclereids occur sparsely in
the Primulales, e.g., in Dionysia (Bokhari and Wendelbo
1976) and Aegiceras (Rao 1971).
Foliar crystals in the form of druses occur widely in the
Myrsinaceae (Grosse 1908) and are also present in Jacquinia
(Votsch 1904; S t a l 1990~).Prismatic crystals also occur in
leaves of many Myrsinaceae (Grosse 1908), either substitut-
ing or mixed with the druses. Although widespread in the
Myrsinaceae, the occurrence of these crystals is often rather
unpredictable, as was noted by Lersten (1977) in Ardisia.
calcium oxalate is evidently absent in vegetative parts of the
Primulaceae (Hegnauer 1969).
Trichomes
Several hair types, both glandular and nonglandular, occur
in the Primulales. Immersed capitate hairs occur in the Theo-
phrastaceae and in most Myrsinaceae but not in Maesa or the
Primulaceae. The capitate hairs generally consist of a foot
cell, a stalk cell, and a head with few to many cells. The head
cells are usually radially arranged but in Aegiceras are nar-
row and standing in two more or less parallel rows (Solereder
1908). Although the entire trichome may be immersed, espe-
cially on thick, coriaceous leaves, it is more common that
Fig. 2. Articulated hairs in Ardisiandra (Primulaceae).
Schlieben 4806 (S). Scale bar = 0.5 mm.
only the foot cell and stalk cell are immersed in the leaf
tissue.
Several genera of the Myrsinaceae have minute, brown
scales that are usually circular in shape. In Myrsine afn'cana
these scales occur on young shoots and undeveloped leaves,
appearing as minute, narrow, erect trichomes rather than
scales, but in Conornorpha, for example, the scales are more
conspicuous and cover-large parts of the leaves and inflores-
cences. The immersed hairs of the Theophrastaceae and
Myrsinaceae could possibly be viewed as precursors of these
brown scale-like hairs, but this still needs to be verified.
Pipoly (1987) referred to the immersed, capitate hairs on
leaves of Grarnrnadenia as hydropotes,
- . i.e., epidermal struc-
tures present in many aquatics.
In a number of genera of the Myrsinaceae, small capitate
trichomes are also found on the adaxial surface of the corolla
(Fig. lM), and these "granules" sometimes cover the entire
surface. Pipoly (1987) used the presence of such trichomes
to diagnose the genus Cybianthus in the wider sense advo-
cated by Agostini (1980). The diagnostic value of this char-
acter is weakened by the fact that similar trichomes are found
also on the corolla of several other genera, e.g., in Ardisia,
Heberdenia, Pleiomeris, and notably Ernbelia, the genus
Pipoly (1987) used as outgroup for his cladistic analysis of
Cybianthus.
It is quite possible that all types of capitate trichomes, on
vegetative aswell as reproductive parts, should be regarded
as homologous structures, being secretory on some organs
and in some taxa and nonsecretory in others. In fact, Miller
et al. (1984) studied the ultrastructure of capitate trichomes
on shoot tips of Ardisia crispa and showed that these lacked
the cellular structures considered necessary for secretory
activity. Mennechet (1902) suggested that the secretory
products produced by the immersed glandular hairs of the
Myrsinaceae were kept in the trichome head cells, whereas
in the Theophrastaceae, they occurred between the head cells
and the upper cuticule. This needs to be verified.
 1704 Can. J. Bot. Vol. 73, 1995

Fig. 3. Primulaceae. (A) Primula rosea, flower in transection. (B) Lysimachia epherneru~n(sect. Palladia),
flower. (C) Soldanella montana, flower in transection. (D) Primula denticulata, umbellate inflorescence.
(A-C) Unvouchered, cultivated in Stockholm (x5). (D) Unvouchered, cultivated in Stockholm ( x 1.5).
Can. J. Bot. Downloaded from www.nrcresearchpress.com by UNIV CHICAGO on 02/16/13
For personal use only.

Grosse (1908) noted that nonglandular hairs are very rare
in the Myrsinaceae, again with the exception of Maesa in
which long, uniseriate hairs with tapering tips are quite com-
mon. In the Primulaceae, long articulated hairs are typical of
virtually all genera (Fig. 2) but characteristically absent from
both Cyclamen and Dodecatheon, as already noted by Schwarz
(1955). Coris has articulated hairs confined to the margins of
the calyx lobes, whereas stalked glandular hairs prevail on
other parts. Interestingly, articulated hairs are prevalent in
Ardisia subg. Bladhia, a group of the Myrsinaceae that are
characterized by a suffrutescent habit. Simple multicellular
trichomes similar to the articulated hairs of the Primulaceae
occur also in Clavija, but these do not collapse when dry and
are probably not homologous.
In species of Androsace, Douglasia, and Vitaliana, white
stellate hairs are common, but such trichomes are not found
elsewhere in the Primulales. A few Myrsinaceae, e.g.,
Corelliana (Grosse 1908; Pipoly 1987), have two-armed
(malpighiaceous) hairs, which consist of a few thin-walled
cells (Grosse 1908) and differ markedly from the thick-
walled, sclereid-like, and mostly unicellular hairs of the
Sapotaceae and Ebenaceae (Solereder 1908; Metcalf and
Chalk 1950).
Many Primula and Dionysia species have a highly charac-
teristic farinose coating on young leaves, stems, and inflores-
cences. The absence or presence of this mealy coating is
often characteristic of different species and is also used in
taxonomic treatments of Primula (e.g., Richards 1993). A
few species of Jacquinia produce a thin covering of needle-
like crystals on their twigs, but this is not the same thing as
the farina in Primula, for example.
Inflorescence
Inflorescences in the Primulales are basically racemose (Sat-
tler 1962), and the flowers are arranged in panicles,
racemes, or umbels. In some Primulaceae, e.g., Anagallis,
the flowers appear solitary in the leaf axils, possibly because
of a reduction of a lateral raceme or panicle. Some Myr-
sinaceae, e.g., Myrsine s.lat., have clusters of flowers that
also could be interpreted as being arranged in reduced
 Anderberg and St%hl

Fig. 4. Maesa. (A) Habit of Maesa lanceolata Forssk. ( B ) Flower of M. lanceolata. (C) Flower of
M. lanceolata from the side showing one of the bracteoles. (D) Fruit of M. japonica showing the two
bracteoles and the superior calyx. (E) Flower of M. japonica (Thunb.) Moritzi in transection. (F) Flower of
M. japonica. (A-C) Anderberg 1645 (S) (life size, flowers X 12). (D) Furuse s.n. (S) ( ~ 8 ) (E
. and F)
Furuse s.n. (S) ( ~ 8 ) .
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For personal use only.

racemes or panicles. In other genera of the Myrsinaceae, the
inflorescences are almost umbellate, with the flowers more
or less condensed towards the tips of the raceme or panicle,
a condition that often becomes more pronounced in fruit (cf.
Fig. 1L). Although the inflorescences appear to be more
umbellate in some genera (e.g., Amblyanthus and Aegiceras)
than in others, we have interpreted these pseudoumbels as a
separate character state, not directly comparable to the per-
fect umbel found in the Primulaceae (Fig. 3D).
Each flower is usually subtended by a single bract, which
is inserted at the junction of the pedicel and the rachis, but
the bracts may sometimes be fused with the pedicel to a
greater or lesser extent. The bracts are usually persistent
throughout anthesis, but in some Myrsinaceae, e.g.,

Fig. 5. Theophrastaceae. (A) Jacquinia, flower from above. (B) Jacquinia, flower in transection.
(C) Clavija, flower from above. (D) Clavija, flower in transection. (A and B) Harling & Andersson
24822 (GB). (C and D) Cult. (GB). All 7 x .
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For personal use only.
Tapeinosperrna and Conandrium, the bracts are caducous. In
Maesa the flowers, apart from the bract, are also subtended
by a pair of opposite or subopposite bracteoles (Fig. 4), a
unique character in the Primulales, but occurring also in
Diospyros (Ebenaceae) and Symplocos (Symplocaceae), for
example (Nooteboom 1975).
Corolla and calyx (Figs. 1, 3, 4, 5)
The flowers are typically hypogynous, and only exception-
ally perigynous as in Maesa and Samolus. In some genera the
calyx is synsepalous, forming a shallow cup, e.g., in Ambly-
anthus and Labisia, and several members of the Primula-
ceae. The corolla is sympetalous, open and short-tubed, or
distinctly campanulate with a well-developed tube. In the
Primulaceae, genera such as Primula and Dionysia have a
conspicuously long corolla tube, which is generally much
longer than the calyx. Embelia of the Myrsinaceae deviates
from the general pattern by having choripetalous flowers,
something that some workers (Bartling 1830; de Candolle
1844; Nakai 1943) have used to place it in a separate tribe
Can. J. Bot. Vol. 73, 1995
(the Embelieae) or family (Embelieae J.G. Agardh 1858,
cited as Embeliaceae Agardh by Takhtajan 1987). However,
choripetalous flowers also characterize the genus Heber-
denia.
In the Primulaceae, most genera have erecto-patent or
spreading corolla lobes, except Dodecatheon and Cyclamen,
both of which have distinctly reflexed lobes. Except for
Aegiceras, no species of the Theophrastaceae or the Myr-
sinaceae have such distinctly reflexed corolla lobes, but some
taxa, e.g., Geissanthus, Parathesis, Solonia, and many spe-
cies of Ardisia (see Fig. 1K) may have recurved or revolute
petals.
Decrock (1901) found that the genera of the Primulaceae
differed in the shape of their corolla epidermis cells and that
members of the tribe Primuleae have short and often iso-
diametric cells, whereas those of the Lysimachieae have
elongated cells. ROsvik (1968) made more thorough investi-
gations of a larger sample of species and confirmed
Decrock's (1901) observation. As a curiosity in this context,
ROsvik (1968) pointed out that Samolus has an inner corolla
 Anderberg and St8hl
epidermis with short isodiametric cells and an outer epi-
dermis with very elongated cells. In the Myrsinaceae, the
corolla epidermis is generally provided with short cells, and
the same is true of the Theophrastaceae.
Pollination
Most Primulales seem to be pollinated by insects, chiefly
bees and flies, but there is some variation in the ways insects
are attracted. Somatic nectaries situated on the gynoecium,
indicating insect pollination, are found in Primula and Samo-
lus, for example, and a gynoecium with nectar-producing tis-
sue as in the Primulaceae was also found in Maesa argentea
Wall. (Vogel 1986; S. Vogel personal communication). A
structure similar to a nectary disc was also found in Ardisian-
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dra (Rosvik 1969), but it is not clear whether it produces
nectar or not. Many Myrsinaceae have connate and some-
times poricidal anthers and seem to be buzz-pollinated, but
more fieldwork is needed before a clear understanding of
their pollination biology can be gained. Buzz-pollination is
known from Cyclamen, Dodecatheon, and some Lysimachia,
genera that do not produce any nectar at all. A unique feature
found only in certain Lysimachia species (e.g., Lysimachia
vulgaris L.) is the presence of oil-producing trichomes on
petals and stamens that attract pollinators from a specific
genus of oil-collecting bees (Macropis, Hymenoptera) (Vogel
1986). Still other species of Lysimachia have nectar produc-
ing trichomes, and such hairs are evidently also present in
For personal use only.
some Theophrastaceae (e.g., Jacquinia). Theop/zrasta is
likely to be pollinated by fungus gnats (Diptera) ( S t b l 1987;
Knudsen and St8hl 1994), although no direct observations of
pollinators have been made so far.
A few taxa (e.g., some Lysimachia, Trientalis) have
flowers that do not produce pollen in large amounts, nectar,
or oil. These may be autogamous forms with flowers repre-
senting various reductions from any of the other flower
types.
Androecium
The flowers in all three families are haplostemonous, with
the stamens situated opposite the petals. The stamens develop
from a common primordium together with the petals, in dif-
ferent temporal order. Sattler (1967) observed that the sta-
mens sometimes arose before the petals, whereas in other
genera the petals arose first. The Theophrastaceae were
found to have early petal initiation, like Ardisia, whereas an
early stamen initiation was found in Myrsine and Aegiceras.
Sundberg (1982) found that both kinds of delayed initiation
occurred also in the Primulaceae and interpreted the delayed
stamen initiation to be the derived condition.
Many taxa in the Primulales have anther filaments fused
into a more or less pronounced ring. Being more or less
fused with the corolla, the presence of this filament tube is
not always easily assessed. In fact, the character is inapplica-
ble in taxa where the filaments are completely fused with the
corolla tube, as in Maesa and in several Primulaceae, as well
as in the Sapotaceae. The stamina1 tube is more evident when
it is partly separated from the corolla, as in Aegiceras and
Myrsine, or when distinct ledges occur between the fila-
ments, so that two or more stamens can be removed as a unit
without rupturing the corolla.
In Primulaceae and Myrsinaceae the anthers are introrse
and open by longitudinal slits. In some Lysimachia, and in
several species of different genera of the Myrsinaceae, the
anthers are poricidal, which is likely to be an adaptation to
buzz-pollination. In Aegiceras the anthers are septate, a fea-
ture described as a peculiarity, although the same type of
anthers are present also in several species of Ardisia, e.g.,
Ardisia elliptica Thunb. (Stone 1993b) (see Figs. 11, 1J).
The Theophrastaceae differ from other Primulales by their
extrorse anthers, which Stihl ( 1 9 9 0 ~ )considered to be one
of three synapomorphies of the Theophrastaceae. The state-
ment by Pax (1889) regarding the stamen dehiscence in
Monotheca is incorrect, since it has extrorse anthers like
other Sapotaceae. The inverted anthers in Actinidia are basi-
cally extrorse and open with very short porelike slits.
In many Myrsinaceae and some Primulaceae, the anthers
form a cone, again something that is likely to be correlated
with buzz-pollination. The flowers of the Theophrastaceae
are protandrous, and the stamens form a cone during their
male phase; however, during the subsequent female phase
they become spreading (Figs. 5A, 5B; StPhl 1990b, Fig. 1).
An exception to this occurrence is found in most species of
Clavija, in which staminate and functionally staminate
flowers have the filaments fused into a persisting tube
(Figs. 5C, 5D).
The anthers of the Theophrastaceae have thecae with
mealy accumulations of calcium oxalate crystals. Being most
abundant in the upper part of the thecae, these accumulations
produce a distinct visual contrast to the yellowish pollen
grains. This highly characteristic feature, considered to be
one of three synapomorphies for the Theophrastaceae by
S t b l (1990a), was first described in Deherainia Decne.
(Pohl 1931) but occurs throughout the family.
Another characteristic and often employed diagnostic fea-
ture of the Theophrastaceae is their whorl of staminodes that
are fused with the corolla and alternating with the petals. Sat-
tler (1962), in a study of the floral development of several
taxa belonging to the Primulales, showed that the staminodial
structures in Theophrastaceae originate from the same whorl
as the stamens and petals. Whether these structures really are
staminodial in nature is still unknown, but at least they are
provided with vascular strands, and therefore the interpreta-
tion that they are vestigial stamens seems reasonable. ~ t a m i -
nodial structures similar to those of the Theophrastaceae are
common also in the Sapotaceae and have also been stated to
occur in a few genera of the Primulaceae, viz. Samolus, Sol-
danella, and Lysimachia subg. Seleucia Bigelow (=Steiro-
nema Rafin.). Thenen (191 1) showed that the antisepalous
scales in Samolus are staminodia, provided with a vascular
trace that is one branch of a three-branched vascular system
in the corolla. between the main corolla veins. Thenen
(19 11) also argued that the three-branched vascularization in
the linear segments between the corolla lobes in Soldanella
proved that these are homologous with staminodia. Vascular
strands are entirely missing from the alleged staminodial
structures of Lysimaclzia, and Thenen (191 1) therefore con-
sidered these to be secondary protuberances rather than
staminodes. In Androsace and Douglasia the corolla throat is
characteristically constricted and provided with scales in the
throat, but these are formed by swelling of the corolla tissue
and lack the three-branched vascular system of Samolus and
Soldanella and therefore cannot be considered staminodial.

Pollen
The pollen of most representatives of the Primulales is tricol-
porate, binucleate, and fairly uniform (Erdtman 1951; Johri
et al. 1992), although exine sculpturing has been shown to
vary somewhat between genera (Punt et al. 1974; B. Stihl
unpublished data). Tetracolporate pollen has been reported
from several species of Myrsine s.lat. (Selling 1947; Vasan-
thy and Pocock 1981) and occurs also in some species of
Clavija. Zonocolpate pollen was found to be diagnostic of
most Primula and Dionysia species (Wendelbo 1961c;
Spanowsky 1962; Nowicke and Skvarla 1977). As far as is
known, the outgroup taxa have tricolporate pollen (Erdtman
1951), except Manilkara that has tetra- or occasionally
penta-colporate pollen (Harley 1990), which seems to be a
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modification of the normally tricolporate pollen of the Sapo-
taceae. A variation between tricolporate and tetracolporate
pollen in the same species was also mentioned by Selling
(1947), thus confirming that the number of apertures
represents variation on a common basic pattern.
Gynoecium
The Myrsinaceae and Theophrastaceae are frequently dio-
ecious or subdioecious, and dioecism is also the dominating
sexual systems in Clavija (Mez 1902; Stihl 1991). Both
families are consistently homostylous, but in the Primulaceae
several genera are heterostylous (Schaeppi 1934). In Prim-
ula, over 90% of the species are heterostylous, but Wen-
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delbo ( 1 9 6 1 ~ ) considered this genus to be primitively
homostylous. He also concluded that the other heterostylous
genera (Hottonia and Dionysia) had evolved from within
Primula.
The Primulales have ovaries with a free central placenta,
a feature that is rare among angiosperms and therefore con-
sidered to be diagnostic of the order as a monophyletic
group. Douglas (1936) accepted that the Primulales could be
derived from a "centrospermal stock," and Dickson (1936)
concurred but stated that the Primulaceae are at an evolution-
ary stage in advance of the Caryophyllaceae. The idea of a
common origin for the Caryophyllaceae and Primulaceae
was, however, rejected by Schaeppi (1937) and Stebbins
(1974), who considered the free placentas in the Primulales
and the "centrospermae" to have evolved independently,
and this is also the view that is prevailing among systematists
today. The Myrsinaceae are supposed to have three to six
carpels (Mez 1902; Cronquist 1981) and the Theophrasta-
ceae two, three (Mez 1903), or perhaps five (Mabberley
1990). In the Primulaceae the ovaries are said to be formed
by five carpels (Douglas 1936; Dickson 1936), but Clincke-
maillie and Smets (1992) found no support for this supposi-
tion other than floral symmetry in general. The absence of
septa in the fruit has apparently contributed to the difficulty
in determining the actual number of carpels in the fruit.
The dry capsular fruits are one of the salient features for
separating the Primulaceae from the other two families. The
Theophrastaceae have berries with fleshy pulp and leathery
exocarp, and the Myrsinaceae have more or less fleshy
drupes. In the perigynous genus Maesa, the fruit is also more
or less fleshy but formed both by the ovary and the surround-
ing tissue and therefore cannot be considered homologous to
that of other Myrsinaceae.
In the Myrsinaceae, the number of ovules can be quite
high initially, although only one mature seed generally
Can. J. Bot. Vol. 73. 1995
develops. In many taxa the ovules are few and well spaced
with placental tissue in between, but in others the ovules are
densely set. The ovules are arranged in one or many series
and more or less deeply immersed in the placenta and some-
times almost concealed by the surrounding tissue. The
arrangement of the ovules was considered to be taxonomi-
cally significant in the family, e.g., for distinguishing
between the two subgenera in Maesa (Mez 1902), the only
genus of the Myrsinaceae that like most genera of the Primu-
laceae has many-seeded fruits.
Fruits of the Theophrastaceae are generally few-seeded
(one-seeded only rarely in a few species of Clavija), in spite
of the fact that the number of ovules may be as high as
200-250 (some species of Jacquinia). In this family the
ovules are arranged in several series but are not immersed in
the placenta, although this was claimed by Pax (1889).
Instead, the mature seeds are more or less surrounded by the
placental pulp.
The Primulaceae usually have many-seeded fruits and
ovules arranged in several series. An exception is Coris, in
which the ovules are inserted in a single series at the top of
the placenta from which the seeds are dispersed, leaving a
shallow starlike impression. All the genera of the tribe
Lysimachieae have ovules immersed in the placenta, but this
is also the case in a few other genera, e.g., Cyclamen, Coris,
and Dionysia (Rdsvik 1966).
Mez (1902), as well as Melchior (1964), suggested that
taxa with one or a few ovules evolved from ancestors with
multiovulate ovaries. It is noteworthy that this scenario is
just the opposite of the one proposed by Stebbins (1974),
who postulated that there had been a general evolutionary
trend in the Primulales towards an increased number of
ovules per gynoecium. Stebbins (1974) exemplified this sup-
posed trend by comparing Glaux and Lysimachia with 3-30
ovules per ovary with the groups he considered more
derived, like Primula (30-150 ovules per ovary), and
Cyclamen (100- 150 ovules per ovary).
Seeds
The seeds of the Primulales exhibit variation in their mor-
phology (Fig. 6) as well as their anatomy (Figs. 7- 1l), and
based on gross morphology, two seed types can be distin-
guished: (i) the subglobose and usually rather large seeds
found in most Theophrastaceae and Myrsinaceae (except
Maesa) and (ii) the angular, medium sized to minute seeds
found in virtually all Primulaceae and in Maesa of the Myr-
sinaceae. The subglobose seeds of Myrsinaceae - Myrsinoi-
deae differ from those of the Theophrastaceae by having a
concave hilum area, appearing as a more or less deep, basal
cavity. In addition, the endosperm in seeds of Myrsinaceae-
Myrsinoideae is often more or less ruminate. These morpho-
logical differences are generally paralleled by differences in
anatomy, so that the subglobose seeds of Myrsinaceae and
Theophrastaceae also have endosperm cell walls with irregu-
lar thickenings and narrow constrictions ("pitted" endo-
sperm walls, Figs. 7 and 8) and a thin testa devoid of
crystals. The small angular seeds of the remaining groups
have evenly thickened, smooth endosperm cell walls and a
thick (usually distinctly two-layered) testa with rhomboid
crystals. Exceptions from this general rule are Cyclamen
(Woodcock 1933), Vitaliana, and Douglasia, which all have
endosperm with pitted cell walls, and Myrsine with smooth
 Anderberg and St%hl
Fig. 6. Seed and embryo morphology. ( A ) Clavija lancifolia Desf. ssp. chermontiana (Standl.) Stihl
(Foster and Terceros 13447). (B) Parathesis belizensis Lundell (Contreras 7957). (C) Maesa lanceolata
Forssk. (Bernardi 11070). (D) Prinzula veris L. (Wall s.n.). (E) Asterolinon Iinum-stellatum (L.) Duby
(unvouchered, cultivated in Lund).
5 mrn
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cell walls. An immersed, concave hilum area similar to the
one found in seeds of Myrsinaceae-Myrsinoideae is also
present in Asterolinon and Pelletiera. In Jacquinia, both seed
types are found, although the "primuloid" seeds in this
genus are considerably larger than those found in Primula-
I
ceae and Maesa.
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I The stored nutritive material in seeds of Deherainia (The-
ophrastaceae), which is of the same morphological (pitted)
type as that of Clavija and Theophrasta, was classified by
I
I Mauritzon (1936) as a perisperm developed from the inner
integument. If this is true then the states pitted versus smooth
should be exchanged for perisperm versus endosperm, but
this observation needs to be verified. Hegnauer (1969, 1973)
and Cronquist (1981) stated that seeds of Primulales do not
store starch but instead store oil and amylose, but despite the
striking difference in endosperm cell wall construction, no
differences in seed chemistry have been reported.
Embryology
The embryological characteristics of the Primulales include
the following features: fibrous endothecium, secretory anther
tapetum with uninucleate cells, anatropous bitegmic tenuinu-
cellate ovules with micropyle formed by both integuments,
multicellular archaesporium, Polygonum-type of embryo sac
with ephemeral antipodals, differentiation of endothelium,
albuminous seeds, and nuclear endosperm formation (Johri
et al. 1992). Two genera, Aegiceras and Cyclamen, differ
from the others in having unitegmic ovules, but Netolitzky
(1926) stated that in some genera of the Primulaceae, i.e.,
Cyclamen and Soldanella, the inner integument is partly
resorbed, indicating that unitegmy is a secondary feature in
the Primulaceae. Aegiceras also differs from all other genera
by having exalbuminous seeds. No endosperm haustoria are
known to occur in the Primulales (Dahlgren 1989, 1991).
Data on embryology and ovule morphology in the Myr-
sinaceae are scarce and restricted to studies of a few taxa
only, viz. Aegiceras, Myrsine, Ardisia, and Maesa (Karsten
1891; Warming 1914; Dahlgren 1916; Schnarf 1931; Carey
and Fraser 1932; Sankara Rao 1972), but it is interesting that
the differences between Maesa and other Myrsinaceae seem
to be confirmed by embryological characters. An integumen-
tary tapetum is missing in Aegiceras, Ardisia, and Myrsine,
as well as in the genera Clavija, Theophrasta, and Jacquinia
of the Theophrastaceae (Mauritzon 1936). In contrast,
Maesa and all investigated Primulaceae (except Coris and
Cyclamen) have an integumentary tapetum (Dahlgren 1916;
Raju 1952; Subramanyam and Narayana 1968). The Primu-
laceae have an embryogeny of the caryophyllad type (e.g.,
Soukges 1937; Raju 1952; Subramanyam and Narayana
1968), but Trientalis seems to have an onagrad-type embry-
ogeny (Johansen 1950). Although the Myrsinaceae are some-
times also stated to have an embryogeny conforming to the
onagrad type (Johansen 1950; Johri et al. 1992), this has
apparently been investigated only in Maesa.
The embryo in Primulaceae and Maesa is short and
provided with short, narrow cotyledons (Saint-Hilaire and
Girard 1839; Martin 1946; Burkardt 1979). In Myrsina-
ceae-Myrsinoideae on the other hand, the embryo is usually
long and somewhat curved and has small, often inconspicu-
ous cotyledons. In the Theophrastaceae, the cotyledons are
often broad and foliaceous, but narrow and poorly differen-
tiated cotyledons were recorded in several species of Jac-
quinia (B. S t h l , unpublished data). The variation in embryo
length may partly be correlated to differences in seed size.
Polyembryony has been recorded from a few species of
Ardisia (Braun 1859) and during the course of this study was
also noted in Suttonia australis A. Rich. and Parathesis
belizensis Lundell.
Davis (1966) mentioned that the endosperm formation in
Ebenaceae is nuclear, but it was shown to be cellular in
Diospyros kaki, Diospyros glaucifolia, and Diospyros vir-
giniana by Yamazaki (1972) and Johri et al. (1992). Further-
more, Yamazaki (1972) stated that both the Ebenaceae and
the Sapotaceae have a Chenopodiad type of embryogeny.
Cytology
Faure (1968) found that the chromosome number in Aegi-
ceras conforms well to the common number of 2n = 46
found in several taxa of the Myrsinaceae, e.g., in many
Ardisia species. With regard to the cytological data, both the
 1710 Can. J . Bot. Vol. 73. 1995

Fig. 7. Pitted endosperm cell walls in Corelliana multi'ora. Fig. 8. Pitted endosperm cell walls in higher magnification (Corelliarla
multif2ora). Fig. 9. Smooth endosperm cell walls in Glaux maritirna. Fig. 10. Smooth endosperm cell walls in higher magnification
(Glaux maritima). Fig. 11. Smooth endosperm cell walls and thick seed wall in Maesa rufescens. Figs. 7 and 8. Frame et al. 179
(S). Figs. 9 and 10. ~ t i h &
l Knudsen s.n. (S). Fig. 11. Wall s.n. (S). All scale bars = 0.1 mm.
Can. J. Bot. Downloaded from www.nrcresearchpress.com by UNIV CHICAGO on 02/16/13
For personal use only.
 Anderberg and St%hl
Theophrastaceae and Maesa differ from the myrsinaceous
pattern. Faure (1968) hypothesized a basic chromosome
number of x = 12 for the Myrsinaceae but x = 18 for the
Theophrastaceae and x = 10 for Maesa. The Primulaceae
show a great variability in chromosome numbers, but chro-
mosome numbers based on x = 10 are very common (e.g.,
Fedorov 1974).
Phytochemistry
In contrast with most other sympetalous plant groups, the
Primulales lack characteristic compounds like iridoids and
ellagic acid (Jensen et al. 1975; Cronquist 1981). Instead, the
Primulales are characterized by the occurrence of various
saponine compounds (Hegnauer 1969), and many genera
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also produce quinoid compounds that are known to be aller-
genic (Hausen 1978).
An interesting compound found in members of the Primu-
laceae is the flavonol gossypetin, which has a very restricted
distribution in flowering plants (Harborne 1969). This sub-
stance has proved to be of systematic value in the Ericaceae
(Anderberg 1992, 1993), and the occurrence of gossypetin in
these two families is interesting considering that they have
both been found to belong in the same major clade of flower-
ing plants (Chase et al. 1993). To date, gossypetin has only
I
been demonstrated to occur in the tribe Primuleae (Primula
I
spp., Dionysia, Douglasia), but an investigation of the distri-
bution of gossypetin from a wider sampleof taxa may reveal
For personal use only.
that it is also a useful taxonomic marker in the Primulaceae.
Whether or not gossypetin occurs in the Myrsinaceae or the
I
Theophrastaceae is as yet unknown.
Fossil record
Records from the Baltic amber (Conwenz 1886) include
flowers of the myrsinaceous genera Myrsinopsis Conwentz
and Berendtia Gopp. (Pax 1889; Mez 1902). Leaves of myr-
sinaceous plants were reported from the Tertiary floras of
Europe and Greenland, and these plant parts were assigned
to a number of different genera (Myrsine L., Myrsinites Ett.,
Ardisia Sw., Pleiomerites Ett., Icacorea Aubl., and Maesa;
e.g., Takhtajan and Uznadze 1963), but Menzel(1921), who
considered the identifications as more or less uncertain,
referred to them in a collective sense as Myrsinophyllum
Menzel. Fruits similar to those of the extant genus Ardisia
were recorded from the Eocene of England (Chandler 1964),
and fossilized pollen of the Myrsinaceae, similar to that of
Suttonia and Rapanea, is known from the Oligocene (Muller
1981). Other Tertiary fossils are inflorescences of the genus
Pleiomerites Ettinghausen, described from Germany (Collin-
son et al. 1993).
. ..... .,
Leaves from the Tertiary in Germany have been referred
to the Theophrastaceae (Clavijopsis, Schindehiitte 1907), but
although similar in shape and venation to leaves of Clavija,
the sparse and incomplete material makes the proposed rela-
tionship with that genus uncertain.
From the Primulaceae, seeds very similar to those of
extant Lysimachia vulgaris are known from the Oligocene
and Miocene (Friis 1985), and according to Muller (1981),
pollen of Samolus valerandi L. was reported from upper
Miocene sediments of Spain. Furthermore, seeds identified
as Androsace are known from the Siberian Miocene (Doro-
feev 1963), but other fossil Primulaceae, including Trientalis
and Primula, are apparently more uncertain (Kirchheimer
1957).
Judging from the fossil record, the Primulales are a rela-
tively young group, first appearing in the Tertiary, in the
Miocene and the Oligocene (about 30 000 000 BP). From
that period fossils are known from at least two of the three
families and in many respects they resemble extant taxa. The
fact that a group like the Myrsinaceae are found as fossils in
areas like Germany and England indicates that a change in
climate has occurred in this part of the world since the Mio-
cene and Oligocene, from a tropical-subtropical to a tem-
perate climate.
Both the putatively related families Ebenaceae and
Sapotacae are also known from the Tertiary, but according
to Menzel (1921), some fossils are as old as from the Upper
Cretaceous.
Circumscription of the Primulales and
their systematic position
In spite of an almost unanimous taxonomic consensus regard-
ing the. monophyly of the Primulales, no clear picture has
emerged concerning their closest relatives, although most
recent classifications seem to agree that the Ebenales are
among the more plausible candidates. Most genera of the
Ebenales, like those of the Primulales, have bitegmic tenui-
nucellate ovules and often have flowers with sympetalous
corollas. The Sapotaceae differ from the other members of
that order by their unitegmic ovules in combination with
tenuinucellus and nuclear endosperm formation (Cronquist
1981). The Sapotaceae also differ from the other families of
the Ebenales in having well-developed laticifers. On the
basis of these differences, Dahlgren (1991) suggested that the
systematic position of the Sapotaceae was in need of further
investigation.
In a recent paper, Geetha et al. (1993) adopted Hutchin-
son's view (Hutchinson 1969) that the Primulales are poly-
phyletic. In their view the Primulaceae would have originated
from the Violales and the two other families directly from the
Theales. This bold hypothesis was based merely on the dis-
tribution of simple phytochemical compounds known to be
highly homoplasious among flowering plants and without
any consideration of morphological data.
Cronquist (1981) included the families Ebenaceae, Sapota-
ceae, Symplocaceae, Styracaceae, and Lissocarpaceae in the
order Ebenales and placed them just next to the Primulales.
He stated that a common ancestor of such a group would
have features like those found in families of the Theales, to
which he referred the Actinidiaceae, for example.
Thorne (1983, 1992) also placed the Primulales and the
Ebenales next to each other, and his circumscription of the
latter differed very little from Cronquist's (1981) view. On
the other hand, Thorne (1983, 1992) had a unique view
regarding the circumscription of the Primulales, which he
subdivided into the suborders Primulineae and Plumbagi-
nineae, with the former including the traditional Primulales
families and the latter comprising the Plumbaginaceae. This
proposed relationship is further discussed below.
Takhtajan's (1987) superorder Ericanae included the orders
Ebenales (Styracaceae, Lissocarpaceae, and Ebenaceae),

Sapotales (Sapotaceae), and Primulales (Theophrastaceae,
Myrsinaceae, Aegicerataceae, and Primulaceae), as well as
the Ericales and Diapensiales. The Sapotales were in this
context placed next to the Primulales, and according to Cron-
quist (1981, p. 508), Takhtajan had earlier suggested a pos-
sible homology of the secretory system of the Myrsinaceae
with that (the laticifers) of the Sapotaceae.
In the last of Dahlgren's systematic schemes (Dahlgren
1989), the superorder Primulanae were composed of the two
orders Primulales and Ebenales. The former comprised the
Myrsinaceae, Theophrastaceae, and Primulaceae, as well as
the two segregate families Coridaceae (Coris) and Aegicer-
ataceae (Aegiceras). The Ebenales comprised the families
Sapotaceae, Styracaceae, Ebenaceae, and Lissocarpaceae.
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In a large analysis of molecular data from the chloroplast
DNA gene rbcL, Chase et al. (1993) corroborated the mono-
phyly of the Primulales. Only three genera from the Primu-
lales, one from each family, were included in the analysis,
but the analysis gave support for the Theophrastaceae (Cla-
vija) as the sister group of the Myrsinaceae (Ardisia) and the
Primulaceae (Anagallis). Unfortunately the analysis left the
question of the sister group of the Primulales open, but
the Primulales as well as the genera of the Ebenales were
found to belong to a larger monophyletic group of families
constituting part of an enlarged Asteridae (clade asterid 111).
This hypothesis obtained from molecular data is supported
by the presence in these families of a number of specialized
For personal use only.
morphological characters, such as unilacunary nodes, tenui-
nucellate ovules, and haplostemonous, sympetalous flowers.
The Primulaceae- Plumbaginaceae connection
The similarity in floral structure on which the supposition of
a close relationship between the Primulaceae and Plum-
baginaceae has been founded (cf. Pax 1889; Thorne 1983,
1992) was recently shown to be due to homoplasy (Clincke-
maillie and Smets 1992). This systematic relationship is fur-
ther contradicted by the fact that the Plumbaginaceae have a
micropyle formed only by the inner integument, and thus
also in this respect conform to the condition of the so-called
Centrospermae (Wunderlich 1967), whereas in the Primu-
lales, the micropyle is formed by both integuments (Johri
et al. 1992). Furthermore, the proposed relationship between
the Primulales and the Plumbaginaceae is at odds with the
molecular work by Chase et al. (1993) that places the Plum-
baginaceae near the traditional centrosperm groups, whereas
the Primulales are included among the sympetalous families
of the Asteridae. Another contradiction is presented by
Frohne and John (1978), who from an investigation of sero-
logical data in the Primulales and supposedly related groups
concluded that there is no support for a close relationship
between Plumbaginales and Primulales. On the other hand,
they found a clear serological correspondence between
Primulales and Ericales, which to some extent is echoed by
the results obtained by Chase et al. (1993) from analyses of
rbcL sequence data. It seems as though the correspondence
in serological data, at least in this case, reflects synapo-
morphic similarity between these groups.
The outgroup taxa
From the analysis by Chase et al. (1993), conclusions can be
drawn about a number of families that could be considered
Can. J . Bot. Vol. 73, 1995
suitable outgroups for an analysis of the Primulales. There
is, however, an important difference between the two topolo-
gies presented for their clade asterid 111. The first scenario
(13A, p. 574) has the families Ebenaceae, Diapensiaceae,
and Polemoniaceae as the nearest relatives of the Primulales,
followed by a wider group of families including, e.g., the
Sapotaceae. The second scenario (13B, p. 575) has the Styra-
caceae and Clethraceae as the closest relatives of the Primu-
lales, with families like the Actinidiaceae, Theaceae,
Cyrillaceae, and Sapotaceae in the close vicinity. Families
like Diapensiaceae, Polemoniaceae, and the Ebenaceae
were, in this scenario, found to be more distantly related to
the Primulales.
In the light of available morphological data, we have con-
cluded that neither the Clethraceae, the Diapensiaceae, nor
the Polemoniaceae are suitable as outgroup taxa for an analy-
sis of phylogenetic interrelationshipsin the Primulales. Instead
we hold to the traditional concept of the Ebenales as a possi-
ble candidate as a sister group to the Primulales, but the dis-
crepancy between the two trees of Chase et al. (1993)
regarding the position of the different families of the Eben-
ales in relation to the Primulales cannot be overlooked. It
must be noted that in neither of the two trees do the Ebenales
appear together as a monophyletic group, and the only family
within the Ebenales that consistently is placed very close,
and basal in relation to the Primulales, are the Sapotaceae.
Because of this uncertainty, we have included representatives
from two families of the Ebenales in the outgroup, as well
as a representative of a group further outside of the
Ebenales - Primulales sphere.
Two genera of the Sapotaceae, Manilkara Adans. and
Monotheca A.DC. (Reptonia A.DC., Edgeworthia Falc.)
were included in the outgroup. Manilkara was chosen mainly
because it was included in the rbcL analysis by Chase et al.
(1993). The second representative, Monotheca, is a mono-
typic genus of northeastern Africa, Arabia, Afghanistan, and
northwestern Pakistan. The genus was once included in the
Theophrastaceae (de Candolle 1844; Hooker 18766; Pax
1889) but was transferred to the Sapotaceae by Radlkofer
(1889) where it has remained ever since (Mez 1903; Stewart
1972; Friis 1978; Malik 1984; Collenette 1985; Stihl
1990~).Because of its former systematic position in the
Primulales, we have found it particularly interesting to
include Monotheca in our analyses.
Apart from these two genera we included Diospyros from
the Ebenaceae to cover more of the character variation in the
Ebenales. This genus was also included in the molecular
study by Chase et al. (1993).
As mentioned above, Chase et al. (1993) consistently
found the families of the Ericales to be associated to the same
clade asterid 111 as the Primulales, and therefore, we have
included also Actinidia Lindley (Actinidiaceae) among the
outgroup taxa. In a number of recent papers, this family was
found to be the basal clade of the Ericales (Anderberg 1993;
Judd and Kron 1993; Kron and Chase 1993), and the choice
of Actinidia also meets the morphological requirements for
a hypothetical ancestor of the Ebenales-Primulales group as
hypothesized by Cronquist (1981, p. 509).
The ingroup taxa
To cover as large a part of the morphological variation as
 Anderberg and St%hl
possible, representatives (genera, subgenera, or species)
included in the ingroup were selected from all currently and
previously recognized suprageneric taxonomic groups. It
was not our main objective to elucidate the interrelationships
within each major group in detail but rather to let the sampled
taxa indicate the delimitation of the major clades in the
Primulales. The taxa included in the analyses are presented
here briefly, together with additional notes that give the
rationale for omitting some genera from the analyses.
Theophrastaceae Link
Theophrasta L. has two closely related species endemic to
the island of Hispaniola (Stihl 1987). Neomezia Votsch is a
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monotypic genus endemic to Cuba, which is similar to Theo-
phrasta and probably the sister group of that genus (Stihl
1990a). Theophrasta was included in our analyses.
Clavija Ruiz & Pavdn is a coherent genus of 50 species
in South and Central America with one species in the West
Indies (Stihl 1991).
Jacquinia L., a genus of about 35 species in the West
Indies, Mexico, Central America, and coastal South
America, is clearly the most heterogeneous genus in the
family. Deherainia Decne. has two species in Mexico and
northern Central America and may be the sister group of
Jacquinia (Stihl 1989, 1990a, 1995). Votschia Stihl is a
monotypic and poorly known genus from Panama that is
morphologically intermediate between Jacquinia and Deher-
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ainia (Stihl 1993). From this group, only Jacquinia was
included in our analyses.
Myrsinaceae R.Br.
The Myrsinaceae have the reputation of being taxonomically
difficult with poorly defined genera and many variable and
sexually heteromorphic species. The delimitation of many
genera is still highly suspect and the classification of the
family is certainly in need of a thorough revision. In our
present study, we have not had the ambition to elucidate the
phylogenetic status of every genus recognized at one time or
the other but have chosen to include a wide sample of taxa
to test the monophyly of the Myrsinaceae. For that purpose,
we believe that the present sample is sufficient.
Aegiceras Gaertn. has two species confined to mangrove
vegetation in southeast Asia and tropical Australia. It has
several distinctive and odd features, and therefore it has often
been placed in a separate family, the Aegicerataceae (de Can-
dolle 1844; Hutchinson 1969; Takhtajan 1987; Dahlgren
1989).
Amblyanthus A.DC., with three species in the eastern
Himalayas, is diagnosed by its connate anthers (but not fila-
ments) and emarginate corolla lobes (de Candolle 184la;
Mez 1902). Amblyanthopsis Mez, also with two species in
the Himalayan region (Stone 1991a), is very similar to
Amblyanthus, according to Mez (1902), and differs only in
having free stamens. From this group, only Amblyanthus was
included in our analyses.
Ardisia Sw. is, in its widest sense, a pantropical genus of
at least 300 species and thus the largest in the family. It was
subdivided by Mez (1902) into 14 subgenera, a classification
followed by most subsequent workers in Asia (Pitard 1930;
Walker 1940; Stone 1982, 1989, 1990, 1993a; Sleumer
1 9 8 8 ~Larsen
; and Hu 1991). From the New World, Lundell
(1964, 1966a, 1971, 1981a, 1981b, 1982) described or recog-
nized several segregate genera, many of which are partly or
entirely based on the subgenera recognized by Mez (1902),
viz. Amatlania Lundell, Auricularardisia Lundell, Chonta-
lesia Lundell, Gentlea Lundell, Graphardisia (Mez) Lun-
dell, Icacorea Aublet, Ibarraea Lundell, Oerstedianthus
Lundell, Valerioanthus Lundell, and Zunilia Lundell. In addi-
tion to these, there are several genera that differ from Ardisia
in some characters but can still be hypothesized to be rather
closely related. We refer to these as an informal Ardisia com-
plex without any a priori assumptions about their phylo-
genetic status. Listed below are a number of these genera that
were all omitted from the analyses. Afrardisia was described
by Mez (1902) to accommodate the African species, which
differ from the main stock by having few ovules arranged in
a single series. It was recognized as a separate genus by
de Wit (1958) but has been included in Ardisia s.lat. by
others (Taton 1980; Halliday 1984). Parardisia Nayar and
Giri (1986), based on two species from Assam and Borneo,
was recently added to the list of synonyms of Ardisia (Stone
1993a). Antistrophe A.DC., with five species in the Indo-
malayan region, is similar to Ardisia s.lat. but with a corolla
twisted to the left and ovules situated in a single series. De
Candolle (1841a) distinguished Antistrophe from Ardisia on
the basis of its anthers ending in a membranous appendix.
Ctenardisia Ducke, with two species in northeastern Brazil,
is distinguished by large oblanceolate leaves, elongated pani-
cles, and erect ovules arranged in a single series around the
base of the placenta (Lundell 1982). The Mesoamerican
genus Yunckeria Lundell, with one to three species in Central
America, has a similar ovule arrangement and was therefore
merged with Ctenardisia by Lundell (1982). Hymenandra
(A.DC.) Spach., with eight species in southeast Asia, is
much like Ardisia but has anthers laterally connate and
ovules in a single series (two in Hymenandra wallichii
A.DC. ; Stone 199Ib). Tetrardisia Mez, with three species in
Java and Queensland, is basically an Ardisia with tetramer-
ous flowers and few ovules in a single series. Systellantha
Stone, with two species from Borneo, has many features in
common with Tetrardisia but differs mainly in having
unisexual flowers and short condensed inflorescences (Stone
1992). Solonia Urban (syn. Walleniella cubana P. Wilson;
Alain 1957), with a single species in Cuba, seems to be an
apomorphic segregate from Ardisia s.lat., with the anther
filaments united into a tube that is largely free from the
corolla. Stylogyne A.DC., a genus of 50 species occurring in
tropical America, is similar to Ardisia s.lat. but often has
unisexual flowers, few ovules and axillary inflorescences
unlike most neotropical Ardisia. The dense lateral venation
and glandular punctation of the leaves may also help to dis-
tinguish the genus from Ardisia, at least in the field (Gentry
1993). A cladistic analysis of the Myrsinaceae with represen-
tatives of all the segregate genera is needed before a better
understanding of the phylogenetic status of the Ardisia com-
plex can be gained. For the present analyses, we have
included two species of Ardisia: Ardisia elliptica Thunb.,
which like Aegiceras has septate anthers, and Ardisia pusilla
A.DC., a species belonging to the mainly suffrutescent
subgenus Bladhia (Thunb.) Mez (Yang and Dwyer 1989).
Cybianthus Martius was treated in a rather narrow sense
by Mez (1902) who included in the genus 36 species from
South America. Agostini (1980) introduced a much broader

concept by including the five neotropical genera Comomyr-
sine Hook.f., Conomorpha A.DC., Corelliana D'Arcy,
Cybianthopsis (Mez) Lundell, and Weigeltia A. DC. and
divided the genus into 10 subgenera. This classification was
adopted by Pipoly (1983, 1987, 1992a) who added an addi-
tional subgenus based on the genus Grarnmadenia Bentham
with seven species in tropical America. In this broad sense
Cybianthus consists of about 150 species in the neotropics. In
our analyses we have included - ~ o n o m o r ~ hA.DC.
a and
Grarnmadenia Benth. (Cybianthus subg. Conomorpha (A.DC.)
Agostini; Cybianthus subg. Grarnmadenia (Benth.) Pipoly).
Embelia Burm.f., with about 130 species in the Old
World tropics, differs from most other members of the
Primulales in its choripetalous flowers, and in addition,
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many species are scandent. Grenacheria Mez, with 10 spe-
cies in the Indomalayan region, is apparently a close relative
of Embelia from which it differs (solely?) by having sym-
petalous flowers. We have included Embelia ribes Burm.f.
in our analyses.
Heberdenia Banks ex A.DC., a monotypic genus of
Macaronesia, resembles Embelia in many characters but dif-
-
fers in its bisexual flowers arranged in subumbellate inflores-
cences and by having ovules in more than one series. A
second, Mexican species, Heberdenia penduliflora (A.DC.)
Mez, was referred to the genus by Mez (1902), but this spe-
cies certainly does not belong there (B. Stihl, unpublished
data). Pleiomeris A.DC. is a monotypic, Macaronesian
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genus, sometimes associated with the Myrsine complex
because of its fasciculate inflorescences and allegedly pau-
ciovulate ovaries (Mez 1902; de Wit 1957; Pipoly 1992b),
but as pointed out by de Wit (1957) and earlier by de Can-
dolle (1844), the ovules in this genus are many and arranged
in two or three series. It is possibly closer to Heberdenia or
Embelia, having granulate petals of the same shape and tex-
ture as these genera but differing in its well developed fila-
ment tube. We have included Heberdenia in our analyses.
Labisia Lindley, with about six species in the Malesian
region (Stone 1988a), is seemingly well diagnosed by
suffrutescent growth habit, leaves with dense lateral venation
and decurrent blade bases, and induplicate corolla lobe mar-
gins. The monotypic genus Emblemantha Stone, described
from Sumatra (Stone 1988b), is evidently a close relative of
Labisia but has an almost tubular corolla. Sadiria Mez, with
six species in the eastern Himalayas, also seems to belong to
this informal group. All three genera have long-apiculate
anthers that are, at least in Labisia and Sadiria, somewhat
curved inwards. We have included Labisia in our analyses.
Maesa Forssk., with ca. 100 species in the Old World,
mainly in the tropics, is distinguished from all other Myr-
sinaceae by its perigynous flowers, many-seeded fruits, and
the pedicels of each flower being supported by a pair of
opposite or subopposite bracteoles. Several micromorpho-
logical, anatomical, and cytological features also seem to set
it apart from other Myrsinaceae (see text for further discus-
sion). A small number of African species have ovules
arranged in only one series and belong to a separate subgenus
(Monotaxis Mez). We have included both subgenera in our
analyses .
Myrsine L. includes in its most narrow sense (Mez 1902;
Taton 1980; Halliday 1984) about seven species and has a
wide distribution in the Old World tropics. However, many
Can. J . Bot. Vol. 73, 1995
workers have given the genus a wider circumscription,
including the large pantropical genus Rapanea Aubl. (King
and Gamble 1905; Perrier de la BIthie 1953; Walker 1959;
Backer and Bakhuizen van der Brink 1965; Stearn 1969;
Liogier 1971; Pipoly 1992b), and sometimes also the Pacific
genus Suttonia A. Rich. (Hosaka 1940). In this wide sense,
Myrsine comprises 150 species, i.e., virtually all Myr-
sinaceae with fasciculate inflorescences and pauciovulate
placentas. In a revision of the Myrsinaceae of the Fijian
region, Smith (1973) merged Suttonia with Rapanea,
whereas Myrsine s.str., which does not occur in that area,
was considered a distinct genus. Nakai (1943) considered
Rapanea to be a New World genus and treated the eastern
Asiatic (mainly Japanese) representatives in three distinct
genera (Anamtia Koidzumi, Athruphyllu~nLour., and Man-
glilla Juss.) distinguished mainly by differences in stigma
morphology, but this classification was later questioned by
Walker (1954). Myrsine s.str. can be distinguished from
Rapanea and Suttonia by its possession of a well-developed
stamina1 tube. Given the uncertainties regarding the relation-
ships within the Myrsine complex, a single species, Myrsine
africana L., was included in the analyses.
Parathesis (A.DC.) Hook.f., with about 70 species in the
neotropics (Lundell 1966b), is easily recognized because of
its usually brownish indumentum of stellate or dendroid
hairs, wide paniculate inflorescences, and narrow, revolute
petals.
Tapeinosperma Hook.f., with some 40 species in New
Guinea, Fiji, and Queensland, is variable and difficult to
define but is characterized by rather showy inflorescences
with caducous bracts, open solid flowers, subhastate-
triangular, sessile or subsessile anthers, rather short style,
and deeply immersed ovules arranged in a single series. The
leaves are medium-sized to very large (Stone and Whitmore
1970) and often pseudoverticillate, and some species have a
pachycaulous growth habit. Several, primarily Asiatic -
Pacific genera share most of these features and for con-
venience are treated here as an informal group under
Tapeinosperma. Discocalyx (A.DC.) Mez, with 50 species
in eastern Malaysia and Polynesia, is close to and perhaps
impossible to keep separate from Tapeinosperma, from
which it ideally differs in unisexual flowers, very short style,
and sessile or subsessile anthers (Smith 1973). Fittingia Mez
with five species in New Guinea (Sleumer 1 9 8 8 ~ is ) similar
to Discocalyx and distinguished by its fruits that have thick
exocarp, which is longitudinally ridged or tubercled (Stone
1991~).Loheria Merr. with six species in the eastern Male-
sian region (Stone 1991a) is a dioecious or functionally dio-
ecious genus characterized by a pachycaulous growth habit
with reduced lateral shoots and a combination of floral fea-
tures that taken together may set it apart from the seemingly
related genera of this group. We have included the species
Tapeinosperma clavatatum Mez in the analyses.
Oncostemum A. Juss., with about 100 species in Mada-
gascar and the Mascarenes (Perrier de la Biithie 1953), is
diagnosed by its tubular androecium, formed by the fila-
ments and most of the anthers. Badula Juss., with 14 species
in the same area, is similar to Oncostemum but has free
anthers, although this distinction is not valid for all species
(Coode 1976); some species are pachycaulous. Monoporus
A.DC. has eight species in Madagascar (Perrier de la BIthie
 Anderberg and Stahl
1953) and is diagnosed by poricidal anthers and erect ovules
inserted at the base of the placenta but is probably a close
relative of Oncosternum. Oncosternurn was included in our
analyses .
Conandrium (K. Schum.) Mez has two Indomalayan spe-
cies (Sleumer 1988b), and because of its connate anthers,
Mez (1902) related this genus to Hymenandra, although the
large differences in inflorescence architecture and floral
morphology indicate that they are not particularly close. The
genus was said (Mez 1902) to have a many-ovuled placenta
with ovules in at least two series, but in the material of
Conandriurn polyanthum (Laut. & K. Schum.) Mez exam-
ined by us (Bergman 440, Brass 28574), only pauciovulate
placentas with deeply immersed ovules in a single series
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were observed. Sleumer (1988b) indicated that only the con-
nate anthers distinguished Conandrium from Ardisia, but he
obviously never examined the ovary.
Wallenia Sw., with 25 species in the West Indies (Stearn
1969), is characterized by coriaceous, sometimes pseudo-
verticillate leaves, usually unisexual flowers, well-developed
corolla tube, exserted stamens, and versatile anthers. This
genus is divided (Mez 1902) into subg. Wallenia, with termi-
nal paniculate inflorescences, and subg. Homowallenia Mez,
with axillary racemose inflorescences. Geissanthus Hook.f.,
with 35 species in tropical South America (chiefly in the
Andes), is diagnosed by recurved petals and by its calyx,
which is closed in bud and opens irregularly into 2 -7 lobes
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(Pipoly 1993), but is otherwise similar to Wallenia. Vegaea
Urban, with one species in Hispaniola, was considered by
Urban (1913) to be a relative of Wallenia but with shorter
stamens and bisexual flowers. We have included Wallenia
subg. Wallenia in our analyses.
Elingamita Baylis is a monotypic tree genus from New
Zealand with entire coriaceous leaves, a tubular corolla that
is shorter than the calyx, few ovules, and red drupaceous,
one-seeded fruit. We have seen no material of this genus, but
it seems to be a typical representative of the Myrsinaceae.
Primulaceae Vent.
Coris L. has two species in the Mediterranean area and in
Somalia, and is diagnosed by its zygomorphic corolla, for
example. The two species are very similar to each other.
Samolus L. has 15 species, occurring especially in salt
marshes. The distribution is worldwide, but most species are
found in the southern Hemisphere. Sarnolus is probably
monophyletic and is diagnosed by its perigynous flowers.
Lysimachia L. is a genus of ca. 150 species, which has a
worldwide distribution. It is heterogeneous and presumably
paraphyletic because other genera of the tribe Lysimachieae,
e.g., Anagallis, Trientalis, the apetalous Glaw, and Astero-
linon, appear to be apomorphic ingroups. A revised circum-
scription of Lysirnachia seems inevitable, but further studies
are needed before a detailed and well-corroborated hypothe-
sis can be formulated regarding the phylogeny of the group.
To account for the considerable variation within Lysirnachia,
we have included four representatives in the analyses, viz.
Lysirnachia insignis Hemsl. (subg. Idiophyton Hand.-Mazz.),
Lysirnachia ciliata L. (subg. Seleucia Bigelow), Lysimachia
glutinosa Rock (subg. Lysirnachiopsis (Hell.) Hand.-Mazz.),
and Lysimachia clethroides Duby (subg. Palladia (Moench)
Hand.-Mazz.).
Trientalis L. has two species in northern temperate areas.
This genus is probably an apomorphic segregate from the
paraphyletic Lysimachia complex. There are some species of
Lysimachia, e.g., Lysimachia evalvis Wall., which have
fruits that much resemble those of Trientalis. Also, several
species of Lysimachia (e.g., Lysimachia paridijormis Franch.
and Lysimachia sciadophylla Chen & Hu), have pseudover-
ticillate leaves like Trientalis. Both species of Trientalis are
pseudoannuals, propagating with rhizomes that end in a bulb-
like winterbud (Warming 1918; Anderson and Loucks 1973;
Anderson and Beare 1983), and such a life cycle is known
to occur also in Lysimachia thyrsiflora L. and in Glaux
(Warming 1918; Jerling 1988). The often heptamerous
corolla of Trientalis is an autapomorphy.
Asterolinon Hoffsgg. & Link. has two species, one in the
Mediterranean region, the other in Ethiopia. The type was
described as a species of Lysimachia by Linnaeus (1753) and
was also included in that genus by Leblebici (1978). Within
the Lysimachia complex, Asterolinon linum-stellatum is prob-
ably closely related to Pelletiera A. St.-Hil., but the second
Asterolinon species has the habit of an Anagallis and may
prpve to be more distantly related. Pelletiera A. St.-Hil. has
two species, one in South America and the other in Macaro-
nesia, which much resemble Asterolinon linum-stellaturn.
These three species share the annual habit, the minute white
petals, the few-seeded capsules, and the irregularly rugose
seeds with a deeply sunken hilum area. As a representative
of this group, we have included Asterolinon linurn-stellatum
in our analyses.
Glaux L. is a monotypic genus occurring in northern tem-
perate areas. Its apetalous species has several features in
common with members of the Lysinzachia complex. Glaux
propagates with rhizomes ending with a bulblike winterbud,
and like Trientalis it also acts like a pseudoannual in a similar
maner (Jerling 1988). Owing to the lack of corolla, many
character states are unknown in the genus, and therefore an
analysis of DNA sequence data will be particularly useful in
finding its proper systematic position. It seems likely, how-
ever, that it belongs in the Lysimachia complex.
Anagallis L. (including Centunculus L.), with ca. 20 spe-
cies, occurs in Africa, South America, and Europe. In most
respects Anagallis agrees with Lysimachia. The salient fea-
ture used to distinguish the genus is the operculate capsules,
but this distinction seems artificial. There is a striking overall
similarity between species of Anagallis to Lysimachia serpyl-
lijolia Schreb., and representatives of Lysirnachia subg. Pal-
ladia, for example. Anagallis may be a monophyletic
ingroup in Lysimachia, but further work is needed to clarify
the relationships.
Androsace L. has more than 150 species found mainly in
northern temperate areas. Androsace is typically recognized
by small flowers with a constricted corolla throat. Most spe-
cies are cushion-forming, but some have longer internodes.
Some species have a Primula-like habit and the relationships
to that genus still have to be investigated further. Richards
(1993) considered Androsace to be primitive in relation to
Primula, whereas Smith and Lowe (1977) believed that
Androsace had evolved from Primula. Stimpsonia C. Wright
ex A. Gray, with one species in eastern Asia, is probably a
derived relative of Androsace (Wendelbo 1961c; RGsvik
1969). Pomatosace Maxim., with one species in the

Himalayas, is traditionally placed in the ~rimuleae-
' resemblance to some Androsace of sect. Pseudoprimula Pax,
Soldanellinae because of its operculate capsules, but all other
characters indicate that it belongs near Androsace. Andro-
sace maxima L. (sect. Andraspis (Duby) Koch) and Andro-
sace obtusifolia All. (sect. Chamaejasme Koch) were
included as representatives of Androsace.
Douglasia Lindley has eight species in mountains of
the U.S.A. and Canada. It is rather homogeneous, but the
characters presented by Kelso (1991) as diagnostic of the
genus are also found in some Androsace, in which it was
included by Wendelbo (1961~).Vitaliana Sesler, with one
species in the European Alps, is usually considered a con-
gener of Douglasia (Pax and Knuth 1905; Constance 1938),
but this relationship needs further study. Douglasia nivalis
Can. J. Bot. Downloaded from www.nrcresearchpress.com by UNIV CHICAGO on 02/16/13
was included in our analyses.
Dionysia Fenzl. (Wendelbo 1961a; Grey-Wilson 1989),
with 39 species occurring in the mountains of central Asia,
is a monophyletic group diagnosed by its very long tubular
flowers. Dionysia is often considered to be closely related to
Primula subg. Sphondylia (Duby) Rupr. (Wendelbo 1961b;
but see Richards 1993).
Primula L., with ca. 400 species, mainly in the northern
Hemisphere, is heterogeneous and probably paraphyletic
(Smith et al. 1977; Richards 1993). Wendelbo (1961~)con-
sidered it likely that the genera Sredinskya, Hottonia, Dio-
nysia, Cortusa, and Dodecatheon all have their closest
relatives within Primula, and Richards (1993) added Ompha-
For personal use only.
logramma and Bryocalpum to that list. If these phylogenetic
schemes corresponds to the true phylogeny of Prirnula and
the other genera of the Primuleae, it would mean that drastic
changes in taxonomy are necessary if paraphyletic groups are
to be abandoned. Sredinskya (Stein.) Fed. is probably an
autapomorphic member of the Primula complex, and the
same is probably true of Omphalogramma (Franch.)
Franch., which is diagnosed by solitary, ebracteate flowers.
There are several Primula species that in habit appear more
or less intermediate between the two genera. The genus
Bryocarpum H0ok.f. & Thoms., with one species in eastern
Himalaya, is either the sister group of Omphalogramma or
more likely an apomorphic ingroup therein (cf. Fletcher
1949, p. 127; Richards 1993, p. 41). Hottonia L. has two
species (one in North America, one in Europe and Asia) that
show distinct affinities to Primula. The flowers in Hottonia
are heterostylous and arranged in several whorls, with the
ultimate whorl being a bracteate umbel, and hence the
inflorescence is structurally the same as the superimposed
umbels of many Primula subg. Aleuritia (Duby) Wendelbo
sect. Proliferae Pax. We have included two species of Prim-
ula in our analyses, viz. Primula veris L. (subg. Primula)
and Primula palinuri Petagna (subg. Auriculastrum).
Cortusa L., with eight species, in the mountains of
Europe and Asia, is probably paraphyletic if the yellow-
flowered Kaujkannia Regel, with one or two species in cen-
tral Asia, is retained as a separate genus.
Ardisiandra H0ok.f. has three species in the African
mountains. This genus has earlier been associated with Cor-
tusa (e.g., Pax 1889), but this opinion was not shared by
ROsvik (1968, 1969), who considered it an isolated genus in
the family and placed it in a separate tribe, the Ardisian-
dreae. In general aspect, Ardisiandra show a remarkable
Can. J. Bot. Vol. 73, 1995
e.g., Androsace axillaris Franch.
Soldanella L., with 10 species in the European Alps, is a
small and monophyletic, very homogeneous group but with
unclear relationships to other genera. The capsule opens by
splitting of the style base but is often said to be operculate
(e.g., Pax and Knuth 1905). This opening mechanism is
probably not homologous to the circumscissile and distinctly
operculate lids of Anagallis.
Cyclamen L. (Grey-Wilson 1988; Anderberg 1994) has
19 species mainly distributed around the Mediterranean.
This is the most distinctive genus in the Primulaceae, and its
monophyly can hardly be questioned. The systematic posi-
tion of Cyclamen was discussed by ROsvik (1966), who
found it isolated in the family.
Dodecatheon L. has 14 species occurring mainly in
eastern North America. Three subgenera were recognized by
Thompson (1953), each with a rather different set of charac-
ters, but the genus as such is probably monophyletic. Apart
from their reflexed corolla lobes, the species of Dodecatheon
much resemble Primula species (e.g., Primula parryi A.
Gray), and Richards (1993) believed the genus to have
evolved from within the Primula stock.
Materials and methods
The present study is based on the investigation of herbarium
specimens housed in the herbaria of AAU, BM, GB, K, S,
and SUNIV, supplemented by information from the litera-
ture, particularly regarding anatomical and embryological
data. Living material in greenhouses, and in the field, was
also investigated.
Cladistic analyses
The data matrix in Table 1 was computed on a 486 PC work-
ing at 66 MHz, using the parsimony program H E N N I G ~ ~
(Farris 1988). The analyses included the genera Actinidia
(Actinidiaceae), Diospyros (Ebenaceae), and the two genera
Manilkara and Monotheca (Sapotaceae) as outgroup for
assessing character polarity and for rooting the cladogram.
All characters were coded as nonadditive except characters
29 and 32, which were coded as additive transformation ser-
ies, with character states 2 and 3 being subsets of states 1 and
2, respectively. The following options were used for the
analyses: multiple HENNIG (MH*,constructing several initial
cladograms by adding taxa in several different sequences,
retaining the shortest cladogram of each), with a subsequent
branch breaker (BB*,generating multiple equally parsimoni-
ous cladograms). The H E N N I G ~successive
~ weighting option
(XSW) was used to evaluate the effect of highly homoplasious
characters on the tree topology (Farris 1969; Carpenter
1988). Strict consensus trees were computed with the NEL-
SEN command, and used for the purpose of the discussion
following the recommendations of Anderberg and Tehler
(1990). Character distributions were analyzed in the H E N N I G ~ ~
DOS EQUIS mode (xx), as well as with the CLADOS program
(Nixon 1992).
The measure of skewness (Hillis 1991), which sometimes
has been used as significance tests of cladistic analysis, was
recently demonstrated to be unreliable for such purposes
 Anderberg and St%hl
Fig. 12. One of 36 equally most parsimonious cladograms obtained from the analysis of the data in Table 1.
(316 steps, CI = 0.35, RI = 0.67). The diagnostic characters are discussed in the text.
Can. J. Bot. Downloaded from www.nrcresearchpress.com by UNIV CHICAGO on 02/16/13
For personal use only.
(Kallersjo et al. 1992). We have instead employed a permu-
tation test (KARA) written by J.S. Farris, to determine
whether the obtained results differed significantly from ran-
domness. The theoretical background and the test as such
was presented by Kallersjo et al. (1992).
Results
The cladistic analysis of the data in Table 1, with the four
genera Actinidia, Diospyros, Manilkara, and Monotheca as
outgroup, gave 36 equally parsimonious cladograms, each
316 steps long with a consistency index (CI) of 0.35 and a
retention index (RI) of 0.67 (Farris 1989). One of the clado-
grams from this analysis is shown in Fig. 12, and the cor-
responding character distribution is discussed below. A strict
consensus tree is shown in Fig. 13.
Five turns of successive weighting gave six equally par-
Oncostemum
simonious cladograms. A strict consensus tree is shown in
Fig. 14.
In a permutation test with the -KARA program, 10 000 per-
mutations showed the data to be significantly nonrandom as
the a,' value for the calculated length was 1 x The
permutation analysis yielded a total support for the real data
of 27, a maximum support of 3. The value of a,' for total
support was 58.8 x lop2, and for maximum support a,'
was 44.6 x lop2.
Discussion
The tropical and subtropical woody representatives in Theo-
phrastaceae and Myrsinaceae were found to be plesio-
morphic in relation to the temperate Primulaceae. This
supports the general conclusion of Judd et al. (1994) that
temperate representatives of tropical and subtropical groups
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For personal use only.
 Anderberg and St2hl
Can. J. Bot. Downloaded from www.nrcresearchpress.com by UNIV CHICAGO on 02/16/13
For personal use only.
In our analyses the Theophrastaceae were diagnosed by char-
acters such as pseudoverticillate leaves with extraxylary
fibres in layers or bundles, anthers initially forming a cone
but later become spreading, upper and lower part of anther
thecae provided with conspicuous clusters of calcium oxalate
crystals, ovules in two or more series, and endosperm with
pitted cell walls. Extrorse anthers are symplesiomorphic in
the family and therefore not diagnostic of it as a monophy-
letic group.
Staminodial flowers are also a symplesiomorphy in the
Theophrastaceae, since flowers lacking staminodes are apo-
morphic in the order. In a few taxa, such as Samolus and Sol-
danella, staminodial structures have evolved in parallel.
The majority of genera in the Primulales belong to a large
monophyletic group consisting of the families Myrsinaceae
and Primulaceae. This clade evolved from ancestors pro-
1719
vided with the following set of characters: leaves with secre-
tory cell cavities, flowers without staminodes, anthers
introrse and longitudinally dehiscent, ovules immersed into
the placenta, and cotyledons not foliaceous in seed. In some
groups within this major clade, the diagnostic features have
reversed in the course of evolution.
Considering that both the Theophrastaceae and Myr-
sinaceae are woody, we can be fairly confident that trees and
shrubs have evolved into herbaceous taxa within the Primu-
lales, and that some shrubby representatives (e.g., Coris,
Dionysia, some Lysimachia) have evolved secondarily from
herbaceous ancestors on more than one occasion. The oppo-
site trend can be seen in the Myrsinaceae, in which a few taxa
have evolved from shrubs into subshrubs, e.g., Ardisia subg.
Bladhia, and it can also be noted that the pachycaulous
growth habit found in Clavija and Theophrasta has evolved
 1720 Can. J. Bot. Vol. 73. 1995

Fig. 13. Strict consensus of 36 equally most parsimonious cladograms obtained from the analysis of the
data in Table 1.
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Primula veris

Androsace maxima
For personal use only.

Androsace obtusifolia

independently from that of the pachycaulous Myrsinaceae
(e.g., in Tapeinosperma). The Myrsinaceae and Theophras-
taceae generally have multiseriate wood rays only, but the
wood of the lignified Hawaiian species of Lysimachia is com-
pletely devoid of rays (Carlquist 1992), and this could sup-
port the hypothesis that these species have evolved in the
islands secondarily from herbaceous ancestors. Such an evo-
lution of a frutescent growth habit in otherwise herbaceous
plants is paralleled by several other Hawaiian plant groups,
e.g., many Asteraceae.
Within the Myrsinaceae - Primulaceae clade, a second
major monophyletic group is constituted by the Myrsinaceae
s.str., a group that may be hypothesized to have evolved
from ancestors with diagnostic characters such as the
presence of brown scale-like trichomes, drupaceous and
more or less fleshy, one-seeded fruits, ovules well spaced on
placenta, and subglobose seeds with depressed hilum area.
The relationships within the Myrsinaceae s.str. are not
well resolved, with the exception of Myrsine africana, which
is consistently placed as the sister group to the remaining
Myrsinaceae (except Maesa). Somewhat surprisingly, this
means that the pauciovulate condition can be viewed as
ancestral in the family, whereas multiovulate ovaries have
evolved several times independently. In this context it must
be noted that Mez' (1902) classification of the family into a
pauciovulate and a multiovulate group is not supported by the
present analyses.
Aegiceras is nested among other genera within the Myr-
sinaceae and linked to them by the following features: leaves
with crystal druses, flowers arranged in pseudoumbels,
bracts caducous, corolla with twisted aestivation, petals lan-
ceolate to ovate, longer than wide, anther connective with
secretory cavities, punctate or truncate stigma, and drupa-
ceous one-seeded fruit. There are also a number of charac-
ters or character states that were found to appear as
parallelisms, reversals, or autapomorphies in Aegiceras:
 Anderberg and St3hl
Fig. 14. Strict consensus of six equally most parsimonious cladograms obtained after five turns of successive
weighting.
Oncostemurn
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For personal use only.
presence of foliar sclereids, terminal inflorescence, strongly
reflexed petals, flowers with nectar-producing trichomes,
versatile anthers with glandular filaments, septate anther the-
cae, very elongated fruit, unitegmic ovules in many series,
densely set on the placenta, and exalbuminous seeds. It is
most parsimonious to hypothesize that although seeds of
Aegiceras are exalbuminous, the ancestor had subglobose
seeds with a depressed hilum area and pitted endosperm cell
walls. Aegiceras is highly apomorphic, but there is no sup-
port for the idea that Aegiceras should be treated as a
separate family as proposed by de Candolle (1844), Hutchin-
son (1969), Takhtajan (1987), and Dahlgren (1989).
The third major monophyletic group in the Primulales is
formed by the the myrsinaceous genus Maesa together with
the Primulaceae. This group can be hypothesized to have
evolved from ancestors provided with character states such
as leaves without immersed glandular hairs, flowers with
somatic gynoecium nectaries, anthers attached at about the
middle of the corolla tube or higher, anthers not sagittate,
ovules in two or more series, seeds sectoroid without
depressed hilum area, testa with rhomboid crystals, and
inner integument with a tapetum.
Within this clade, Maesa is itself monophyletic and diag-
nosed by variously serrate leaf margins, presence of pris-
matic foliar crystal druses, pedicel provided with a pair of
bracteoles subtending the flowers, perigynous flowers, fruit
formed by the ovary and the surrounding tissue, and an
embryogeny of the onagrad type.
The Primulaceae, the sister group of Maesa, are diag-
nosed by the following set of characters: herbaceous growth
habit, terminal inflorescence, thin delicate corolla texture,
synsepalous, often cup-shaped calyx, dry capsular fruit
opening with teeth or valves, ovules not immersed in the
placenta, and an embryology of the caryophyllad type.
In the initial, unweighted, analysis, Sarnolus, which also
constitutes the monogeneric tribe Sarnoleae of Pax and Knuth

(1905), was found to be the sister group of all other
Primulaceae. The perigynous flowers in Maesa and Samolus
could be interpreted as parallelisms, but an equally par-
simonious interpretation is to assume that perigyny once
evolved in the ancestor of the Maesa - Primulaceae clade and
transformed back into hypogyny above the Samolus node
(Fig. 12). After successive weighting, the position of Samo-
lus changes in the topology to a position higher up in the
cladogram, near the genera of the tribe Primuleae (Fig. 14),
and this may support the notion that perigyny has actually
evolved in parallel, particularly since some Samolus species
have a less pronounced perigyny and almost appear to be
hypogynous.
The monogeneric tribes Ardisiandreae and Cyclamineae
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were both found to be ingroups in the Primuleae, but the
position of the Corideae is somewhat more difficult to ascer-
tain. In the initial analysis, Coris came out as the sister group
of a monophyletic Lysimachieae, sharing elongated corolla
epidermis cells, glandular anther filaments, and ovules
immersed in the placenta. After successive weighting,
however, Coris is placed at the base of the tribe Primuleae
together with Samolus. The unique zygomorphic flowers of
Coris must be interpreted as an autapomorphy evolved from
ordinary actinomorphic flowers in the ancestor, independent
of the position of the genus.
It was anticipated by us that the tribe Lysimachieae would
come out paraphyletic in relation to the tribe Primuleae, con-
For personal use only.
sidering their open deeply lobed flowers and their secretory
cavities. However, the monophyly of the Lysimachieae was
supported by our unweighted analysis, although the subse-
quent analysis with successive weighting indicated them to
be a paraphyletic grade in the very base of the family, just
next to Maesa.
Compared with the character states occurring in Maesa,
the Lysimachieae (excluding Coris) could be diagnosed by
the following characters: open rotate corolla, twisted corolla
aestivation, sepals more or less free to base, flowers with
nectar-producing trichomes, and anther filaments attached at
base of the corolla tube. The genus Lysimachia was found to
be paraphyletic, with Anagallis, Trientalis, Glaux, and
Asterolinon being specialized ingroups. The genus Glaux
was found to be the sister group of Asterolinon, which is
interesting since both genera show a reduction in petal size.
In Asterolinon the petals are minute and inconspicuous,
whereas Glaux is completely apetalous.
The Primuleae clade was diagnosed by the following
characters: basal rosulate leaves, absence of secretory cell
cavities, and umbellate inflorescence subtended by a whorl
of bracts. The position of the scape is hard to assess, but the
ancestral condition can be hypothesized to be a laterally axil-
lary position, as in Ardisiandra. There is strong support for
a sister-group relationship between Dodecatheon and Cycla-
men, which is expressed by a striking similarity in the
general aspect of the flowers. The strongly reflexed corolla
lobes are actually one of the synapomorphies uniting Dode-
catheon and Cyclamen, but features like absence of articu-
lated hairs, acute petals being longer than wide, and punctate
or truncate stigma are other character states uniting these
genera. Furthermore, Dodecatheon and Cyclamen belong to
a group within the Primuleae that is supported by features
like flowers without nectaries, sagittate anthers forming a
Can. A. Bot. Vol. 73, 1995
cone that become spreading at anthesis, anther filaments
shorter than the anthers and attached at base of the corolla
tube, and style exserted far beyond the anthers. Some of
these features are also shared with the other genera of this
group, viz. Ardisiandra, Soldanella, and Cortusa. The stated
affinity of Dodecatheon to North American species of Prim-
ula (e.g., Richards 1993) is probably due to symplesio-
morphic similarity.
The systematic position of Maesa
The systematic position of the genus Maesa has always been
with the Myrsinaceae, but most workers regarded the genus
as a rather aberrant off-shoot from the main core of genera.
De Candolle (1844) placed Maesa in a separate tribe, the
Maeseae, and Pax (1889) described the subfamily Mae-
soideae to account for the genus within the family. In this
context, it is also noteworthy that Maesa differs from most
other Myrsinaceae by having mixed uniseriate and multiseri-
ate rays, and it was differences in wood anatomy that led
Janssonius (1920) to suggest that Maesa ought to be excluded
from the Myrsinaceae.
Mez (1902) followed Pax' (1889) treatment and retained
Maesa in the Myrsinaceae but remarked that Maesa was inti-
mately related to Samolus, which perhaps should be better
lifted out of the Primulaceae and put into a separate group
together with Maesa. A similar view had been expressed
earlier by Bartling (1830). The differing features between the
genera according to Mez (1902), were the woody habit of
Maesa, its noncapsular fruit, and its presence of bracteoles.
There are many characters indicating that Maesa and the
Primulaceae evolved from a common ancestor, but our
analyses do not place Samolus and Maesa in a monophyletic
group, and the affinity referred to by Mez (1902) is probably
due to parallel evolution.
W e did not include any cytological data in our cladistic
analysis, because of the coding difficulties due to a consider-
able variation. However, if our tree topologies are compared
with Faure's hypothesis of chromosome number evolution
(Faure 1968), the position of Aegiceras in the Myrsinaceae
gain further support, and the aberrant chromosome numbers
of the Theouhrastaceae reflect their uosition outside of the
other two families. It is also noteworthy that the chromosome
number in Maesa (x = 10; e.g., Gajapathy 1962; Faure
1968: Paiva and Leitao 1987) is different from those found
in thk Myrsinaceae proper but agrees well with those of
many Primulaceae (see, e.g., Fedorov 1974, for further
references).
In many characters Maesa differ from the fairly homo-
geneous Myrsinaceae: the leaves lack immersed glandular
hairs, the inflorescence is not fascicled or pseudoumbellate,
the flowers are perigynous, the pedicel has a pair of bracte-
oles subtending the calyx, the corolla is campanulate or
shortly tubular, the stamens are adnate to the corolla at about
the middle or higher, the anthers are not sagittate, the gyno-
ecium has somatic nectaries, the fruit is inferior, many-
seeded, and formed by the ovary as well as the surrounding
tissue, the ovules are densely set on the placenta and have an
inner integument with a tapetum, and the seeds are sectoroid
without depressed hilum area and provided with rhomboid
crystals in the testa.
Three options are available to resolve the situation with
 Anderberg and St%hl
the paraphyletic Myrsinaceae: (i) Maesa only o r (ii) the
entire Myrsinaceae are included in the Primulaceae, o r
(iii) a less drastic option is to give Maesa equal rank as a
family. W e believe that taxonomic disorder would b e the
result if two so well known families as the Primulaceae and
the Myrsinaceae were united into one. W e also believe that
it would b e unsatisfactory to unite the frutescent Maesa with
I the herbaceous Primulaceae.
I
W e believe that the best taxonomic solution would be to
raise Maesa to the family level. This would result in four
well-diagnosed families within the Primulales and in some
ways would also b e congruent with the view of earlier
authors. Our conclusion echoes the view by, e.g., d e Can-
dolle (1841b), who at one time considered this possibility but
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eventually refrained because h e considered the similarity
between Maesa and the Myrsinaceae to b e greater than the
corresponding differences between them and neighboring
families such as Sapotaceae o r Ebenaceae. In the light of the
present analyses, the overall similarity in habit between
Maesa and the Myrsinaceae can b e interpreted as sym-
plesiomorphies o r parallelisms, and therefore, the words of
d e Candolle (1841b) have a new significance:
Les MaesCes s'kloignent cependant plus gros des MyrsinCacCes
que toute autre tribu, et peuvent Ctre considCrCes par ce motif
comme une famille distincte ou comme une sous-famille
(subordo); j'incline ?i ce dernier mode parceque leur distance
des vraies MyrsinCacCes est plus faible que celle des EbCnacCes
For personal use only.
et SapotacCes, des ~ r i m u l a c ~ eets ~~rsinCacCes,admises
ordinairement comme familles.
(de Candolle 1841b, p. 167)
i Acknowledgements
1 Qiu, Y.-L., Kron, K.A., Rettig, J.H., Conti, E., Palmer, J.D.,
W e are indebted to Pollyanna Lidmark for preparing the
illustrations, and to Else Marie Friis and Stefan Vogel for
valuable comments on various issues. W e a r e also grateful to
Steve Farris for providing the software used for the permuta-
tion tests, to Anna-Lena Anderberg for technical assistance,
and to o n e anonymous reviewer for valuable comments. W e
also acknowledge the financial support of Swedish Natural
Science Research Council grants B-BU 08950-303 (to A.A.),
and B-BU 06389-303 (to B.S.).
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 Anderberg and St%hl

anomalous and normal pollen of Rapanea: morphology, elemen- Leaves

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349 -379. lamina and petiole indistinct
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1, present
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E. Warming. KGbenhavn. pp. 1-45. 1, articulated present
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Selsk. Skr. Nat. Mat. 8, 11: 6. Copenhagen.
Two-armed hairs
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0, absent
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..... . .
.. ...,
Primula subgenus Sphondylia (Syn. Sect. Floribundae) with a Leaves
review of the subdivision of the genus. Arb. Univ. Bergen. 0, without immersed glandular hairs
Mat. Nat. Ser. 11: 1-49. 1, with immersed glandular hairs
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related to crimula with special reference to their pollen mor- 0, not farinose
phology. Arb. Univ. Bergen. Mat. Nat. Ser. 19: 1 - 15
For personal use only.

1, farinose
Woodcock, E.F. 1933. Seed studies in Cyclamen persicurn Mill.
Annu. Rep. Mich. Acad. Sci. 17: 415-419. Secretory cell cavities
Wunderlich, R. 1967. Some remarks on the taxonomic significance 0, absent
of the seed coat. Phytomorphology, 17: 301 -31 1. 1, present
Yamazaki, T. 1972. Embryological studies in Ebenales (4). Foliar sclereids
Ebenaceae. J. Jpn. Bot. 47: 20-272. 0, absent
Yang, Y-P., and Dwyer, J.D. 1989. Taxonomy of subgenus Blad- 1, present
hia of Ardisia (Myrsinaceae). Taiwania, 34: 192-298. Extraxy lary fibres
0 , absent
1, in layers or bundles
Appendix 1 2, scattered
Characters used for the cladistic analyses of the Primulales. Prismatic foliar crystal druses
0 , absent
1. Habit 1, present
0, trees or shrubs Inner wall of leaf epidermis cell
1, dwarf shrubs 0, not gelatinized
2, suffrutices or perennial herbs 1, gelatinized
3, annual herbs Inflorescence
2. Growth form 0, unbranched
0, not pachycaulous (branched) 1, branched
1, with pachycaulous growth form 2, fascicled
3. Rhizomes Inforescence
0, not long and slender 0, cymose
1, long and slender 1, racemose
4. Leaves Flowers
0, alternate 0, neither solitary nor arranged in umbels or
1, opposite or whorled pseudoumbels
5. Leaves 1, arranged in umbels, generally subtended by a whorl of
0, neither rosulate nor pseudoverticillate bracts
1, basal, rosulate 2, arranged in condensed racemes, pseudoumbels
2, terminal, pseudoverticillate 3, solitary
6. Leaf margins Inflorescences
0, entire 0, lateral or axillary
1, variously serrrate or dentate 1, terminal
 Can. J. Bot. Vol. 73, 1995

Appendix 1 (continued). 41. Petals

0, longer than sepals or subequal
Bracts 1, much smaller than sepals
0, persistent at time of flowering 2, missing
1, soon caducous 42. Petals
Pedicel 0, broadly ovate-rotund, shorter or about as long as wide
0, without a pair of bracteoles 1, lanceolate-ovate, more or less acute and usually
1, with a pair of bracteoles distinctly longer than wide
Flowers 43. Adaxial petal surface
0, hypogynous 0, without capitate trichomes (granules)
1, perigynous 1, with capitate trichomes (granules)
Flowers 44. Petal epidermis cells
0, homostylous 0, rounded, more or less isodiametric
1, heterostylous 1, elongated, about twice as long as broad
Can. J. Bot. Downloaded from www.nrcresearchpress.com by UNIV CHICAGO on 02/16/13

Flowers 45. Flowers

0, unisexual 0, without nectaries and oil (pollen flowers)
1, bisexual 1, with nectar-producing trichomes
Flowers 2, with oil-producing trichomes
0, polystemonous 3, with somatic gynoecium nectaries
1, diplostemonous to triplostemonous
46. Anther filaments
2, haplostemonous with the functional stamens opposite 0, without long septate trichomes
the corolla lobes
1, with long septate trichomes
Perianth
47. Anther filaments
0, 5-merous
0, free from each other
1, 4-merous
2, 6-merous 1, fused into a more or less pronounced ring
3, usually 7-merous 48. Anther filaments
0, attached at base of the corolla tube
For personal use only.

Corolla
0, lobes free or almost free at the base 1, attached at about the middle of the corolla tube or
1, distinctly sympetalous higher
Corolla 2, filaments or filament tube fused to and decurrent on
0, open, more or less rotate lower part of corolla
1, campanulate to deeply bowl-shaped 49. Anther filaments
2, with short narrow tube 0, eglandular
3, with long tube, more or less hypocrateriform 1, glandular
Corolla 50. Anther filaments
0, without staminodes 0, longer than the anthers (i.e., anther shorter than its
1, with staminodes distance from the receptacle)
Corolla aestivation 1, shorter than anthers (i.e., anther longer than its
0, imbricate or quincuncial distance from the receptacle)
1, twisted 51. Anthers
2, valvate or valvate-involute 0, longitudinally dehiscent, extrorse
Corolla 1, longitudinally dehiscent, introrse
0, not constricted at the throat 2, poricidal
1, constricted at the throat 52. Anthers
Corolla throat 0, broadest at the base, sagittate, conspicuously triangular
0, without scales and sometimes acuminate
1, with scales 1, oblong rounded at apex and broadest at the middle, not
Corolla texture sagittate
0, thick
53. Anthers
1, thin, delicate
0, longer than wide
Sepals
1, shorter, or about as long as wide
0, more or less free to base
1, united, often cup-shaped (synsepal) 54. Anthers
Petals 0, versatile
0, erect to spreading, not reflexed 1, not versatile
1, strongly and abruptly reflexed 55. Anther thecae
2, evenly recuwed, but not abruptly reflexed 0, not septate
Petals 1, septate
0, entire 56. Anthers
1, emarginate 0, not curved after anthesis
2, fringed 1, curved after anthesis
 Anderberg and Stahl

Appendix 1 (concluded). 74. Ovules

0, in two or more series, usually many
57. Anthers 1, in a single series, few or very few
0, not forming a cone 75. Seeds
1, initially forming a cone, but eventually spreading 0, ellipsoid, without depressed hilum area
2, forming a cone throughout anthesis 1, subglobose to obtusely angled, without depressed
58. Anther connective hilum area
0, without secretory cavities 2, subglobose, with a depressed hilum area
1, with secretory cavities 3, sectoroid, more or less angled, without depressed
59. Anthers hilum area
0, without large apical appendage 4, laterally compressed, often sickle-shaped (e.g.,
1, with large apical appendage Roosmalen 1985)
60. Anther thecae 76. Seed coat
0 , without calcium oxalate crystals 0, not sclerotic, or with few scleric cells
Can. J. Bot. Downloaded from www.nrcresearchpress.com by UNIV CHICAGO on 02/16/13

1, with calcium oxalate crystals in upper and lower part 1, sclerotic with several layers of scleric cells
61. Pollen 77. Cotyledons
0, 3-, 4-, or 5-colporate, or colporoidate 0, not foliaceous in seed
1, zonocolpate 1, foliaceous in seed
62. Style 78. Testa
0, included, shorter than, or slightly longer than the 0, without rhomboid crystals
anthers 1, with rhomboid crystals
1, exerted, protruding far beyond the anthers 79. Endosperm cell walls
63. Style 0, smooth
0, shorter or as long as ovary 1, pitted
1, longer than ovary 80. Inner integument
64. Style 0, with tapetum
0 , more or less deeply cleft 1, without tapetum
For personal use only.

1, entire 8 1. Embryogeny
65. Stigma 0 , of the solanad type
0 , punctate or truncate 1, of the caryophyllad type
1, capitate or subcapitate, sometimes slightly lobed 2, of the onagrad type
82. Endosperm formation
66. Fruit
0, cellular
0 , a berry
1, nuclear
1, drupaceous, more or less fleshy
2, a capsule
3, dry or fleshy, formed by the ovary and surrounding
tissue Appendix 2
67. Fruit
Material used for the anatomical study of seeds (specimens in
0 , normally many-seeded
Swedish Museum of Natural History (S), unless otherwise
1, normally one-seeded
stated).
68. Fruit opening - -

0, by irregular disintegration of the fruit wall Sapotaceae

1, with teeth or valves Manilkara zapota (L.) van Roy. (Lundell and Contreras
2, with a circular lid 19114)
69. Fruit Monotheca blcxifolia (Falconer) A.DC. (Hedge et al. W7429)
0, ovoid to ellipsoid or cylindrical Ebenaceae
1, globose or subglobose Diospyros tetraspema Sw. (Stearn 68)
2, very elongated Diospyros icrenensis Britton (Broadway 6586)
My rsinaceae
70. Placenta
Aegiceras corniculatum (L.) Blanco (Brass 25462)
0, not free central
Ardisia pusilla A.DC. (Maximowicz s.n.)
1, free central
Conandriurn polyanthum (Laut. & K . Schum.) Mez (Brass
71. Ovules 25475)
0, densely set on placenta Conornorpha pseudo-icacora (Miq.) Mez (Prance and
1, well spaced on placenta Pennington 1725)
72. Ovules Corelliana multiflora (A.C.Sm.) D'Arcy (Frame et al. 179)
0, unitegmic Embelia ribes B u m . (Tsang 21967)
1, bitegmic Grammadenia assymetrica Mez (Neil1 and Palacios 7660,
73. Ovules AAU)
0, not immersed in the placenta Heberdenia bahamensis (Gaertn.) Sprague (Mand6n 177, Stihl
1, immersed in the placenta et al. 801)
 Can. J. Bot. Vol. 73. 1995

Appendix 2 (concluded).

Labisia pumila (Blume) F.-Vil. (Rahmat Si Boeea 6986)

Maesa alnifolia Haw. (Hilliard and Burtt 16756)
Maesa lanceolata Forssk. (Bernardi 11070)
Maesa rufescens A.DC. (Wall s.n.)
Myrsine afn'cana L. (Armitage 21822)
Oncostemum sp. (Nicoll et al. 538)
Parathesis belizensis Lundell (Contreras 7957)
Tapeinosperma clavatum Mez (A. C. Smith 5379)
Wallenia laurifolia Swartz (Ekman 1047).
Theophrastaceae
Clavija eggersiana Mez (Knudsen 105, GB)
Clavija lancifolia Desf. ssp. chermontiana (Standl.) St&]
Can. J. Bot. Downloaded from www.nrcresearchpress.com by UNIV CHICAGO on 02/16/13

(Foster and Terceros 13447)

Jacquinia armillaris Jacq. (Glaziou 12933)
Jacquinia arborea Vahl (Webster and Wilson 5231)
Jacquinia nitida Stihl (Moreno 23478, GB)
Jacquinia umbellata A.DC. (Stihl 155, GB)
Theophrasta americana L. (St&] and Lindstrom 178, GB)
Primulaceae
Anagallis monellii L. (Anderberg 7278)
Androsace m i m a L. (Steinitz 307)
Androsace obtusifolia All. (Grenier s.n.)
Ardisiandra sibthorpoides H0ok.f. (Schlieben 4806)
Asterolinon linum-stellatum (L.) Duby (Anderberg 1143)
Centunculus minimus L. (Houfek 317)
For personal use only.

Coris monspeliensis L. (Ross 182)

Cortusa matthioli L. (Aitchinson 725)
Cyclamen coum Miller (cult., Stockholm)
Dionysia balsamea Wendelbo & Rech.f. (Rechinger 18918)
Dodecatheon frigidurt~ Cham. & Sch. (Spetzman 3092)
Douglasia nivalis Lindl. (Sandberg and Leiberg 545)
Glaux maritirna L. (Stihl and Knudsen s.n.)
Lysimachia ciliata L. (Biltmore 3477b)
Lysimachia clethroides Duby (Murata and Shimizu 1302)
Lysimachia glutinosa Rock (Sparre H 34)
Lysimachia insignis (Cavalerie 2721, K)
Pelletiera verna St. Hil. (Sparre 179)
Primula farinosa L. (Wanntorp s.n., SUNIV)
Primula veris L. (Wall s.n.)
Samolus valerandi L. (Samuelsson 1043)
Soldanella alpina (Anderberg and Anderberg 73 10)
Trientalis europaea L. (Anderberg and Anderberg s.n.)