26 Araceae

Araceae VEGETATIVE MoRPHOLOGY. The stem varies

from an elongated aerial axis with extended
S.J. MAYO, J. BoGNER, and P.C. BoYcE internodes, as in the many climbers and climb-
ing hemiepiphytes, to a hypogeal rhizome or
depressed-globose (rarely elongate to napiform)
tuber. Genera with rhizomes or tubers are com-
mon in Lasioideae and Aroideae. The climbing
habit is characteristic of subfamilies Pothoideae
and Monsteroideae. Abbreviated aerial stems
Araceae Juss., Gen. Pl.: 23 (1789), nom. cons. are also frequent, producing rosulate plants, as in
certain epiphytic species of Philodendron and
Herbs, diminutive to gigantic, terrestrial, geo- Anthurium. Erect stems with relatively short
phytic, epiphytic, or climbing, rarely aquatic; internodes occur in some tribes of Aroideae
with true or false stems; tissues with raphides; (e.g. Aglaonemateae, Dieffenbachieae ). Some,
laticifers commonly present; mature shoots com- generally larger, species have an arborescent habit
posed of articles forming a sympodium (except in which the main axis is a fleshy (Alocasia,
Pothoideae, Heteropsis ); leaf bifacial, usually dif- Montrichardia, Xanthosoma sect. Xanthosoma)
ferentiated into petiole and expanded blade of or fibrous (Philodendron) stem or a false stem
very variable shape; midrib compound; primary of petiole sheaths (Arisaema, Dracunculus,
venation usually pinnate but sometimes pedate Typhonodorum). Shoots specialised for vege-
or arcuate from the base or strictly parallel, fine tative reproduction include flagelliform shoots
venation ± reticulated or parallel-pinnate (i.e. (e.g. Amydrium, Cercestis, Philodendron, Rhodo-
parallel to primary lateral veins); inflorescences spatha, Syngonium), bulbils adapted for animal
1-many, each inflorescence a pseudanthium dispersal (Remusatia), cormlets (Dracontium,
(especially so when flowers unisexual) consisting Dracontioides) and hypogeal stolons (Ari-
of a spadix (a dense spike of minute, ebracteate saema, Colocasia, Cryptocoryne, Lasimorpha,
flowers) subtended by a spathe (last leaf of article, Spathiphyllum ).
usually specialised in form and colour, incon- In virtually all genera the mature stem is a
spicuous in Gymnostachys, Orontium); flowers sympodium composed of articles ending with
sessile (except Pedicellarum), bisexual or un- an inflorescence or aborted inflorescence. Foliage
isexual (unisexual inflorescences in Arisaema - leaves and cataphylls occur in an often predictable
paradioecy), when unisexual the female flowers sequence within each article; the constancy of the
occupying the proximal part and males the me- sequence varies from genus to genus. With the
dian or distal part of the spadix; perigone present termination of the article by an inflorescence,
or absent, tepals free, partly or completely con- growth of the stem is continued in most genera by
nate; stamens usually 3 + 3 or 2 + 2, or sometimes a branch axillary to the foliage leaf or cataphyll
less or more, opposite tepals in bisexual flowers, situated at the second node below spathe
free to connate, anthers extrorse (except insertion. In the Orontioideae this continuation
Zamioculcas, Pedicellarum); gynoecium syncar- shoot arises at the first node below the spathe.
pous, ovary usually 1-3 locular ( 1-8 locular in "Enrichment" of the flowering shoot to form
Spathicarpeae, 2-47-locular in Philodendron), a synflorescence takes place commonly by the
ovules bitegmic, atropous, anatropous, ana- development of a sequence of short articles,
campylotropous or of intermediate form; style each consisting at least of prophyll, spathe and
usually inconspicuous, stigma wet at anthesis, inflorescence, beginning in the axil of the leaf
sometimes distinctly lobed; fruit a berry (basally immediately below the spathe. The resulting
dehiscent in Lagenandra), usually free and borne synflorescences exhibit a range of structural varia-
in ± cylindric, dense, fleshy spike, rarely connate tion which may be quite complex; extreme forms
into syncarp; seeds small to very large, testa very are found in Gymnostachys and Homalo-
thin to very thick, rarely absent, endosperm mena. Anomalous shoot arcliitecture occurs in
present or absent, embryo minute to large, both Gymnostachys, Potheae and Heteropsis, in which
with or without starch, chlorophyllous or not. flowering is confined to lateral short shoots from
A family comprising 104 genera and over 3300 the monopodia! main vegetative axis.
species, with a subcosmopolitan distribution but Roots in Araceae are always shoot born; the
most diverse in tropical regions. primary root withers soon after germination or is
K. Kubitzki (ed.), Flowering Plants · Monocotyledons
© Springer-Verlag Berlin Heidelberg 1998
 Araceae 27

absent (Pistia). In root climbers and hemi- be arcuate-parallel (i.e. acrodromous, campylo-
epiphytes, root dimorphism is frequent in; non- dromous, convergate or curvipalmate, e.g.
geotropic anchor roots, adapted for clinging on Anthurium sect. Digitinervium), pedate (e.g.
trees, are anatomically or morphologically distin- Sauromatum) or radiate (e.g. many Arisaema
guishable from larger feeding roots which grow spp.) but are most commonly pinnately organised.
down to the forest floor, as in Monstera, Philoden- Even in pedately and radiately lobed leaves, the
dron, Rhaphidophora. In some geophytic genera, primary veins of the lobes are generally pinnate.
e.g. Arisarum, Arum, Biarum, and in Crypto- In pinnately veined leaves the major branches of
coryne, which is rhizomatous, contractile roots the midrib are thus usually primary veins, but are
occur. referred to as primary lateral veins. Except possi-
In virtually all genera the leaf is clearly differen- bly in some compound leaves, the primary veins
tiated into a blade, petiole and petiole sheath; ex- always run throughout the leaf blade, ultimately
ceptions are Gymnostachys, some Biarum spp. joining together at the leaf apex. In many genera
and Pistia. The sheath normally clasps the sub- either one or several of the primary veins form
tended internode, at least basally, and has an an- a submarginal collective vein on each side in ad-
nular insertion (except many Potheae and most dition to the marginal vein. The finer venation
Heteropsis ). The foliage leaves of mature shoot ar- may be reticulated (e.g. Anthurium) or parallel-
ticles often have short or very reduced sheaths. pinnate (i.e. more or less parallel to the major
Leaf blade size may range from diminutive lateral veins); the latter is normally referred to as
(e.g. Ambrosina) to gigantic (e.g. Amorphophallus, "parallel" or "striate" venation in the taxonomic
Colocasia, Cyrtosperma, Xanthosoma). Shape literature on aroids. A third, intermediate type
varies from linear (Biarum, Typhonium) to of fine venation is recognised for the tribes
dracontioid (Anchomanes, Dracontium, Pseudo- Colocasieae and Caladieae ("colocasioid vena-
hydrosme, Pycnospatha, Taccarum, Thom- tion"). Here the fine veins arising from major lat-
sonieae), through cordate, sagittate, hastate, eral veins arch towards the leaf margin, and may
trilobed or trisect, pedately, pinnately and radi- run virtually without reticulation (Colocasia) or
ately lobed or -sect. Bipinnatifid, tripinnatifid fuse to form sinuose sympodial collective veins
and combined pedatisect-pinnatifid leaves also ("interprimary veins") between the major lateral
occur. Dracontioid leaves are generally trisect veins (most Alocasia spp., Xanthosoma). Strictly
at the petiole insertion and thereafter more or parallel venation of the typical monocot type is
less dichotomously subdivided. Zamioculcas, found only in Gymnostachys. Intermediate forms
one species of Anaphyllum and one species of occur between virtually all recognised patterns
Anaphyllopsis have truly pinnate compound and this makes venation a difficult character to
leaves. use. Ertl's study (1932) is the only large-scale
Heteroblasty is a striking feature and defines a survey yet made.
number of climbing genera [Cercestis, Monstera,
Philodendron, Pathos (except subgen. Pathos), VEGETATIVE ANATOMY. 1 The rhizodermis is 1-,
Rhaphidophora, Rhodospatha, Syngonium]. It oc- less commonly several-layered. True multiple root
curs both in ontogeny from seedling to mature epidermis resembling a velamen occurs in various
plant and in association with the development Anthurium spp. The cells are not or at most
of flagelliform shoots. Fenestrate or perforated vaguely differentiated into long and short cells.
leaves are another peculiarity well shown in gen- A sclerotic root hypodermis occurs in Culcasieae,
era such as Amydrium, Cercestis, Dracontioides, Montrichardia, Philodendron, Homalomeneae
Monstera and Rhaphidophora. In genera with di- and Anubias. Lobing of the central vascular cylin-
vided leaves the first foliage leaf is normally entire, der occurs in the roots of some Philodendron and
although in Gonatopus and Thomsonieae it is Cercestis species.
already divided. A number of genera have spe- In the stem, a cortical vascular system, distinct
cies with peltate leaves (Ariopsis, Caladium, from the central cylinder, is present in some gen-
Colocasieae and a few species of Anthurium and era (Pathos, Pothoidium, Heteropsis, Anthurium,
Homalomena).
Leaf venation in Araceae is complex and varied
in detail but little studied and the terminology
used is confusing in practice. The primary veins 'This account is based on the treatment by J.C. French in
(i.e. the major veins including the midrib) may Mayo eta!. (1997).
28 Araceae

Monstereae, Philodendron, Dieffenbachia, Syn- and Zantedeschia, necrotic areas provide another
gonium, Caladium, Chlorospatha, Xanthosoma), mode of attactive leaf variegation.
which in Colocasieae often forms a character- Trichosclereids are characteristic of the Spathi-
istic anastomosing system. An unusual type of phylleae and Monstereae (sparsely present in
endodermis, which surrounds each axial vas- Amydrium).
cular bundle individually but not the central Laticifers, normally associated with the phloem
cylinder or leaf traces, is widespread in the of the vascular bundles, occur in the majority of
Schismatoglottideae. Araceae, although absent from bisexual-flowered
Bud trace insertion patterns are also variable. genera except Calla and Orontium. Anastomos-
Three patterns are recognisable in Araceae. Two ing laticifers occur in tribes Colocasieae (in
of them are found mainly in climbing genera and Ariopsis the laticifers are simple), Caladieae, and
a third is characteristic of genera with rhizomes or Zomicarpeae. Latex composition varies, being
tubers (French and Tomlinson 1981a-d, 1983, clear or milky with a variety of constituent latex
1984). particles and chemical constituents. Large quanti-
In the leaf, trichomes and larger epidermal ties of tannin have also been detected in aroid
scalelike processes are rare (occurring in Bognera, latex.
Callopsis, Pedicellarum, Pistia and certain species Resin canals containing terpenoids are known
of Anubias, Cryptocoryne, Homalomena, Philo- from the leaves, stems, roots and inflorescen-
dendron, Schismatoglottis, Stylochaeton, Syn- ces of Cercestis, Culcasia, Homalomena and
gonium, Xanthosoma). Prickles are found on Philodendron.
stems, petioles and peduncles in various species Extrafloral nectaries occur on the leaves, peti-
of Lasioideae, Nephthytideae, Homalomena, oles and spathes of Philodendron and Culcasia,
Anubias, and Montrichardia. Warty petioles and wax glands in the Alocasia species previously
peduncles occur in some genera of Lasioideae and recognised as Xenophya and intravaginal
in Amorphophallus and Arisaema. Stomata tend squamules, of unknown function, occur in the leaf
to be concentrated on the abaxial leaf surface. The axils or at the nodes in Cryptocoryne, Lagenandra
number of subsidiary cells varies from none and Philodendron.
(anomocytic, e.g. in Lysichiton, Orontium, Pistia, Calcium oxalate has been recorded from almost
Symplocarpus) to 2 (paracytic, e.g. in Amor- all genera and is found in various crystalline
phophallus, Arisaema, Arum, Rhaphidophora, forms, including raphides, druses and specialised
Typhonium) to 4-8 (tetracytic, e.g. in Dieffen- idioblasts (biforines). The latter have a "blowgun"
bachia, Lagenandra, Philodendron and Rhaphido- explosive discharge and contain large, often
phora); paracytic is the commonest type. Leaf grooved, raphides neatly arranged in bundles. For
mesophyll is predominantly bifacial and in illustration of the various crystal forms see
many species of Homalomena, Philodendron, Seubert (1993).
Piptospatha, Schismatoglottis and Typhonium is Vessels have been found in the roots of many
distinctively chambered. Petiole anatomy has so genera and in the stems of a few species of
far been little studied. The occurrence of a Epipremnum, Philodendron, Pathos, Rhaphido-
geniculum (pulvinus), usually at the petiole apex, phora and Scindapsus.
is a useful taxonomic character and commonest in
genera with bisexual flowers. In Gonatopus, INFLORESCENCE AND FLOWER STRUCTURE. The
genicula also occur at the base of the major leaf inflorescence of almost all Araceae is an un-
pinnae, and in G. boivinii and in at least one branched, fleshy spike of numerous, very small
species of Anthurium (A. oerstedianum) the flowers, and is known as a spadix. The family is
geniculum is located in the middle of the petiole characterized by the absence of floral bracts
instead of the apex or nearly so. and the lack of pedicels in all genera except
Variegation of the leaf blade, petiole and pe- Pedicellarum. Occasionally the spadix is not
duncle is a striking feature of many genera, in- fleshy or only few-flowered. The flowers are
cluding several which are well known in usually spirally arranged and tightly packed, al-
horticulture (Aglaonema, Dieffenbachia, Cala- though in some species of Amorphophallus (male
dium). Variegated leaves also occur in the juvenile and female flowers), most species of Arisaema
plants of some species of Monstera, Philodendron, and Arisarum (male flowers), Dieffenbachieae
Scindapsus and Syngonium and in general seem to (female flowers), Pedicellarum and Pathos series
be more common in plants growing in tropical Goniuri, they may be somewhat distant from one
rainforests on or near the ground. In Caladium another.
 Araceae 29

The last leaf of each flowering article is called the Flowers in Araceae may be 2- or 3-merous. In
spathe and is part of the inflorescence. The spathe perigoniate flowers the tepals, when free, are
is usually distinctive, although in Gymnostachys organised in 2 whorls. The tepals are usually ±
and Orontium it remains an inconspicuous bract. fleshy and fornicate apically (except Potheae)
The internode between spathe and spadix (termed and in some genera or sections (Anadendrum,
the spadix stipe) is usually rudimentary, whereas Holochlamys, Pedicellarum, Spathiphyllum sect.
that between spathe and uppermost foliage leaf Massowia, Stylochaeton) they are fused so that the
(termed peduncle) is much longer. In some con- perigone is a single, cuplike structure. Stamens
servative genera, however, this arrangement is re- in perigoniate flowers and in the naked bisexual
versed (Gymnostachys, Orontioideae, some Pathos flowers of most Monsteroideae have essentially
spp.). Gymnostachys, Orontioideae and Poth- the typical structure of a distinct (usually
oideae seem to represent earlier, less uniform flattened) filament, terminal basifixed anther
phases of organisation. The typical araceous and slender, inconspicuous connective. In the
inflorescence has given rise to a wide range unisexual flowers of Philodendron, Anubias and
of variant forms in different genera, which are Homalomeneae, however, the stamens typically
seen as representing an overall evolutionary lack filaments and have a thick, fleshy connective
trend of increasing integration towards a pseu- which is probably osmophoric. Stamens of the
danthium. The major phyletic modifications are Colocasieae and Caladieae are similar but always
(1) loss of perigone in the flowers; (2) differentia- fused into synandria. In Arophyteae and other
tion of flowers on the spadix into a lower female Aroideae the stamens may be fused or not but
zone, upper male zone and, often, one or several generally lack conspicuous filaments. Large, fused
sterile zones of sterile flowers or axial regions connectives also occur in the Spathicarpeae but
lacking flowers - terminal appendices are a well- differ from those of the Caladieae and Coloca-
known feature of more advanced genera (e.g. sieae. Well-developed connectives also occur in
Areae); (3) differentiation of the spathe into a Zomicarpeae, especially in Filarum, where they
lower, convolute tube and an upper, expanded are elongated and thread-like. Anthers are
blade. almost always extrorse (introrse in Zamioculcas,
Spathe and spadix modifications are closely re- latrorse in Pedicellarum). The thecae dehisce by
lated, so that in a phyletic sense the spathe has longitudinal slits (most genera with bisexual
become increasingly integrated into the pseu- flowers and some unisexual flowered genera:
danthium. In extreme cases (e.g. Cryptocoryneae, Anubias, some Areae, Arisaema, Arisarum,
Ambrosina, some Typhonium spp., Pinellia, Stylochaeton) or by apical or subapical pores or
Pistia, Spathantheum and Spathicarpa), fusion short slits. In Philodendron, Anubias, Homalo-
and still more elaborate modifications have meneae, Caladieae and Colocasieae, the dehis-
brought about division of the spathe into separate cence of each theca is usually by a subapical pore,
chambers. In some genera (e.g. Philodendron) the which seems to be correlated with the extrusion of
entire spathe persists until fruit, but in most the pollen in strands; similar structures occur in
Caladieae, Colocasieae and Schismatoglottideae Amorphophallus and Dracunculus.
the blade withers or drops off immediately after The number of locules in the ovary normally
anthesis and only the tube persists until fruiting. varies between 1 and 3, and, when unilocular, the
In Monstereae the entire spathe withers or drops gynoecium often shows traces of 2- or 3-merous
off soon after flowering, a behaviour which seems origin through the presence of a several-lobed
to be correlated with the presence in this tribe of stigma (e.g. Typhonodorum) or more than 1 pla-
numerous protective trichosclereids in the style centa (e.g. most Schismatoglottis spp.). Gynoecia
tissue. Terminal appendices of the spadix are with more than 3 locules are less common but are
found in some genera of the Colocasieae and found in the Spathicarpeae (1-8-locular) and are
Schismatoglottideae, all genera of the Zomi- normal in Philodendron (2-47-locular). Placenta-
carpeae, Thomsonieae, Areae and Arisaemateae tion varies from axile to parietal or basal or apical
and sporadically elsewhere in the family. The or both basal and apical, with many intermediates.
function of the appendix, where studied, is to pro- Ovules may be anatropous, amphitropous,
duce odours which attract pollinators. Structur- anacampylotropous, atropous or intermediate be-
ally the appendix varies from being covered by tween these types. Ovular trichomes are usually
staminodes (e.g. Pseudodracontium) to entirely present and secrete a clear, mucilaginous sub-
smooth with no vestige of floral organs (e.g. most stance which in many genera (e.g. Monstereae,
Areae). Philodendron) entirely fills the ovary locules; this
30 Araceae

secretion may play a role in pollen-tube growth. form a parietal cell or not), most Araceae so far
The style may be narrow and elongated (e.g. investigated, except Symplocarpus and Calla, have
Dracontium) but in most genera it is superficially tenuinucellate ovules.
inconspicuous, even when a thick stylar region Linear megaspore tetrads are the commonest
is developed above the ovary locules. Stigmas type, but T-shaped tetrads are also found and
are wet in Araceae and in some genera (Anthu- sometimes both types occur in the same species.
rium, Arum, several Lasieae) produce con- The mature embryo sac is usually of the 8-nucleate
spicuous nectar droplets at anthesis. In Alo- type (10-12-nucleate in Nephthytis). Embryogeny
casia, Amorphophallus, Dieffenbachia and is clearly understood in only nine genera. Onagrad
some Spathicarpeae the lobing of the stigma is and asterad embryogeny are known in unisexual-
pronounced or the stigma is relatively massive. flowered genera and caryophyllad and solanad
types are known in bisexual-flowered genera.
INFLORESCENCE AND FLOWER ANATOMY. Eyde et Cellular and free-nuclear types of endosperm
al. (1967) established that in floral anatomy, as in development both occur but, according to
other characters, Acorus differed markedly from Grayum (1991), endosperm development in
the Araceae and confirmed the absence of floral Araceae is best interpreted as a form of the
bracts in Araceae. Studies of inflorescence and flo- Helobial type. Chalaza! and micropylar cells
ral anatomy in Philodendron have demonstrated ("chambers") are formed by the first division of
the existence of spathe tissues adapted for opening the endosperm nucleus, followed by cellular
and closing movements, resin secretion of various or free nuclear divisions in the micropylar
types from the spathe and a wide variety of chamber. The chalazal cell remains undivided,
gynoecial and androecial structure (e.g. Mayo usually with a single, hypertrophied nucleus,
1986). Barabe and coworkers (e.g. Barabe and forming a "haustorium" or "basal apparatus". An
Forget 1988) have studied the floral anatomy of endothelium, derived from the inner surface
representatives of a range of tribes with par- of the inner integument, is reported for most
ticular emphasis on floral vasculature. Family- genera investigated.
wide surveys by French of endothecial thickenings
(French 1985a,b, 1986a), stamen vasculature PALYNOLOGY. Major contributions: Thanikai-
(French 1986b), ovule vasculature (French 1986c) moni (1969), Grayum (1984, 1985, 1986a, b, 1992).
and ovular trichomes (French 1987) have further The pollen grains may be sulcate, extensive-
confirmed the distinctness of Acorus and revealed sulcate, meridionosulcate (zonate), disulcate
taxonomically useful character variation. (diaperturate), forate (periporate) or inaperturate
(omniaperturate) (Fig. 10).
EMBRYOLOGY. The embryology of Araceae has Sulcate pollen grains are commonest in the
been reviewed by Jiissen (1928) and Grayum more conservative genera, i.e. those with bisexual,
(1984, 1991). Thorough treatments of all embryo- perigoniate flowers. Spathiphyllum, in spite of
logical aspects have been published for Peltan- several primitive characters, has inaperturate
dra virginica, Synandrospadix vermitoxicus and pollen grains as do all monoecious taxa except
Theriophonum minutum; other genera are known Zamioculcadeae.
less completely. Generalisations about the family Exine ornamentation may be smooth (psilate),
are therefore based on rather fragmentary scabrate, foveolate, reticulate, spinulose, spinose,
coverage. fossulate, rarely gemmate, verrucate, areolate,
Araceae have a periplasmodial anther tapetum, rugulate or striate. Foveolate or reticulate exines,
thus differing from Acorus, which has the secre- widespread among monocots, are considered to
tory type. Pollen mother cell division is probably be primitive in Araceae.
always successive. Both 2-nucleate and 3-nucleate Pollen-grain shape is globose to ellipsoid (boat-
pollen is widespread in the family, but pollen shed shaped) or hamburger-shaped (Gonatopus, some
at the 2-nuclear stage is the condition normally Heteropsis spp., most Monstereae, Zamioculcas).
found in the less advanced genera (Grayum The pollen grains are usually shed as monads, but
1986a). in two relatively advanced genera (Chlorospatha,
The nucellar epidermis of virtually all Araceae Xanthosoma) they are release as tetrades; the tet-
forms a "nucellar cap", usually 2-3 cell layers rads are arranged tetrahedrally or in a serial se-
thick (1 in Pistia), and for this reason the Araceae quence (Chlorospatha longipoda).
have been termed crassinucellate. However, in a The size of the pollen grains varies from small
strict sense (division of the archesporia! cell to (12 f!m in Homalomena versteegii) to large (ca.
 Araceae 31

Fig. lOA-E. Araceae. Pollen SEM micrographs. A Gymno-

stachys anceps, sulcate, exine foveolate. X3800. B Gonatopus
angustus, zonasulcate, exine foveolate. X 1200. C
Sauromatum venosum, inaperturate, exine spinose. XS60. D
Spathiphyllum cochlearispathum, inaperturate, exine striate.
X2200. E Pistia stratiotes, inaperturate, exine plicate with
undulate frills. X900. (Photo W. Barthlott)

60 ~-tm) or very large (106 ~-tm in Pseudohydrosme The pollen of most genera contains starch (73%
gabunensis), but in the majority of genera (68%) of the examined species), the remainder being
the grains are medium-sized (between 25 and starchless; in Schismatoglottis both types have
50 ~-tm diam., mean 37~-tm). been found in different species.
32 Araceae

It seems probable that exine ornamentation is The size and shape of the chromosomes are also
correlated with pollinator type. Smooth pollen quite variable. The exceptionally large chromo-
(and probably pollen extruded in strands) is usu- somes in Anchomanes and Nephthytis (Marchant
ally associated with beetle pollination according to 1971a) is an important reason for placing these
existing observations, while spinose pollen is usu- genera in the same tribe.
ally found in fly-pollinated genera. Exceptions
have, however, been observed. In Cryptocoryne, PHYTOCHEMISTRY. The Araceae are well known
smooth pollen grains are associated with pollina- as a family which includes poisonous plants. Many
tion by minute flies; in bee-pollinated Peltandra ornamentals (e.g. species of Alocasia, Caladium,
sagittifolia pollen is smooth; Amorphophallus Dieffenbachia, Philodendron, Zantedeschia) are
maculatus, with somewhat scabrous pollen, known to cause poisoning, particularly wheu veg-
appears to be pollinated by both beetles and etative plant parts are accidentally eaten by chil-
blowflies. dren and pets. The red berries of Arum are known
to be poisonous and the tubers of all e :lible
KARYOLOGY. Petersen (1989) has provided a criti- Araceae must be cooked or treated before the'r can
cal and comprehensive review. Chromosome be eaten. The presence of active compounds i~ a: so
numbers in Araceae vary from 2n = 14 (Ulearum) evident from the use of Araceae for medicinal
up to 2n = 168 (Arisaema). The number may vary purposes by people in many parts of the world;
considerably within one genus, e.g. in Crypto- e.g. see Plowman (1969); Knecht (1983); !:own
coryne (2n = 20 to 132) or in Arisaema (2n = 20 (1988). The chemical basis of the toxicity of
to 168). Other genera have very stable numbers, Araceae is not completely understood, but it is
as in Anthurium, the largest genus of the family generally assumed that calcium oxalate raphicles
(2n = 30 in the great majority of spp.), Dracon- can cause minute lesions in the mouth, thus fac:Ji-
tium (2n = 26), and Zantedeschia (2n = 32). tating the access of toxic chemical compounds to
The genera with bisexual, perigoniate flowers the blood system.
tend to have less variation in chromosome num- Ethereal oils are present in Homalor.1ena
ber and neither very high nor very low numbers; but do not occur in specialised idioblast1:. In
for example, Pothos has 2n = 24 and 2n = 36 and Araceae, major groups of compounds include
Spathiphyllum 2n = 30 and 2n = 60. The highest saponins, proanthocyanidins, cinnamic acids,
numbers appear in tribes with advanced floral and flavonoids, cyanogenic glucosides, alkaloids a1d
inflorescence morphology, e.g. the Areae, the amines; cyanogenesis is widespread. Irritant
Arisaemateae and the Cryptocoryneae. In certain compounds so far identified include protocate-
groups with advanced floral characters (e.g. chuic aldehyde, homogentisic acid and :h·~ir
Zomicarpeae), there has been a reduction in chro- glycosides (Alocasia, Colocasia, Pinellia) nd
mosome number. allergenic 5-alkyl- and 5-alkenylresorci nols
Although a primary base number of x = 7 has (Philodendron). The flavonoids and antho-
been proposed by other authors, Petersen (1989) cyanins have been surveyed in Araceae (Will ams
considered that the base numbers x = 14 or x = 12 et al. 1981). C-glycoflavones, flavonols and pro-
may have been the starting point for the deriva- cyanidins are the main flavonoids, while flav,)nes
tion of all the chromosome numbers in extant are of limited ocurrence. Broadly speaking, tlle
genera of Araceae. These numbers may represent chemistry of the Araceae is comparable to th 1t of
secondary base numbers. the Liliiflorae (Hegnauer 1987, 1997 in Mayo et :tl.
High and diverse chromosome numbers in 1997).
Araceae are thought to have been derived by aneu-
ploidy and polyploidy. Thus the aneuploid series PoLLINATION. Araceae inflorescences are almost
x = 11, 10, 9, 8 (and probably 13, 12) could have always insect pollinated, although "wind-tunnel"
been derived by reduction, and x = 15, 16, 17, 18, pollination has been proposed for Pinellia (Uhlarz
19, 20, 21 by an increase in chromosome number 1985). Pollinators so far reported for Araceae
from a primitive (but possibly secondary) base (Grayum 1984) include trigonid bees (Mons,'era,
number of x = 14. Spathiphyllum), euglossine bees (Anthur,'um,
Diploids, triploids, tetraploids and still higher Spathiphyllum), beetles of the families Asilidae
levels of ploidy can be found within one species, as (Amorphophallus ), Cetoniidae (Amorphophallu.; ),
in Cryptocoryne ciliata with diploids and triploids Curculionidae (Anthurium, ?Pistia), Dermestidae
(2n = 22, 33), and Cryptocoryne cordata with (Dracunculus), Nitidulidae (Alocasia, Amcr-
2n = 34,68, 85,102. phophallus, Anchomanes, Anubias, Aridarum,
 Araceae 33

Cercestis, Culcasia, Cyrtosperma, Nephthytis, and many Spathiphyllum species have pure white
Typhonium), Scaphidiidae (Pseudohydrosme), spathes.
Scarabaeidae (Alocasia, Amorphophallus, Odour is evidently a prime factor in attracting
Anubias, Caladium, Dieffenbachia, neotropical pollinators, and Araceae are famous for their foul
Homalomena, Philodendron, Syngonium), Scyd- blossom odours. Many genera, however, are
maenidae (Typhonium), Silphidae (Amorpho- less offensive. Spathiphyllum, Philodendron and
phallus), Staphylinidae (Alocasia, Anthurium, Xanthosoma have heavy, spicy odours and in An-
Chlorospatha, Dracunculus, Lysichiton, Pseu- thurium, odours range from spicy to the smell of
dohydrosme, Typhonium), and flies of the fam- decaying apples. Amorphophallus galbra is known
ilies Anthomyiidae (Alocasia), Calliphoridae to have a pleasant smell. Few studies of odour
(Dracunculus, Helicodiceros), Ceratopogonidae chemistry have been made, and this field is still in
(Arum, Cryptocoryne), Chloropidae (Peltandra), its beginnings.
Choridae (Pseudohydrosme), Drosophilidae Inflorescence heating (thermogenesis) in con-
(Alocasia, Colocasia, Culcasia, Homalomena, nection with flowering has been examined mainly
Nephthytis), Ephydridae (Cryptocoryne), Myceto- in Alocasia, Arum, Philodendron, Sauromatum
philidae (Arisaema, Arisarum), Neurochaetidae and Symplocarpus. This phenomenon appears to
(Alocasia), Phoridae (Cryptocoryne), Psychodidae be fairly widespread in the family, although by no
(Arum), Sciaridae (Arisaema, Arum), Simuli- means universal. Its function is generally agreed
idae (Arum), Sphaeroceridae (Arum, Pseudo- to be volatolisation of odour compounds. Attrac-
hydrosme) and Syrphidae (Peltandra). tion to a warm site is also thought to have a role,
The kettle trap mechanisms (in German litera- at least in cases involving carrion beetles and flies.
ture referred to as Kesselfallenblumen) of genera In many genera, however, odours are produced
with unisexual flowers and complex spathes apparently without heating.
(e.g. Arisaema, Arisarum, Arum, Cryptocoryne, Odour production and thermogenesis (when
Lagenandra, Philodendron, etc.) have attracted present) take place mainly in terminal appendices
most attention. Much less is known about pollina- (e.g. Arum) or in the male or sterile male zones of
tion mechanisms in genera with bisexual flowers the spadix (e.g. Philodendron). Odour production
and simpler or spreading spathes (e.g. Anthurium, from the spathe is suspected in Dracontium and
Monstereae, Lasioideae). While kettle "trap" probably occurs in other genera. The thickened
seems an adequate description for Arum and connectives of stamens, staminodes, synandria
Arisaema, in other genera (e.g. Amorphophallus, or synandrodes in Philodendron, Anubias,
Philodendron) it is unclear whether the pollinators Homalomeneae, Caladieae and Colocasieae prob-
are unable or merely unwilling to leave the inflo- ably represent adaptations for osmophore func-
rescence once they have entered it, kept there tion. In Cryptocoryneae, the so-called olfactory
perhaps by attractants such as stigma secretions, bodies are probably osmophoric, although Vogel
food bodies, pollen, odour compounds (male (1963) found osmophores on the spathe limb
euglossine bees) or sites for reproduction. in Cryptocoryne. In genera with well-developed
The colour of the spathe, and to a lesser extent of terminal spadix appendices (e.g. Areae, Arisae-
the spadix, varies considerably within the family, mateae, Thomsonieae), thickened connectives are
ranging from inconspicuous greens (e.g. Anthu- generally absent. Odour production in genera
rium, Nephthytis) to elaborate patterns (e.g. with bisexual flowers is poorly understood. In
Colletogyne, Sauromatum) or striking "flags" Spathiphyllum the stigma is thought to play this
(e.g. Anthurium andraeanum). Fleshlike colours role and in Anthurium the thickened tepal apices
play an important role in attracting pollinators may be involved.
(myiophilous colours) in some genera. Differenti- Araceae are always protogynous and the female
ated colour zones are frequent. In Philodendron (stigma receptivity) and male (anther dehiscence)
many species have purple zones inside the spathe phases usually do not overlap, so that obligate
tube, while the blade is white or pale green. In outcrossing is the general rule. Some cases of self-
Arisaema and Arisarum, the reverse situation is pollination or apomixis are known or suspected
found, with white spathe tubes and dark purple (Anthurium gracile, triploid Amorphophallus
blades which are often striped. The foul odours of paeoniifolius, Arum idaeum, A. hygrophilum,
such species as Amorphophallus konjac are very Pinellia).
often associated with flesh-coloured or livid Manipulation of pollinator behaviour within the
spathes, resembling carrion. By contrast, the per- inflorescence may be the basis for many of the
fumed inflorescences of Zantedeschia aethiopica specialised features of the spadix, particularly in
34 Araceae

unisexual-flowered genera. Spathe constnctwns the inner layer of the pericarp may also be
may act as "skid zones" (Arum) or, in conjunction mucilaginous. This makes the seeds sticky and
with closing movements, as "brooms" to eject aids their dispersal by birds or mammals.
pollinators from the female chamber after pollina- The seeds are usually straight, but in Lasieae,
tion (Philodendron). The various types of hairs, Monstereae and Pothoideae they may be curved
scales and warts found on the inner surface of the or even strongly horseshoe- or kidney-shaped
spathe (e.g. Amorphophallus) or the wide range of (anacampylotropous). Endosperm may be abun-
staminodial or pistillodial structures on the spadixdant or absent and all intermediate states exist.
(e.g. Areae, Bucephalandra) have less obvious Sometimes the endosperm is reduced at seed ma-
functions; in Arum the filamentous pistillodes and turity to a single cell layer. The embryo is usually
staminodes are thought to exclude inappropri- hardly differentiated. Rarely, the plumule is highly
ately large insect visitors and in Dieffenbachia, the
developed, e.g. as in Peltandra and Typhono-
staminodes are food bodies (Young 1986). In dorum. In Cryptocoryne ciliata, which grows
Helicodiceros the spathe and spadix hairs may act in freshwater tidal zones, between 20 and 40
to give the inflorescence the appearance of a mam- cataphylls are formed in the embryo and these
malian corpse. probably serve to fix the seed to the substrate,
Observations have been made of various sec- preventing it from being swept away with the tide.
ondary associations between insects and aroid in- Seeds with well-developed plumules, except
florescences, which do not involve pollination. Cryptocoryne, usually lack endosperm and have
Some species of drosophilid flies are known to only a very thin, papery testa. The testa may be
breed on the inflorescences of Alocasia, Colocasia smooth or variously sculptured, thin to thick or
and Homalomena (Okada 1986). They also exhibit occasionally absent (Gymnostachys, Nephthytis,
specialisation in their behaviour on the spadix: Orontium). In some Lasieae the testa is strongly
stamenicolous species lay their eggs in the male warty, hard and thickened other genera (Areae)
zone, pistilicolous species in the female zone. Fur-have strophiolate seeds, while in Pistia the seed
thermore, several different pairs of fly species, one
has a true operculum.
stamenicolous and the other pistilicolous, are The Araceae have the most varied endosperm
known to breed in association on one aroid spe- of any family. Genera conservative in this re-
cies (synhospitality). Hymenopteran parasitism spect, such as Epipremnum and Spathiphyllum,
in Philodendron inflorescences has also been possess abundant starch-free endosperm rich
observed (Mayo 1989). in lipids and aleurone. Accumulation of starch
is a feature of many of the specialised genera,
FRUIT AND SEED (see Seubert 1993). The fruits of beginning with Anthurium, where the endo-
the Araceae are usually juicy berries (more rarely sperm cells contain both lipids and starch grains,
rather dry and firmly carnose or leathery), densely leading to pure starch endosperms (e.g.
packed into a cylindric (sometimes subspherical) Xanthosoma), and finally to the "cereal type
infructescence. The berries are normally free, but endosperm" differentiated into a peripheral
in Syngonium they form an indehiscent syncarp, aleurone layer sharply delimited from the lipid-
and in Cryptocoryne a syncarp which opens free starchy remainder, as in Areae and Pistia.
apically, releasing the seeds. The berries are usu- Another trend leads to reduction and finally
ally indehiscent and may decay at maturity, but loss of endosperm, as in Lasia and Lysichiton,
the ripe berries of Lagenandra actively open at the where the endosperm is reduced to a single cell
base to release the seeds. The colour may be vivid layer, or Dieffenbachia, Orontium and Pothos,
orange (Nephthytis) to red (Arum) or purple-red where no trace of endosperm is left in the ripe
(Sauromatum) or white (many Philodendron spp., seeds. In a few genera (e.g. Anthurium, Gymno-
most Stenospermation spp., some Biarum spp.), stachys, Zomicarpa) the endosperm is chloro-
sometimes green (Arophyton buchetii, Lysichi- phyllous, the embryo more frequently so.
ton, Orontium, Peltandra virginica, Pinellia) or Chlorophyllous embryos occur both in genera
yellow (Typhonodorum), rarely blue (Gymno- with storage embryos (i.e. seeds lacking en-
stachys, Amorphophallus yunnanensis) or brown- dosperm, as in Anaphyllum and Pothos) and in
ish (!asarum). genera with a well-developed endosperm (e.g.
The seeds are often embedded in a mucilaginous Ambrosina, Arisarum, Calla, Dracunculus).
pulp secreted by the ovular and placental Gymnostachys is unique in having both green
trichomes. In many epiphytic Anthurium species endosperm and an embryo.
 Araceae 35

DISPERSAL. Dispersal of the genera with red- or west to E Africa and east to N America.
orange-coloured berries is probably carried out Amorphophallus, Remusatia, Rhaphidophora and
mainly by birds, though data are scarce. Mature Sauromatum are shared between tropical and
berries have been observed to stay fresh for a subtropical Africa and Asia. Amorphophallus
relatively long time while awaiting dispersal and Remusatia also occur in Madagascar.
(Arum, Nephthytis). In many genera (Cala- Pathos is found in Madagascar as well as in
dium, Colocasia, Philodendron, Syngonium), tropical Asia and NE Australia. Lysichiton and
the infructescences are markedly ephemeral, Symplocarpus occur in NE Asia as well as in tem-
disappearing almost as soon as they are ex- perate N America. Calla and Lysichiton extend
posed. Dispersal by ants (myrmecochory) has northwards to the subarctic zone. An unusual
been observed in Biarum and some species of geographical pattern is represented by the tribe
Amazonian Araceae (e.g. Anthurium ernestii, Peltandreae, which consists of Peltandra (eastern
Philodendron megalophyllum) are reported to N America) and Typhonodorum (Madagascar).
be associated with ant-gardens. Dispersal by Perhaps the most interesting feature of aroid
water (hydrochory) is also known (Cryp- geography is the distribution of three genera
tocoryne, Lagenandra, Montrichardia, Pistia, Homalomena, Schismatoglottis and Spathiphyl-
Typhonodorum). Dispersal by mammals, includ- lum which are disjunct between tropical SE Asia or
ing primates, has been observed (e.g. Anchomanes Melanesia on the one hand and tropical America
difformis - chimpanzee; Philodendron bipin- on the other. These genera are rainforest herbs
natijidum monkeys, bats; Xanthosoma whose long-distance dispersal by water or far-
robustum - bats). The seeds of Araceae, particu- ranging animal vectors is implausible. It is also
larly those with large chlorophyllous embryos, difficult to imagine them being rafted by the
do not remain viable for a long time. Bulbils southern Gondwanic route via Antarctica or via
with hooks or hairlike processes are found in the Bering Straits because of their intolerance of
Remusatia, which must be dispersed by birds or subtropical conditions, much less temperate cli-
mammals. mates. No really plausible historical explanation
A number of species which are important as has yet been proposed for these disjunctions. It is,
human food (Alocasia macrorrhizos, Colocasia nevertheless, tempting to speculate that these pat-
esculenta, Cyrtosperma merkusii, Xanthosoma terns are the relicts of a once-continuous distribu-
sagittifolium complex) or ornamentals (Epiprem- tion during the Cretaceous.
num pinnatum 'Aureum', Monstera spp., Philo-
dendron spp., Syngonium spp., Zantedeschia FOSSIL RECORD. Gregor and Bogner (1984, 1989),
aethiopica) have been widely dispersed through- Herendeen and Crane (1995) and Mayo et al.
out the tropics by man and have become (1997) have reviewed the literature on Araceae
naturalised as well. Some Typhonium species, fossils. Fossil leaves, spadices and infructescences,
possibly dispersed widely because of their medici- fruits and seeds are well documented from the
nal value, have become weedy in many tropical Eocene onwards; Spathiphyllum-like pollen has
regions. been described from the Palaeocene.

DISTRIBUTION AND HABITATS. The genera of EcoLOGY AND LIFE FoRMS. The Araceae include
Araceae are concentrated in the tropics of a wide range of life forms: (1) terrestrial plants
America and SE Asia. Tropical and southern with aerial stems; (2) geophytic plants with sub-
Africa is the next richest region, followed terranean stems modified into subglobose tubers
in order of decreasing diversity by temperate (sometimes cormlike) or thickened rhizomes; (3)
Eurasia, Madagascar and the Seychelles, and N root climbers, epiphytes, hemiepiphytes and litho-
America (including northern Mexico). Australia phytes; and (4) aquatic and semiaquatic plants.
has only one endemic genus (Gymnostachys). The least specialised life form appears to be that
The other native Australian Araceae represent shown by terrestrial rainforest genera such as
an extension of the tropical Asian araceous Aglaonema, Chlorospatha, Dieffenbachia, Homa-
flora. lomena, Schismatoglottis and Steudnera. The stem
The majority of genera are endemic to the major of such plants is aerial, decumbent or erect, with
continental regions, but some extend further; short, green internodes. In some other genera,
Pistia is pantropical and Calla is circumboreal. certain species may exhibit this habit. e.g. Anthu-
Arisaema is most diverse in SW China and extends rium, Culcasia.
36 Araceae

About 1/3 of the genera of Araceae prefer wet to Some Araceae grow in association with rocks
swampy conditions, or grow near open water. (lithophytes, epilithic or saxicolous plants), with
Among the most specialised aroids are the their stems, rhizomes or tubers growing in humus
rheophytes, which are flood-resistant plants grow- deposits in holes or crevices of steep outcrops (e.g.
ing up to the flood level in or along swift-running Colocasia gigantea, Typhonium albispathum in
streams or rivers, a habitat in which competition limestone hills in Thailand and the Malay Penin-
from other plants is low. They typically have sula) or in eroded limestone cavities in Mada-
narrow, leathery leaves and short stems, cling- gascar (Amorphophallus ankarana, Colletogyne),
ing tightly to rocks. All species of the genera or on granitic or sandstone mountains in Eastern
Aridarum, Bucephalandra, Hottarum and Pipto- Brazil (thick- and brittle-leaved Anthurium
spatha are true rheophytes while other genera species such as A. erskinei). Limestones, with their
contain a few species which are true or facultative eroded cavities, are especially favourable for the
rheophytes, as in Homalomena and Schis- collection of vegetable debris and leaf litter, and
matoglottis. Araceae often prefer these habitats.
]asarum steyermarkii and many Cryptocoryne The root climbers and hemiepiphytes normally
species are rhizomatous, permanently submerged grow in humid forests, often very high in the
hydrophytes. In flower, the inflorescence as a canopy. When losing connection with the forest
whole (]asarum) or the tip of the spathe floor, they may send down long roots to the
(Cryptocoryne) is held above the water surface. ground; such plants may be referred to as
Pistia stratiotes is the only free-floating aroid. secondary hemiepiphytes. Hemiepiphytes and
Many Araceae are helophytes (emergent rooted root climbers cling tightly to the bark of the host
aquatics or swamp plants), which show a great with negatively phototropic roots. Skototropism,
range of habit, including rhizomatous, creeping, an orientation movement towards a well-defined
arborescent or with simple, erect, aerial stems. dark area, has been observed in Monstera tenuis
Typhonodorum, Lasiomorpha and Montrichardia and other genera and is an adaptation in juvenile
form extensive pure stands along rivers and in plants for seeking a suitable host tree, following
freshwater lagoons. Aglaodorum griffithii occurs germination on the forest floor.
in the freshwater tidal zone in both fresh and True epiphytes are known in Anthurium, Philo-
brackish water. dendron and Stenospermation. Here the plants are
Many of the geophytic species grow in decidu- always independent of the ground, their seeds ger-
ous forests or woodlands in seasonally dry cli- minating directly on the host tree after dispersal,
mates, but some also grow in rainforests (e.g. probably by birds. The bird's nest epiphytes of
many Amorphophallus and Dracontium spp.). Anthurium sect. Pachyneurium and in the genus
Only one species of Araceae is known from true Philodendron (e.g. P. insigne) collect leaflitter and
desert conditions (Eminium spiculatum subsp. humus in their leaf rosettes and root balls.
negevense), but several genera are represented in Remusatia vivipara is a common epiphyte in the
semiarid areas, e.g. Arum and Eminium inC Asia, Old World, apparently dispersed by its hooked
Arum and Biarum in N Africa and the Orient, bulbils.
Arisaema and Sauromatum venosum in the Ara-
bian Peninsula and E Africa, and a few species of CoNSERVATION. The main threat to the long-
Stylochaeton from the Sahel zone of Africa. In the term survival of many Araceae species is the loss
northern temperate region, the tuberous life form and reduction in quality of their natural habitats,
enables genera like Arisaema, Arum and Pinellia especially the rainforests of tropical Asia, Africa
to survive the winter by means of dormancy. and America. Some Araceae particularly highly
These genera usually grow in woodlands or in adapted to specific habitats cannot survive
shady places under trees or shrubs. A few tuber- the transformation of their environments; e.g.
ous genera are found at high altitudes in tropical Chlorospatha and rheophytic species of the tribe
regions where warm temperate climates exist, Schismatoglottideae. These taxa have proved diffi-
as, for example, in the Andes (Gorgonidium to cult to cultivate and are unlikely to survive in the
ca. 3000m) or African mountains (Arisaema long term in botanic gardens. This is also true
ruwenzoricum up to 3000m in the Ruwenzori). of forest aquatics such as ]asarum (found only
The highest altitude is reached in the Himalayas, in blackwater rivers) and several species of
where Arisaema lobatum and A. jacquemontii Cryptocoryne. Currently, no species of Araceae is
grow up to 4500 m and A. flavum to 4400 m. listed in CITES (Convention on International
Trade in Endangered Species).
 Araceae 37

EcoNOMIC IMPORTANCE. Colocasia esculenta Anthurium, Caladium, Dieffenbachia, Epiprem-

(taro), with edible tubers and leaves, is the major num, Monstera, Philodendron, Spathiphyllum,
traditional staple food of the Pacific Islands and an Syngonium and Zantedeschia. Some large and
important subsistence food crop throughout the especially beautiful tropical species have been
tropics. Xanthosoma sagittifolium (cocoyam; important for modern garden design in the trop-
treated here in the broad sense including various ics, particularly in the work of the famous Brazil-
races and possibly species), originally from tropi- ian landscape architect, Roberto Burle Marx.
cal America, is grown throughout the tropics Anthurium andraeanum, Zantedeschia aethiopica
for its tuberous stems which are even more nutri- and their hybrids are an important source of cut
tious than those of Colocasia. Amorphophallus flowers, and the former is the basis of an impor-
paeoniifolius (elephant yam) is cultivated for its tant industry in Hawaii, the Netherlands and else-
starch-rich tubers in tropical Asia, particularly where. Several species of Cryptocoryne are much
India. Amorphophallus konjac (konjac) is impor- prized as aquarium plants. In general, it may be
tant in Japan, where "Konnyaku", a tasteless gel- said that the Araceae are probably the most im-
ling agent much used in food, is made from the portant foliage ornamental plant group in horti-
glucomannan-rich tubers. Cyrtosperma merkusii culture, worldwide. A fuller account of economic
is grown from SE Asia to Oceania as a famine and medicinal aspects of Araceae is given by Bown
food or for animals, and likewise Alocasia {1988).
macrorrhizos. Typhonodorum lindleyanum seeds
are cooked and eaten in Madagascar, and SUBDIVISION AND RELATIONSHIPS. Gymno-
Montrichardia seeds are roasted and eaten by stachys differs markedly in from all other Araceae.
the Amerindians of tropical South America. The Spathiphylleae are conservative morpho-
Many Araceae are poisonous when fresh, and in logically and the Orontioideae and Lasioideae
virtually all edible species, the stems, rhizomes, have many primitive characters, notably bisexual,
tubers and leaves must be cooked before they perigoniate flowers. Cladistic analyses based
can be consumed (Alocasia, Amorphophallus, on morphological and molecular data (French
Colocasia, Cyrtosperma, Xanthosoma). In classical et al. 1995; Mayo et al. 1997) showed that all
Greece the starchy tubers of Arum italicum genera with unisexual flowers form one mono-
were used as food and in the Middle Ages Arum phyletic group, recognised here as subfamily
maculatum tubers were used as famine food Aroideae. The new classification of Mayo et
in England. The infructescences of Monstera al. {1997), based on these studies, is adopted
deliciosa are delicious when fresh, with a here.
pineapple-like flavour.
The frequent use of numerous different Araceae AFFINITIES. The affinities of the Araceae have
by indigenous people in folk medicine, as arrow been considered obscure until recently. Compre-
poisons etc. may indicate the rich occurrence in hensive reviews by Grayum {1984-1992) and the
the family of biodynamic substances. cladistic studies of French et al. ( 1995) and Mayo
Aerial roots of Heteropsis and Philodendron are et al. {1997) suggest the following as ancestral
or have been a source of fibre for Amerindians in characters for Araceae: laticifers absent, raphides
tropical South America, the former being particu- present; the stem short, upright, unspecialised
larly highly regarded. Even today, roots of either as a climber or a geophyte; leaves pro-
Heteropsis species are used for basket-weaving bably simple, bifacial, with an expanded blade
in Amazonia. (?distichously arranged) and a geniculate petiole;
The Araceae include many very handsome orna- finer leaf venation may have been either reticulate
mental plants and are cultivated everywhere. In or parallel-pinnate; inflorescence a spike (axis not
temperate regions, the genera Arisaema and necessarily fleshy), subtended by a relatively in-
Arum, and to a lesser extent Biarum, Lysichiton, conspicuous bract (spathe); flowers bractless, bi-
Orontium and Symplocarpus, provide much-loved sexual, 3-merous (?2-merous), with 2 whorls of 3
perennial herbs for woodland, rock and bog (?2) free tepals, 2 whorls of 3 (?2) free stamens,
gardens. Arisaema species are particularly valued anthers terminal, basifixed, extrorse, on ± elon-
for this purpose in China and Japan and are. gated filaments, thecae dehiscing by longitudinal
increasingly sought after in Europe as woodland slits, tapetum probably periplasmodial, pollen
garden plants. Some aroids are ubiquitous monosulcate and ellipsoid, starchless, binucleate;
house plants and in the tropics are also grown ovary superior, syncarpous, 3-locular, with several
outside in gardens, e.g. species of Aglaonema, anatropous ovules per locule and axile placenta-
38 Araceae

tion; the seeds have endosperm with cellular Clear evidence of the "monocot type" was
of free-nucleate development in the micropylar recognised in the floral structure, but an inde-
chamber and a 1-celled haustoria! chalazal pendent evolution from a primitive monocot form
chamber, embryo small with caryophyllad or was regarded as the best explanation. Engler
solanad embrogeny; base chromosome number (1920a) went so far as to state that the most primi-
x = 7 or 14 (or 12). Following the proposal by tive Araceae (his subfamily Pothoideae) differed
Tillich (1985) and the reviews by Grayum (1987, from the Liliaceae essentially only in having a
1990), Acorus has been removed from Araceae fleshy outer seed integument. In his classification,
to its own family. Strong support for this change however, Engler (e.g. Engler and Gilg 1919) did
has come from the molecular studies of Duvall not associate Araceae and Liliaceae closely and
et al. (1993) and Chase et al. (1995), who placed it is reasonable to suppose that he saw the simi-
it at the base of the monocot clade. It differs larities between them in terms of shared primi-
from Araceae in a number of important char- tive characters (plesiomorphies). Hutchinson
acters: unifacial (ensiform) leaves, specialised (1934) visualised a close relationship with the
idioblasts containing essential oils, 2 separate Liliaceae. He considered that Araceae evolved
vascular systems in the peduncle, introrse anthers from tribe Aspidistreae of Liliaceae (his circum-
(in Araceae known only in Zamioculcas), stellate scriptions) due to their common possession of
pattern of endothecial thickenings, distinct mor- densely spicate inflorescences. Araceae have also
phology of ovular trichomes and their location been associated in the past with Pandanaceae
on the outer and inner integuments, presence of and Typhales.
perisperm, absence of raphides, secretory anther Dahlgren and coworkers (Dahlgren and Clifford
tapetum, unique bud trace insertion pattern 1982; Dahlgren et al. 1985) argued for a closer
and absence of dorsal vascular bundles in the relationship of Arales (Araceae, Lemnaceae) to the
gynoecium. Alismatiflorae than to any other group, pointing
The Lemnaceae have been considered by most out many important differences from the palms.
authors to be closely related to Araceae and to The characters that best link the Alismatiflorae to
have evolved from them by neotenous reduction. the more primitive tribes of Araceae appear to
This view was based mainly on the similarity be extrorse anthers, periplasmodial tapetum
between Pistia seedlings and Spirodela, embryo- and caryophyllad embryogeny. The Alismatiflorae
logical characters, the putative homology between also differ, however, in important respects:
the aroid spathe and the "spathe" of Spirodela absence of raphides, usually apocarpous gyno-
and Lemna, presence of grooved raphides, oper- ecium, helobial or nuclear endosperm develop-
culate seeds and a similar pattern of germina- ment, non-berried fruits and usually absence of
tion in Pistia and Lemnaceae. More recently, new endosperm.
lines of evidence confirming the close link be- Grayum (1984) agreed with Dahlgren's views.
tween the two families have emerged. French et His re-interpretation of aroid endosperm devel-
al. (1995), in a detailed cpDNA study, showed that opment as helobial further emphasises a relation-
Lemnaceae are embedded within the Araceae, ship with Alismatiflorae. Recent molecular studies
while the fossil genus Limnobiophyllum (Stockey by Chase et al. (1995) have further strengthened
et al. 1997) appears to bridge the gap between the link between Arales and Alismatiflorae. Their
Araceae and Lemnaceae. There is a growing body cladograms show the two taxa as sister groups,
of opinion that favours the formal reduction of forming a well-supported clade near the base of
Lemnaceae to a subgroup of Araceae (e.g. Mayo the monocotyledons.
et al. 1997). The pollen morphology differs signifi-
cantly, however; in Lemnaceae the pollen is ulcer- CLASS! FICA TION OF THE ARACEAE
ate and spinose while in Pistia it is inaperturate
and plicate. a. Flowers bisexual
I. Subfamily Gymnostachydoideae Bogner & Nicolson
Earlier authorities, such as Engler and Gilg (1991).
(1919) and Wettstein (1935), grouped the Araceae (genus 1)
together with the Palmae and Cyclanthaceae. This II. Subfamily Orontioideae Mayo, Bogner & Boyce (1997).
view was based on the common occurrence in (genera 2-4)
these families of a tendency to floral reduction, III. Subfamily Pothoideae Engler (1876).
1. Tribe Potheae Engler (1876).
"spathe" development, the condensation of the (genera 5-7)
inflorescence into a spadix or spadices and an as- 2. Tribe Anthurieae Engler (1876).
sociated tendency to pseudanthial inflorescences. (genus 8)
 Araceae 39

IV. Subfamily Monsteroideae Engler (1876). 26. Tribe Pistieae Blume (1836).
1. Tribe Spathiphylleae Engler (1887). (genus 105)
(genera 9-10)
2. Tribe Anadendreae Bogner & French (1984).
KEY TO THE GENERA OF ARACEAE
(genus 11)
3. Tribe Heteropsideae Engler (1906). AND AcoRACEAE
(genus 12)
1. Plants free-floating aquatics; leaves rosulate, hairy;
4. Tribe Monstereae Engler (1876).
flowers unisexual, naked; inflorescence with a single
(genera 13-20)
female flower and a few male flowers 104. Pistia
V. Subfamily Lasioideae Engler (1876).
- Plants terrestrial or helophytes, climbers, hemiepiphytes,
(genera 21-30)
epiphytes or lithophytes or other but never floating 2
VI. Subfamily Calloideae Endlicher (1837).
2. Leaves not differentiated into petiole and blade, primary
(genus 31)
venation strictly parallel; inflorescence borne on a
b. Flowers unisexual culmlike axis 3
- Leaves with distinct petiole and expanded blade, primary
VII. Subfamily Aroideae Engler (1887). venation never strictly parallel 4
1. Tribe Zamioculcadeae Engler (1876). 3. Leaves ensiform, unifacial; spadix solitary, pseudolateral
(genera 32-33) and overtopped by a single, erect, leaflike spathe; flowers
2. Tribe Stylochaetoneae Schott (1856). 3-merous, tepals 6 Acorus (Acoraceae)
(genus 34) - Leaves dorsiventrally flattened, bifacial; flowering shoot
3. Tribe Dieffenbachieae Engler (1876). with long culmlike axis, bearing numerous spadices dis-
(genera 35-36) tally, these borne in axillary clusters subtended by elon-
4. Tribe Spathicarpeae Schott (1856). gate bracts; flowers 2-merous, tepals 4 1. Gymnostachys
(genera 37-44) 4. Flowers with obvious perigone of free or fused tepals
5. Tribe Philodendreae Schott (1856). (except Pycnospatha which lacks perigone, but has
(genus 45) dracontioid leaf, tuberous stem and boat -shaped, forni-
6. Tribe Homalomeneae (Schott) M. Hotta (1970). cate spathe - see lead 22) 5
(genera 46-47) - Flowers without perigone of free or fused tepals 27
7. Tribe Anubiadeae Engler (1876). 5. Flowers bisexual, spadix uniform in appearance with
(genus 48) flowers of only one type 6
8. Tribe Schismatoglottideae Nakai ( 1943). - Flowers unisexual, spadix clearly divided into basal
(genera 49-55) female zone and apical male zone; tropical Africa 25
9. Tribe Cryptocoryneae Blume (1836). 6. Higher-order leaf venation parallel to primary lateral
(genera 56-57) veins; tissues with abundant trichosclereids 7
10. Tribe Zomicarpeae Schott (1856). - Higher-order leaf venation clearly reticulate; tissues
(genera 58-61) without trichosclereids or trichosclereids very few 8
11. Tribe Caladieae Schott (1832). 7. Spathe persistent; tepals free or connate; ovary 2-4-
(genera 62-68) locular; ovules 2-8 per locule, placenta axile
12. Tribe Nephthytideae Engler (1887). 9. Spathiphyllum
(genera 69-71) - Spathe deliquescent; tepals connate; ovary 1-locular;
13. Tribe Aglaonemateae Engler (1876). ovules several, placenta basal 10. Holochlamys
(genera 72-73) 8. Stem aerial, not tuberous or rhizomatous, never aculeate;
14. Tribe Culcasieae Engler (1887). plant usually a climber, hemiepiphyte or epiphyte, less
(genera 74-75) often lithophyte or terrestrial, only very rarely helophytic
15. Tribe Montrichardieae Engler (1876). (some spp. of Anthurium) 9
(genus 76) - Stem typically subterranean, tuberous or rhizomatous,
16. Tribe Zantedeschieae Engler (1887). sometimes aerial and creeping or scrambling, but then
(genus 77) aculeate; plant frequently a helophyte 13
17. Tribe Callopsideae Engler ( 1906). 9. Neotropical plants; seeds with copious endosperm;
(genus 78) pollen usually forate, never monosulcate 8. Anthurium
18. Tribe Thomsonieae Blume (1837). - Palaeotropical plants; seeds without endosperm; pollen
(genera 79-80) monosulcate or inaperturate 10
19. Tribe Arophyteae Bogner (1972). 10. Stigma transversely oblong; stamens always 4 per flower;
(genera 81-83) pollen inaperturate; perigone consisting of a single
20. Tribe Peltandreae Engler ( 1876). cuplike structure 11. Anadendrum
(genera 84-85) - Stigma hemispheric to discoid; stamens usually 6 per
21. Tribe Arisareae Dumortier (1827). flower; pollen sulcate; perigone usually consisting of free
(genus 86) tepals or when connate and cuplike the flowers are borne
22. Tribe Ambrosineae Schott (1832). on short pedicels 11
(genus 87) 11. Ovary 3-locular; locules 1-ovulate; flowering shoot with
23. Tribe Areae Engler (1876). inflorescences always axillary 12
(genera 88-96) - Ovary 1-locular; flowering shoots terminating in a
24. Tribe Arisaemateae Nakai (1943). branching system of spadices 7. Pothoidium
(genera 97-98) 12. Flowers sessile; tepals free, very rarely basally united
25. Tribe Colocasieae Engler (1876). 5. Pothos
(genera 99-104) - Flowers pedicellate; tepals connate 6. Pedicellarum
40 Araceae

13. Plants of temperate regions (N America, NE Asia); leaf flowers of only one type (sometimes with sterile flowers
blade always entire, ovate to elliptic 14 at spadix base) 28
- Plants of tropical and subtropical regions; leaf blade - Flowers unisexual; spadix clearly divided into basal
sagittate, pinnatifid, pinnatisect or dracontioid 16 female zone and apical or intermediate male zone,
14. Ovary 2-locular; ovules 2 per locule, placentation axile flowers very rarely in longitudinal rows (Spathicarpa) 37
3. Lysichiton 28. Helophytes from temperate regions of northern hemi-
- Ovary 1-locular; ovule 1, placenta apical or basal 15 sphere; petiole sheath with long apical ligule 31. Calla
15. Placentation basal; spathe inconspicuous; spadix cylin- - Climbers, hemiepiphytes or sometimes epiphytes from
dric, stipe very long 2. Orontium tropical regions; petiole sheath non-ligulate or ligule only
- Placenta apical; spathe thick, ventricose, enclosing ~~ ~
spadix; spadix subglobose, stipe short 4. Symplocarpus 29. Petiole usually very short with non-annular insertion;
16. Leaf deeply sagittate, anterior division not pinnatifid or trichosclereids not present in tissues, leaf never per-
pinnatisect 17 forated or lobed; primary lateral veins forming distinct
- Leaf blade pinnatifid, pinnatisect, dracontioid or some- submarginal vein 8. Heteropsis
times :±:: pedatifid; anterior division always pinnately - Petiole well developed with annular insertion and usually
divided, either pinnatifid, pinnatisect or yet more highly conspicuous sheath; trichosclereids present in tissues,
divided 21 or if absent (or nearly so) then leaf with conspicuously
17. Ovary many- to 2-ovulate, rarely 1-ovulate; seeds with reticulate higher-order venation and often perforated or
endosperm 18 lobed (Amydrium); primary lateral veins usually not
- Ovary 1-ovulate, rarely 2-ovulate; seeds without en- forming distinct submarginal vein 30
dosperm or rarely with a little endosperm 19 30. Trichosclereids rare or nearly absent; higher-order leaf
18. Plants without stolons; petiole spines dispersed; stamen venation completely reticulate; ovary 1-locular, placenta
filaments free; tropical Asia to Oceania 26. Cyrtosperma 1, intrusive-parietal, ovules 2 13. Amydrium
- Plants with long stolons; petiole spines in ridges; stamen - Trichosclereids abundant; higher-order leaf venation
filaments free or connate; tropical W Africa parallel to primary lateral veins, or only finest venation
27. Lasimorpha reticulate 31
19. Petiole aculeate, with obvious spines; Malay Archipelago 31. Ovary 1-locular or incompletely 2-locular 32
28. Podolasia - Ovary 2-5 locular 35
- Petiole smooth to scabrid-verrucose, never aculeate; 32. Ovules anatropous, more than 1 33
tropical America 20 - Ovules amphitropous to anatropous, solitary, basal 34
20. Leaf blade never perforated; spathe lanceolate, very long- 33. Ovules numerous, superposed on 2 (rarely 3) parietal
acuminate and usually spirally twisted; ovary locules with placentas; seeds fusiform, straight, 1.3-3.2mm long,
(1- )2 to several ovules; neotropics but not east coastal 0.6-l.Omm wide 14. Rhaphidophora
Brazil 30. Urospatha - Ovules 2-4 (-6) at base of a single intrusive placenta;
- Leaf blade often with a few perforations of irregular size seeds curved, 3-7mm long, 1.5-4.0mm wide
between primary lateral veins; spathe fornicate; endemic 15. Epipremnum
to east coastal Brazil 22. Dracontioides 34. Adult leaf blade entire; palaeotropical 16. Scindapsus
21. Stem aculeate, aerial and scrambling to prostrate, intern- - Adult leaf blade pinnatifid; neotropical (Amazonia)
odes distinct, green 29. Lasia 18. Alloschemone
- Stem not aculeate, subterranean, internodes very abbre- 35. Seeds fusiform, claviform or lenticular, less than 3mm
viated, not green 22 long, endosperm present; ovules (2-)3-many per locule;
22. Leaf blade dracontioid, anterior division bipinnatifid or leaf blade entire 36
yet more highly divided; stem a depressed-globose tuber; - Seeds globose to oblong, 6-22mm long, the raphe S-
spathe fornicate 23 shaped; endosperm absent; ovules 2 per locule; leaf blade
- Leaf blade pinnatifid, pinnatisect, or sometimes :±:: variously shaped, often perforated or pinnatifid or both
pedatifid, anterior division pinnatifid to pinnatisect; stem 17. Monstera
a vertical or horizontal rhizome; spathe erect, not forni- 36. Placentation basal; seeds fusiform to claviform; leaf
cate, blade often spirally twisted apically 24 blade thickly coriaceous 20. Stenospermation
23. Tropical America; flowers with perigone of 4-8 free - Placentation axile; seeds lenticular and flattened, strongly
tepals; berries smooth, red 21. Dracontium curved; leaf blade mostly membranous
- Tropical SE Asia; flowers without perigone; berries ac- 19. Rhodospatha
uleate, dark green 24. Pycnospatha 37. Spadix fused laterally on both sides to spathe and entirely
24. Tropical America; testa thick, verrucose; embryo curved enclosed by it, forming a septum dividing the spathe into
23. Anaphyllopsis 2 chambers, with a single gynoecium on one side and the
- S India; testa membranous, smooth; embryo straight male flowers arranged in 2 rows on the other; very small,
25. Anaphyllum seasonally dormant plants endemic to western
25. Leaf pinnatisect to tri- or quadripinnatifid; tepals free; Mediterranean 87. Ambrosina
spathe margins free 26 - Spadix free or fused to spathe in various degrees but
- Leaf entire, linear to cordate, sagittate or hastate; tepals never fused laterally on both sides to spathe to form 2
connate into cup; spathe margins connate basally internal chambers with a single gynoecium on one side
34. Stylochaeton and the male flowers on the other 38
26. Leaf blade pinnatisect; stamens free 32. Zamioculcas 38. Stamens of each male flower free or only the filaments
- Leaf blade bipinnatifid to quadripinnatifid, at least connate 39
in lowest pinnae; stamen filaments connate - Stamens of each male flower entirely connate into a dis-
33. Gonatopus tinct synandrium, synandrium rarely reduced to single
27. Flowers bisexual; spadix uniform in appearance with stamen and then endemic to Madagascar ( Colletogyne) or
 Araceae 41

stamens free and basally connate with remote glo- 50. Stigma smaller than ovary; upper part of spadix sterile
bose thecae and then endemic to Andean S America with a distinct appendix of hornless sterile flowers;
(Gorgonidium mirabile), or only filaments connate and spathe constricted or not; stamens never excavated
then stigma stellate and 5-8-lobed (Spathantheum) 78 apically 51
39. Spadix never entirely enclosed by spathe in a basal - Stigma as broad as ovary; upper part of spadix mostly
"kettle" formed of connate spathe margins (or if spathe fertile to apex and without a distinct appendix; spathe not
margins basally connate, then plant never aquatic) 40 conspicuously constricted; stamens all or mostly exca-
- Spadix entirely enclosed by spathe in a basal "kettle" vated apically 52
formed of connate spathe margins, plants always 51. Spathe not constricted; male flowers smooth or verru-
helophytic or aquatic 77 cose; sterile flowers between male and female flowers
40. Higher-order leaf venation parallel-pinnate 41 flattened 52. Bucephalandra
- Higher-order leaf venation reticulate 53 - Spathe constricted; male flowers densely tuberculate;
41. Upper part of spathe persisting as long as lower part; sterile flowers between male and female flowers
petiole sheath lacking ligule; ovary !-many-locular; subcylindric 53. Phymatarum
thecae dehiscing by subapical pores or longitudinal slits; 52. Stamens all excavated; placenta basal 54. Aridarum
connective usually conspicuously thickened 42 - Two lateral stamens of each male flower excavated and
- Upper part of spathe marcescent or caducous at anthesis, thecae horned, central stamen truncate and thecae horn-
lower part long-persistent; petiole sheath with long, less; placentas basal (fertile ovules) and apical (appar-
marcescent ligule (except most Schismatoglottis spp.); ently sterile) 55. Heteroaridarum
ovary !-locular; thecae dehiscing by apical pores, connec- 53. Leaf blade dracontioid, leaf solitary in each growth
tive not conspicuously thickened 47 period 54
42. Spathe variously shaped, never campanulate; plants - Leaf blade shape of various types but never dracontioid;
tropical American or tropical Asian; peduncle usually usually several leaves present 57
short, if long, then female flowers in single whorl 43 54. Petiole usually aculeate; at least some of the ultimate leaf
- Spathe obconic to campanulate; plants from southern lobes trapezoid, truncate or shallowly bifid, veins not
Africa (naturalised in America and Asia); peduncle long, forming regular submarginal collective vein on each side
sometimes longer than leaves 77. Zantedeschia 55
43. Plant always terrestrial, rarely aquatic, never climbing or - Petiole usually smooth, sometimes rugose but never
epiphytic; inflorescences not secreting resin at anthesis; aculeate; ultimate leaf lobes usually oblong-elliptic,
endothecium with cell wall thickenings; ovary !-locular acuminate, with primary lateral veins forming regular
or incompletely 2-5-locular; mostly tropical Asian submarginal collective veins on each side 56
(except Homalomena sect. Curmeria) 44 55. Peduncle long; ovary !-locular 70. Anchomanes
- Plant usually climbers, hemiepiphytes or epiphytes; in- - Peduncle very short; ovary 2-locular
florescences secreting resin from spathe or spadix at an- 71. Pseudohydrosme
thesis; endothecium nearly always lacking cell-wall 56. Ovary 1-4 locular; terminal appendix smooth, rugose,
thickenings; ovary completely 2-many locular, placenta rarely verrucose, or staminodial (appendix absent in
axile to basal; tropical America 45. Philodendron Amorphophallus margaritifer and A. coudercii)
44. Seed without endosperm, embryo large; ovule 1, placenta 79. Amorphophallus
basal or parietal 45 - Ovary always !-locular; terminal appendix staminodial,
- Seed with copious endosperm, embryo relatively small; separated from male zone by naked axial region
ovules several to many, placenta basal, parietal or axile 80. Pseudodracontium
46 57. Spadix fertile to apex, terminal appendix absent 58
45. Inflorescence with short peduncle; female flowers in spi- - Spadix with :±: smooth terminal appendix 62
rals; stem erect to repent; placentation basal; forest plants 58. Helophytes with robust, erect stems and an apical crown
72. Aglaonema of sagittate to hastate (rarely trisect) leaves; tropical
- Inflorescence with long peduncle; female flowers in a America 76. Montrichardia
single whorl; stem repent; placenta parietal; on tidal - Terrestrial, climbing or epiphytic plants, leaf blade
mudflats 73. Aglaodorum variously shaped; tropical Africa 59
46. Male flower consisting of solitary stamen overtopped by 59. Leaf blade usually with pellucid resin canals (lines or
flask-shaped pistillode; ovary !-locular, placenta basal points); plants mostly climbers; hemiepiphytes; spathe
46. Furtadoa boat-shaped, convolute basally; anthers lacking endo-
- Male flower consisting of ( 1) 2-6 stamens, pistillodes thecial thickenings 60
absent; ovary incompletely 2-5 locular, placentas parietal - Leaf blade lacking pellucid resin glands; plants terrestrial;
and axile or nearly basal 47. Homalomena spathe :±: fully expanded, not convolute; anthers with
47. Placentation parietal; thecae truncate 48 endothecial thickenings 61
- Placentation basal or basal and apical; thecae truncate or 60. Laticifers absent; flagelliform shoots absent; leaf blade
horned 49 always simple, acute to rounded at base; ovary 1-3-
48. Spathe constricted; ovules anatropous to hemi- locular; spadix stipitate or sessile 74. Culcasia
anatropous; petiole sheath usually not ligulate; upper - Laticifers present; flagelliform shoots present; leaf blade
part of spadix usually sterile 49. Schismatoglottis oblong-lanceolate to cordate, sagittate, hastate, trifid or
- Spathe not constricted; ovules hemiatropous to atropous; laciniate to pinnatifid; ovary !-locular; spadix sessile
petiole sheath with long, marcescent ligule; spadix fertile 75. Cercestis
almost to apex SO. Piptospatha 61. Leaf cordate-sagittate or subtriangular, deeply sagittate
49. Thecae truncate; placentation basal 51. Hottarum or trifid, glabrous; spathe green; spadix entirely free of
- Thecae horned; placentation basal or basal and apical spathe 69. Nephthytis
so
42 Araceae

- Leaf cordate-ovate, minutely hispid abaxially; spathe - Ovary 1-ovulate; female zone of spadix adnate to spathe;
pure white; female zone of spadix adnate to spathe spathe usually with transverse septum between male and
78. Callopsis female zones 96. Pinellia
62. Laticifers anastomosing; tropical S America 63 76. Flowers of both sexes always present in a single inflores-
- Laticifers simple; temperate Eurasia and palaeotropics cence; male flowers 1-androus; lower spathe margins
66 connate; leaf blade ovate or sagittate 86. Arisarum
63. Ovary 6- to 9-ovulate; leaf blade trisect to pedatisect; stem - Flowers of both sexes present in a single inflorescence, or
a subglobose tuber 58. Zomicarpa more often with male and female flowers appearing on
- Ovary 1- to 6-ovulate; leaf blade cordate-sagittate; stem a separate inflorescences; male flowers 2-5-androus; lower
subglobose tuber or rhizome 64 spathe margins convolute; leaf blade normally trisect,
64. Appendix slender 65 pedatisect or radiatisect, rarely simple and ovate
- Appendix relatively thick and subcylindric; stem a creep- 97. Arisaema
ing rhizome; ovary 1-ovulate 60. Ulearum 77. Female flowers spirally arranged (pseudowhorl in
65. Stamen connective much prolonged, threadlike; stem Lagenandra nairii, whorled in L. gomezii) and free;
tuberous; ovary 1-ovulate 61. Filarum spathe tube kettle with connate margins occupying
- Stamen connective not at all prolonged; ovary 1-6- entire spathe tube; spathe blade opening by a straight or
ovulate; stem a creeping rhizome 58. Zomicarpella twisted slit; berries free, opening from base; leaf ptyxis
66. Spadix ± elongated, filiform to clavate with zone of involute 56. Lagenandra
sterile flowers between male and female zones, rarely - Female flowers in a single whorl, connate; spathe tube
with a naked axis between female and male zones of kettle occupying only lower part of spathe tube, remain-
spadix (Arum pictum) or with fertile zones contiguous der also with connate margins (except Cryptocoryne
(Dracunculus) 67 spiral is), blade spreading or twisted; berries connate into
- Spadix usually without sterile flowers (sometimes present a syncarp which opens from the apex; leaf ptyxis
in Arisaema) 75 convolute 57. Cryptocoryne
67. Placenta parietal; leaf blade sagittate or hastate 78. Laticifers simple 79
88.Arum - Laticifers anastomosing 95
- Placentas apical and/or basal; leaf blade variously shaped 79. Synandria connate, thecae of adjacent synandria encir-
68 cling pits in the spadix, each with a somewhat prominent
68. Placentas basal and apical 69 upper margin; leaf peltate; Myanmar to India
- Placenta basal 71 98. Ariopsis
69. Male zone of spadix contiguous with female zone; leaf - Synandria free; leaf not peltate; Africa, Madagascar or
blade pedatifid but lobes not spirally twisted up Americas 80
90. Dracunculus 80. Higher-order leaf venation parallel-pinnate or if reti-
- Male zone of spadix separated from female zone by subu- culate then stem a creeping rhizome and plant from
late to filiform sterile flowers; leaf blade variously shaped tropical S America (Bognera) 81
70 - Higher -order leaf venation reticulate; stem usually a
70. Appendix covered with subulate to setiform sterile flow- subglobose tuber, if rhizomatous then plant Madagascan
ers; leaf blade pedatifid, lobes twisting up on each side 85
91. Helicodiceros 81. Ovules anatropous; primary lateral veins ofleaf forming
- Appendix smooth; leaf blade oblong-lanceolate or a single marginal vein, no submarginal collective vein
sagittate-hastate 92. Theriophonum present; plant from tropical America or continental
71. Lower spathe margins free 72 tropical Africa 82
- Lower spathe margins connate for an appreciable dis- - Ovules atropous to hemiatropous; primary veins of leaf
tance (entirely free in Biarum aleppicum) 73 forming submarginal collective vein and 1-2 marginal
72. Infructescence borne above ground level, berries dark veins; plants from temperate eastern N America or
red to purple, pericarp juicy; sterile zone between male Madagascar 84
and female zones of spadix relatively long, often partially 82. Female zone of spadix free; plant from tropical W and
naked; tropical and subtropical to warm temperate Asia Central Africa 48. Anubias
to Australia 93. Typhonium - Female zone of spadix entirely adnate to spathe; plant
- Infructescence borne at or below ground, berries white to from tropical America 83
pale lilac, pericarp firm, not juicy; sterile zone between 83. Female flowers each with whorl of several staminodes;
male and female zones relatively short and covered en- higher-order leaf venation strictly parallel-pinnate
tirely with subulate sterile flowers; Turkey, western N 35. Dieffenbachia
Africa, Middle East, Central Asia 89. Eminium - Female flowers without staminodes; higher-order leaf
73. Leaf usually solitary, blade pedate; ovary 2-several- venation reticulate 36. Bognera
ovulate 94. Sauromatum 84. Giant herbs (to 4 m) with massive false stem of petiole
- Leaves several; blade linear to ovate, elliptic or obovate; sheaths; staminodes of female flowers free; Madagascar
ovary 1-ovulate 74 (also naturalised in Pemba, Zanzibar and Mascarene Is.)
74. Leaf blade broad elliptic, spathe tube septate 85. Typhonodorum
93. Typhonium pro parte - Relatively small herbs (less than 1m) without false stem;
- Leaf blade linear, ovate, elliptic-oblong or obovate; staminodes of female flowers connate into a cuplike
spathe tube not septate 95. Biarum synandrodium; eastern N America 84. Peltandra
75. Ovary several-ovulate; female zone of spadix free from 85. Madagascan; seed lacking endosperm; ovary 1-locular;
spathe; spathe without a transverse septum separating leaf venation with primary lateral veins forming submar-
male and female zones 76 ginal collective vein and 1-2 marginal veins on each side
86
 Araceae 43

- S American; seed with abundant endosperm; ovary with hastate or pedatifid 99

more than 1 locule (except Spathicarpa); leaf venation - Submerged aquatic; leaf blade linear 64. ]asarum
with primary lateral veins usually forming single 99. Pollen shed in tetrads; style usually laterally thickened
marginal vein on each side, submarginal collective veins or expanded into a diaphanous mantle; leaf blade entire
usually absent 88 or pedatifid 100
86. Stamens either completely connate with marginal thecae - Pollen shed in monads; stylar region not laterally ex-
or only partially connate by filaments; bisexual flowers panded; leaf blade entire or trifid 101
often present between male and female zones of the 100. Spathe tube subglobose, inflated; female zone of spadix
spadix 82. Carlephyton free; styles normally discoid (laterally swollen) and co-
- Stamens completely connate into truncate synandria or herent (except Xanthosoma plowmanii); synandrodes
synandria reduced to 1 stamen 87 (sterile flowers) between male and female flowers well-
87. Synandria reduced to 1 stamen, thecae apical on conical developed, :±:: prismatic 65. Xanthosoma
filament; spadix fertile to apex; leaf blade always cordate - Spathe tube narrow, elongate; female zone of spadix
83. Colletogyne mostly adnate to spathe; stylar region thin, spreading,
- Thecae apical on a truncate synandrium; spadix appen- diaphanous, mantlelike; synandrodes (sterile flowers)
dix present or not; leaf blade cordate, hastate, trifid, between male and female flowers usually irregular or
trisect or pedatisect 81. Arophyton fungiform, not prismatic 66. Chlorospatha
88. Spadix free or only female zone adnate to spathe 89 101. Spathe tube always convolute; stylar region as broad as
- Spadix usually entirely adnate to spathe (male zone free ovary (Caladium paradoxum has discoid, coherent stylar
in Spathantheum intermedium) 94 regions); synandrodes between male and female flowers
89. Ovules anatropous 90 well developed, prismatic; placentas 1-2(-3), parietal;
- Ovules atropous 92 seeds several (rarely 1-2) 63. Caladium
90. Ovules 2 per locule; leaf blade entire, linear to Spathe tube gaping widely at anthesis; style much nar-
subsagittate; spadix free, with terminal appendix of rower than ovary; synandrodes (sterile flowers) lacking,
synandrodes 37.~angonia male and female zones contiguous; placenta 1, basal; seed
- Ovules 1 per locule; leaf blade entire, pinnatifid to solitary 62. Scaphispatha
subdracontioid; spadix fertile to apex 91 102. Ovules more than 1; leaf blade pel tate 103
91. Leaf blade pinnatifid to bipinnatifid or subdracontioid; - Ovule solitary; leaf blade not peltate 68. Hapaline
synandria elongate; stigma capitate or lobed; staminodes 103. Female flowers with staminodes; spathe not constricted;
offemale flowers free (connate in Taccarum caudatum) stem trunk-like or creeping 100. Steudnera
38. Taccarum - Female flowers without staminodes; spathe with 1 or 2
- Leaf blade usually pinnatisect, rarely entire; synandria constrictions; stem tuberous, producing erect or spread-
short and domed; stigma deeply lobed; staminodes of ing stolons bearing small tubercles covered in hooked
female flowers free or connate 39. Asterostigma scales 101. Remusatia
92. Styles and synandria elongate 93 104. Leaf blade pedatisect; female flowers each with several
- Styles and synandria short and squat; staminodes in large staminodes 99. Protarum
female flowers obovate 42 . .Gearum - Leaf blade entire or pinnatifid; female flowers without
93. Staminodes in female flowers filiform to subclavate; staminodes (except single small ones in Colocasia
synandria with free filament apices or not; leaf blade esculenta) 105
pinnatifid, pinnatisect or bipinnatifid 40. Gorgonidium 105. Placentas parietal; ovules many; leaf blade always entire
- Staminodes in female flowers elongate-triangular; 102. Colocasia
synandria entirely connate; leaf blade ovate-emarginate - Placenta basal; ovules few; leaf blade entire or pinnatifid
to cordate 41. Synandrospadix 103. Alocasia
94. Ovary 6-8-locular; female flowers below, male above; leaf
blade entire or pinnately lobed 43. Spathantheum
- Ovary 1-locular; female and male flowers intermixed I. Subfam. Gymnostachydoideae Bogner &
(2 central rows of male flowers, 2 outer rows of female Nicolson (1991)
flowers); leaf blade entire 44. Spathicarpa
95. Plants climbers or hemiepiphytes, sometimes creeping Laticifers absent; stem a short rhizome; leaves dis-
on ground in submature growth, internodes long; berries tichous, linear, not differentiated into blade and
connate into a syncarp 67. Syngonium petiole, venation parallel; flowering shoot a com-
- Plants terrestrial or geophytic, rarely aquatic, not climb-
ing; internodes very short, berries free from each other plex synflorescence consisting of 3-7 successive,
96 short, perennating floral sympodia, borne on an
96. Spadix without an appendix (present in Hapaline erect peduncular axis, each sympodium axillary
appendiculata, included here, occasionally absent in to a bract and composed of numerous inflores-
Colocasia esculenta, excluded here) 97
cences; spathe inconspicuous; spadix long-
Spadix with an appendix (occasionally absent in
Colocasia esculenta); palaeotropical 104 stipitate; flowers bisexual, 2-merous, tepals 4,
97. Ovary completely to incompletely 2- to several-locular fornicate, stamens 4, thecae dehiscing by longitu-
with deeply intrusive parietal palcentas (1-locular with dinal slit; ovary 1-locular, ovule 1, atropous, pla-
basal placenta in ]asarum, Scaphispatha and a few species centa apical; berries deep blue, exserted beyond
of Caladium and Xanthosoma); neotropical 98
- Ovary clearly 1-locular, placentas not intrusive;
tepals; seed obovoid, testa absent, embryo axile,
palaeotropical 102 endosperm copious, starchy apart from peripheral
98. Helophytes or terrestrial; leaf blade ovate, sagittate to layer.
44 Araceae

@
1. Gymnostachys R. Br.
Gymnostachys R. Br., Prodr.: 337 (1810}.
~~
E~
Leaves basal, midrib distinct, margins erose-
serrate; spathe short, alternating with prophylls in
each floral sympodium; spadix cylindric, erect in F
flower, pendent in fruit. 2n = 48. Only one sp., G.
anceps R. Br., Australia (Queensland, New South
Wales), in moist hardwood forests.

II. Subfam. Orontioideae Mayo, Bogner &

P.C. Boyce (1997).

1. Tribe Orontieae R. Br. ex Dumort. (1827).

Laticifers absent (except Orontium); swamp
plants, helophytes; stem a stout, erect rhizome,
continuation shoot in axil of last leaf preceding
spathe; leaf blade entire, oblong-elliptic or cor-
date; flowers bisexual, with perigone of fornicate
tepals, stamens with distinct filament, anther ter-
minal, connective inconspicuous, thecae dehisc-
ing by longitudinal slits; ovary ± immersed in
spadix axis; testa smooth, embryo large, en-
dosperm absent in Orontium, reduced to 1 cell
layer in the other genera.

2. Orontium L.
Orontium L., Sp. Pl.: 324 (1753); Huttleston, Temp. N Am.
Arac.: 50-55 (1953) (unpubl. thesis).

Laticifers present, simple; leaf blade aerial or float-

ing; primary veins arcuate-parallel; inflorescences
Fig. llA-F. Araceae-Orontioideae. A-C Lysichiton camts-
1-2; peduncle suppressed; spathe inconspicuous, chatcensis. A Habit. B Inflorescence. C Flower, one tepal and
short, inserted at attachment to rhizome; spadix one stamen removed. D-F Symplocarpus foetidus . D Inflores-
stipe very long, swollen distally; spadix emergent, cence. E Flower, seen from above. F Flower, longitudinal
conical, slender, golden yellow; tepals (4-)6; section; note single ovule with apical placentation. (Takhtajan
stamens 6 or 4; ovary 1-locular, ovule 1, 1982)
hemianatropous, placenta basal; berry depressed-
globose, green; testa thin. 2n = 26. Only one sp., 0.
aquaticum L., temperate and warm temperate spadix stipe, distally boat-shaped, gaping; spadix
eastern N America, partly submersed. cylindric, stipe very long; tepals 4; stamens 4;
gynoecium elongate-ovoid, style distinct, atten-
uate, ovary 2-locular or incompletely 2-locular,
3. Lysichiton Schott Fig. 11A-C ovules atropous, 1-2 per locule, placenta axile,
Lysichiton Schott, Oesterr. Bot. Wochenbl. 7: 62 (1857); Hulten stigma capitate; seed ellipsoid or ovoid. 2n = 28.
& St. John, Sven. Bot. Tidsk. 25:453-464 (1931); Huttleston, Two spp., ·temperate E Asia and western N
Temp. N. Am. Arac.: 31-38 (1953) (unpubl. thesis). America, in swamps and wet woodlands.

Robust, rosulate; petiole flattened; primary veins

arcuate-parallel to subpinnate; inflorescence 1; 4. Symplocarpus Salish. ex Nuttall Fig. 11D- F
peduncle suppressed; spathe inserted at attach- Symplocarpus Salish. ex Nuttall, Gen. 1: 105 (1818), nom.
ment to rhizome, proximally narrow and clasping cons.; Lee, Ill. Fl. Korea: 183 (1989).
 Araceae 45

Robust, rosulate; leaf blade subcordate or cor- New Guinea, NE Australia; in tropical humid
date-ovate, primary veins arcuate-parallel to forests.
subpinnate; inflorescence 1(-2); peduncle short;
spathe thick, ventricose or conchiform, cucullate-
6. Pedicellarum M. Hotta
convolute below, open above; spadix short,
subglobose, stipe ± short; teplas 4, stamens 4; Pedicellarum M. Hotta, Acta Phytotaxon. Geobot. 27: 61
ovary 1-locular, ovule 1, subatropous, placenta (1976); Nicolson, Aroideana 7: 56-57 (1984).
apical-parietal, style long-attenuate; infructes-
cence ± globular, berries immersed in spongy Petiole sheath narrow, ± involute; leaf blade
axis; seed globose, testa thin. 2n = 30, 60. Three lanceolate to elliptic, venation pinnate, forming
spp., temperate E Asia and eastern N America, submarginal collecting veins; flowering articles
swamps and water courses. infra-axillary, with cataphylls basally; spathe
ovate; spadix axis flexuose; flowers sparse, distant,
on short pedicels, swollen below perigone into a
III. Subfam. Pothoideae Engler (1876). receptacle-like structure, tepals connate; thecae
latrorse; ovary 3-locular, placenta basal; berry
Laticifers absent; stem usually aerial; petiole obovoid, red, 1-3(?)-seeded; seed compressed-
geniculate apically; higher-order leaf venation re- ellipsoid. n = ? Only one sp., P. paiei M. Hotta,
ticulate; spathe simple, spreading to reflexed, not Borneo, in humid forests.
enclosing spadix; flowers bisexual, perigoniate;
stamen filaments distinct, anthers terminal, 7. Pothoidium Schott
thecae dehiscing by longitudinal slits, connective
inconspicuous. Pothoidium Schott, Aroideae 26 (1856-57); Oesterr. Bot.
Wochenbl. 7: 70 (1857).

1. Tribe Potheae Engler (1876). Petiole broad, flattened, apically rounded; leaf
Shrubby climbers with tough woody stems; main blade triangular-lanceolate, venation ± arcuate-
shoot monopodia!, flowering shoots axillary or parallel; flowering shoot terminating in a branch-
infra-axillary; leaves distichous; petiole sheath ing system of spadices; spathe lanceolate, short,
long and often broad, flattened and apically au- or lacking entirely in the most apical inflores-
riculate (reduced in Pedicellarum and Pothos se- cences; spadix cylindric, stipitate; ovary 1-locular,
ries Goniuri); flowers (2-)3-merous; tepals (4-)6, placenta parietal-basal; berry ovoid, acute;
unthickened and without truncate apices, stamens seed oblong, testa smooth. 2n = 24. Only one
6; ovules 1 per locule, anatropous, stigma sessile, sp., P. lobbianum Schott, Philippines, Celebes,
usually umbonate; embryo large, endosperm Moluccas, Taiwan; in humid forests.
absent.
2. Tribe Anthurieae Engler (1876).
5. Pothos L. Sympodial articles composed of prophyll, cata-
Pothos L., Sp. Pl.: 968 (1753).
phyll, foliage leaf, inflorescence (often aborted);
mature petiole with very short sheath, geniculate
Lower branches of stem rooting, the upper apically and basally; primary lateral veins pinnate
ones free; petiole sheath broad, flattened and (rarely arcuate-parallel), usually forming sub-
auriculate apically or much narrower and ± marginal collective veins; spathe usually simple,
involute, leaf blade linear- to ovate-lanceolate; spreading, reflexed or erect; spadix usually
flowering articles axillary or infra-axillary, tapering to subcylindric; flowers bisexual, with
usually with basal cataphylls; peduncle some- perigone of 4 fornicate tepals, stamens 4, filaments
times reflexed; spathe ovate to linear; spadix distinct, anther terminal, connective inconspicu-
globose, ellipsoid or narrowly cylindric, sessile ous; ovary 2-locular, locules 1-2-ovulate (rarely
to long-stipitate; tepals ± fornicate; ovary more), ovules anatropous to campylotropous,
3-locular, placenta axile; berry ellipsoid or abo- placenta axile-subapical; mature berry exserted
void, red, 1-3-seeded; seed compressed -ellipsoid, and usually dangling; seed often sticky, endo-
testa thickish, smooth. 2n = 24, 36. About 80 sperm copious.
spp., Madagascar and Comores Is., tropical S
and SE Asia, Indonesia, Borneo, Philippines,
46 Araceae

8. Anthurium Schott 9. Spathiphyllum Schott

Anthurium Schott, Wiener Z. Kunst 1829 (3): 828 {1829); Spathiphyllum Schott in Schott & Endlicher, Melet. Bot.: 22
Madison, Selbyana 2: 239-282 {1978); Croat & Bunting, {1832); Bunting, Mem. N.Y. Bot. Gard. 10: 1-53 {1960), rev.;
Aroideana 2: 15-25 {1979); Mayo, Kew Bull. 36: 691-719 Nicolson, Blumea 16: 119-121 {1968) (rev. Old World spp.).
{1982); Croat & Sheffer, Aroideana 6: 85-123 {1983); Croat,
Ann. Mo. Bot. Gard. 70: 211-420 {1984) & Monogr. Mo. Bot. Peduncle subequal to or longer than leaf; spathe
Gard. 14: 1-204 {1986) & Ann. Mo. Bot. Gard. 78: 539-855.
cuspidate, usually spreading, rarely enclosing spa-
dix, persistent; spadix sessile or stipitate, erect,
Stem creeping to erect, sometimes root-climbing;
shorter than spathe; flowers (2- )3-merous; tepals
leaf blade entire and linear to cordate, hastate or
4-6, fornicate at apex or if connate, then forming a
sagittate, trilobed or trisect, pedately or radiately
truncate cup; stamens as many as tepals; ovary
lobed or -sect (primary lobes rarely themselves
(2- )3(4)-locular, ovules 2-8 per locule, placenta
pinnately lobed); inflorescence 1; peduncle rela-
axile, style comic or inconspicuous; berry rounded
tively long, more rarely short; spathe usually per-
or :::t::: attenuate apically, 1-8-seeded; embryo axile,
sistent, :::t::: rarely brightly coloured; spadix sessile
narrowly cylindric. 2n = 30, 60. Forty one spp.,
to long-stipitate, sometimes clavate, rarely glo-
tropical America, Melanesia, E Malesia, Pacific,
bose; mature berry globose to elongate-fusiform,
in damp and humid places; Bunting (1960) re-
variously and often brightly coloured, 2-4-seeded
cognised four sections; S. solomonense Nicolson is
(rarely more); seed usually :::t::: oblong, testa :::t:::
very distinct and recognised as Sect. Chlaeno-
verruculose, embryo axile, subcylindric. 2n = 30,
phyllum by Nicolson (1994).
60, 90 (20, 24, 28, 40, 48, 56, 84). About 800 spp.
(probably more according toT. Croat), Mexico to
Argentina, from humid to dry tropical lowland 10. Holochlamys Engler
to lower montane forest, normally epiphytic or
Holochlamys Engler in Beccari, Malesia 1: 265 {1883).
climbing hemiepiphytic, also terrestrial, litho-
phytic or rarely rheophytic; A. scherzerianum
Peduncle shorter than petiole; spathe tightly
Schott and A. andraeanum Linden important in
clasping the spadix, not spreading, drying after
horticulture; 19 sections recognised (Croat and
an thesis and long persistent, gradually decompos-
Sheffer 1983).
ing; spadix sessile; tepals 4, connate into a trun-
cate cup; stamens 4; ovary 1-locular, ovules many,
IV. Subfam. Monsteroideae Engler (1876). anatropous, placenta basal; berry few- to 1-
seeded. 2n = 60. Only one sp., H. beccarii (Engler)
Laticifers absent; trichosclereids (H- or T-shaped) Engler, Melanesia (New Britain, New Guinea), on
abundant (except Anadendreae, Heteropsideae; forest floor.
in Amydrium present in inflorescence and very
sparse elsewhere); terrestrial, climbers, hemi- 2. Tribe Anadendreae Bogner & French (1984).
epiphytes or more rarely epiphytes; petiole
geniculate apically, sheath usually long; leaf blade Trichosclereids absent; climbers; petiole geni-
never sagittate, sometimes pinnatifid or perfo- culate apically, sheathed almost to apex; finer
rated; spathe not constricted centrally, expanded venation reticulate; peduncle relatively long;
or boat-shaped; spadix fertile to apex; flowers spathe boat-shaped, longer than spadix; perigone
bisexual, perigone present or absent, 2-merous membranaceous, urceolate, shorter than gyno-
(2-3-merous in Spathiphylleae). ecium; stamens 4, thecae dehiscing by longi-
tudinal slit; gynoecium truncate apically, ovary
1. Tribe Spathiphylleae Engler (1887).
1-locular, ovule 1, hemianatropous, placenta
basal, stigma transversely oblong or subspheroid;
Tissues with small trichosclereids arranged in endosperm absent, embryo large.
bundles; stem short, upright; leaves (sub- )disti-
chous, petiole geniculate apically; leaf blade ob-
11. Anadendrum Schott
long, cuspidate-acuminate, primary leaf venation
pinnate, finer veins parallel to them; inflorescence Anadendrum Schott, Bonplandia 5: 45 (1857).
solitary; flowers bisexual, 2-3-merous, tepals free
or connate; thecae dehiscing by longitudinal slit; Leaf blade obliquely ovate-oblong, entire; spathe
ovules anatropous to hemianatropous; seed :::t::: oblong-ovate, white; spadix cylindric, stipitate;
oblong, endosperm copious. berry subglobose, truncate apically; seed sub-
 Araceae 47

globose, testa thickish. 2n = 60. About seven spp., cence usually 1; spathe conchiform to ovate, re-
SE Asia, Indonesia, Borneo, hemiepiphytes. flexed at anthesis; spadix sessile to long-stipitate;
ovary 1-locular, ovules 2, anatropous, placenta
near base of single, deeply intrusive septum;
3. Tribe Heteropsideae Engler (1905). berry white-green or orange; seed subglobose,
Trichosclereids absent; bushy root climbers or endosperm present. 2n = 60. Six spp., tropical
hemiepiphytes; main shoot monopodia!, flower- SE Asia and Malesia, climbing hemiepiphytes.
ing articles usually short, axillary; petiole usually
almost entirely adnate to succeeding internode,
14. Rhaphidophora Hasskarl
leaving only apical geniculum free; primary and
secondary leaf venation parallel-pinnate, veins Rhaphidophora Hasskarl, Flora 25, Beibl. 1: 11 (1842).
fused distally into submarginal collective vein, Raphidophora Schott (1858), orth. var.
finer venation ± reticulate; spathe convolute, Afrorhaphidophora Engler (1906).
opening at anthesis; perigone absent; thecae
dehiscing by apical slit; gynoecium truncate, Sometimes repent, often robust; leaf blade entire,
ovary incompletely 2-locular, ovules 2 per locule, irregularly perforate or pinnately lobed, finer
anatropous, axile at base of partial septum; venation subparallel-pinnate; inflorescences 1-3;
stigma oblong or rounded; endosperm absent. spadix sessile, lowermost flowers often female;
ovary partially 2-locular, septa deeply intrusive,
placentas basal and parietal on septa, ovules
12. Heteropsis Kunth many, anatropous; berry red, many-seeded;
seed oblong, testa thin, smooth, endosperm
Heteropsis Kunth, Enum. pl. 3: 59 (1841).
copious. 2n = 60, 120 (42, 54, 56). About 100
spp., tropical Africa and from S and SE Asia
Inflorescence 1, axillary or terminal on short pe-
to Melanesia, Australia, and the Pacific, climbing
duncle; spathe ovate or ovate-oblong; spadix stipi-
hemiepiphytes, rarely rheophytic (R. beccarii
tate, shorter than spathe, cylindric or ellipsoid;
Engler).
berry shortly obovoid or obpyramidal, 1-4-
seeded; seed obovoid. 2n = 28. About 13 spp.,
Costa Rica to S Brazil. 15. Epipremnum Schott
Epipremnum Schott, Bonplandia 5: 45 (1857).
4. Tribe Monstereae Engler (1876).
Leaf blade entire, perforate or pinnately lobed;
Climbing hemiepiphytes (except Stenosperma-
finer venation subparallel-pinnate; ovary 1-
tion); leaves distichous; petiole geniculate
locular; ovules 2, more rarely 4-8, anatropous,
apically, sheath long (except Amydrium), almost
equalling petiole, leaf blade normally oblique; pe- placenta parietal or near base of parietal partial
septa; berry 1-8-seeded; seed kidney-shaped, testa
duncle relatively short (except Stenospermation),
thickish, endosperm copious. 2n = 60 (56, 84).
spathe usually erect and boat-shaped (except
Seventeen spp., tropical S and SE Asia to Australia
Amydrium); spadix equalling or shorter than
and the Pacific, climbing hemiepiphytes.
spathe; perigone absent, thecae dehiscing by
longitudinal slit not reaching base.
16. Scindapsus Schott
13. Amydrium Schott Scindapsus Schott in Schott & Endlicher, Melet. Bot. 21 (1832).

Amydrium Schott, Ann. Mus. Bot. Lugd.-Batav. 1: 127 (1863);

Nicolson, Blumea 16: 123-127 (1968); Carvell, Am. J. Bot. 6
Sometimes robust; petiole sheath rather broad, in
(Suppl.): 231 (1989). some spp., decomposing to form a highly irritant,
Epipremnopsis Engler (1908). fibrous mass; leaf blade entire, fine venation
parallel-pinnate; spadix sessile; ovary 1-locular,
Trichosclereids sparsely present (most abundant ovule 1, anatropous, placenta basal; berry rather
in style according to Carvell 1989); stem some- large; seed rounded, subreniform, compressed,
times creeping; petiole sheath short; leaf blade testa thickish, endosperm present. 2n = 60 (42, 56,
entire or pinnately lobed, usually perforate; pri- 58, 64, 70, 112). Thirty spp., from tropical Sand SE
mary lateral veins forming obscure, submarginal Asia through Melanesia into the Pacific, hemi-
collective veins, finer venation reticulate; inflores- epiphytic climbers or rupicolous.
48 Araceae

17. Monstera Adanson Fig. 12A-C

Monstera Adanson, Fam. 2: 470 (1763), nom. cons.; Madison,
Contrib. Gray Herb. 207: 1-100 (1977), rev.

Leaf blade entire or perforate and/or pinnatifid,

fine venation parallel-pinnate or ± reticulated;
inflorescences !-several; spadix sessile; ovary 2-
locular, ovules 2 per locule, anatropous, placenta
at base of septum; seed obovoid or subcordate, a
little compressed, raphe S-shaped, testa smooth,
thickish, endosperm absent, embryo large. 2n
= 60 (24, 48, 56, 58, 70). About 40 spp., trop-
ical America (Mexico to S Brazil), climbers,
hemiepiphytes in humid forest; the variegated,
juvenile leaves of M. dubia (H.B.K.) Engler &
Krause, adpressed to tree boles, frequently attract
the attention of travellers in Amazonian forests;
the pulp surrounding the seeds of M. deliciosa
Liebm. is delicious and made into ice cream
and soft drinks in Mexico; M. deliciosa is one of
the world's most ubiquitous ornamental foliage
plants; four sections are recognised.

18. Alloschemone Schott

Alloschemone Schott, Gen. Aroid.: 99 (1858); Madison,
Selbyana 1: 325-327 (1976).

Leaves distant; petiole ± terete; leaf blade pin-

nately lobed, subcordate, lobes acute, falcate, 4-6
per side, each with only 1 primary lateral vein,
finer venation parallel-pinnate; spadix sessile;
stamens free to connate; ovary 1-locular, ovule
1, amphitropous, placenta basal; fruit and seed
unknown. 2n = 84. Two spp., Amazonian Brazil,
climbers or hemiepiphytes in forest.

19. Rhodospatha Poeppig

Rhodospatha Poeppig in Poeppig & Endlicher, Nova Gen. Sp.
3: 91 (1845).
Anepsias Schott (1858). Fig. 12A-I. Araceae-Monsteroideae. A-C Monstera obliqua.
A Habit. B Spadix. C Stamen. D-1 Stenospermation spru-
Leaf blade entire, fine venation parallel-pinnate; ceanum. D Spadix. E Flowers seen from above. F Flower,
longitudinal section, showing two stamens, basal placenta
spathe sometimes rose-coloured within; spadix ±
with anatropous ovules, and broad stylar region with small
long-stipitate, flowers small; pollen extruded in stigma. G Ovule, note papillose lower part of funicle. H
strands; ovary 2( -3-6)-locular, ovules usually Stamen, front view. I Stamen, side view. (Takhtajan 1982)
numerous per locule, amphitropous, placentas
axile; berry small, 10-12-seeded; seeds ::±:: flat-
tened, rounded-reniform, testa hard, verruculose,
endosperm present, embryo rather large. 2n = 28, 20. Stenospermation Schott Fig. 12D-I
56. About 75 spp., tropical America (Mexico to Stenospermation Schott, Gen. Aroid.: t 70 (1858); Gomez, Rev.
SE Brazil), climbers, hemiepiphytes in humid Stenospermation (Araceae) in Central America, Masters
forest. Thesis, St. Louis University, 114 pp. (1983).
 Araceae 49

Shoots usually short; leaf blade entire; fine vena- Rhizome hypogeal, erect; leaves several; petiole
tion parallel-pinnate; inflorescence 1, nodding or geniculate apically, sheath long; leaf blade stron-
erect; peduncle relatively long; spadix stipitate; gly sagittate, often perforated; inflorescence 1;
ovary 2-locular, ovules 4 to many per locule, ana- peduncle long; spathe convolute, basally forming
tropous, placenta basal; berry obovoid, white, distinct tube, blade strongly fornicate; spadix
rarely orange, several- to many-seeded; seed sessile, much shorter than spathe; tepals 4; sta-
clavate-cylindric, slender, testa thickish, smooth, mens 4; ovary 2-locular, locules 1-ovulate, ovules
endosperm copious. 2n = 28. About 36 spp., tro- anatropous, placenta axile, style conical; berry
pical America (Guatemala to SE Brazil), in humid obovoid, purplish red; seed campylotropous, testa
forest, epiphytic or on the ground. thick, crested and warty. 2n = 26. Only one sp., D.
desciscens (Schott) Engler, E Brazil, in humid
forest and wet, shaded places.
V. Subfam. Lasioideae Engler (1876).

Laticifers absent; terrestrial or rooted aquatics; 23. Anaphyllopsis A. Hay

stem tuberous or rhizomatous, usually geophytic Anaphyllopsis A. Hay, Aroideana 11: 25-31 (1989, "1988").
(except Lasia, Podolasia); petiole usually aculeate
or warty or with striking coloration, long, usually Rhizome hypogeal, erect; leaf 1(-2); petiole
± geniculate apically; primary lateral veins of ma- geniculate apically; leaf blade broadly hastate-
jor leaf blade divisions pinnate to arcuate-parallel, sagittate in outline, pinnately lobed or pinnate,
higher-order venation reticulate; spadix flowering often irregularly perforated, ultimate lobes often
and fruiting in basipetal sequence; flowers bi- bifidtruncate; inflorescence 1(-2); peduncle long;
sexual, perigoniate (except Pycnospatha); tepals spathe marcescent, convolute basally, spirally
fornicate, ± truncate, free, in 2 or more whorls; twisted apically; spadix stipitate, much shorter
stamens free (except Lasimorpha), anther termi- than spathe; tepals 4; stamens 4; ovary 1-locular,
nal; embryo large, endosperm present (except ovules 1-2, anatropous, placenta basal; seed
Anaphyllum), forming a thin but distinct layer. campylotropous. 2n = 26. Three spp., northern S
America, in humid forest and wet places.
21. Dracontium L.
Dracontium L., Sp. Pl.: 967 (1753). 24. Pycnospatha Thorel ex Gagnep.
Echidnium Schott (1857).
Pycnospatha Thorel ex Gagnep., Bull. Soc. Bot. Fr. 88: 511
Sometimes gigantic, tuber depressed-subglobose; (1941); Bogner, Oesterr. Bot. Z. 122: 202 (1973).
leaf usually 1; petiole sometimes geniculate api-
cally; leaf blade dracontioid, ultimate lobes with Tuber depressed-subglobose; leaves 1-2; petiole
acuminate tips, leaf dissection attained at least smooth, rough or aculeate, strikingly patterned,
partly by perforation; inflorescence 1; spathe ± not geniculate apically; leaf blade dracontioid, tri-
boat-shaped, persistent; spadix stipitate, much sect; inflorescence 1, appearing before or with leaf;
shorter than spathe; tepals 4-8; stamens 4-6(-12); peduncle short; spathe thick, ± convolute below
pollen extruded in masses; ovary incompletely 1- but margins not overlapping; spadix stipitate,
6-locular, locules 1(2)-ovulate, ovules anatropous much shorter than spathe; flowers without
to subcampylotropous, placenta axile to subbasal, perigone; stamens ca. 6 or more, crowded densely
style long, attenuate; berry obpyramidal to obo- together; ovary 1-locular, ovule 1, anatropous to
void, red to purplish red; seed campylotropous, hemianatropous, placenta basal to subparietal;
testa thick, crested and warty. 2n = 26. About 23 berry spiny, globose, dark green; seed ellipsoid,
spp., tropical America (Mexico to Paraguay and testa thick, verrucose, embryo large, endosperm
Bolivia), in forest and savanna, ground-dwelling, sparse. 2n = 26. Two spp., Laos, Thailand,
seasonally dormant or evergreen. Vietnam on forest floor, seasonally dormant.

22. Dracontioides Engler 25. Anaphyllum Schott

Dracontioides Engler, Pfianzenreich IV.23C: 36 (1911); Mayo, Anaphyllum Schott, Bonplandia 5: 126 (1857); Sivadasan,
Aroideana 1: 4-10 (1978). Taxonomic study of Araceae of S India, unpubl. doctoral
Dracontium sect. Urospathopsis Engler (1884). thesis, University of Calicut, Kerala (1982).
50 Araceae

Clump- or colony-forming, rhizome creeping; dosperm present, sparse. 2n = 26. Only one sp., L.
leaves few; petiole geniculate apically; leaf blade senegalensis Schott, Senegal to Angola, helophytic.
pinnately lobed or pinnate; inflorescence 1; pe-
duncle long; spathe marcescent, either convolute
basally, spirally twisted and long-acuminate 28. Podolasia N.E. Brown
above, or oblong-ovate and fully expanded; spadix Podolasia N.E. Brown, Gard. Chron. II, 18: 70 (1882); Hay,
stipitate, rather short; tepals 4; stamens 4; ovary Blumea 33: 463-465 (1988).
1-locular, ovule 1, anatropous, placenta parietal,
intrusive; berry ovoid, red; seed ovoid, testa Rhizome short, erect to decumbent, petiole ac-
unlignified, membranous, smooth, embryo large, uleate, spines usually pointing downwards; leaf
endosperm absent. 2n = 26. Two spp., southern blade deeply sagittate to hastate; inflorescence 1;
India, in evergreen forest, swamp forest peduncle long; spathe linear-lanceolate, fully ex-
undergrowth. panded; spadix relatively long, usually stipitate;
tepals 4-6; stamens 4-6; ovary 1-locular, ovule 1,
26. Cyrtosperma Griffith anatropous, placenta parietal to basal; berry
obovoid, red; seed campylotropous, ± spherical,
Cyrtosperma Griffith, Notul. Pl. Asiat. (Posthum. Pap.) 3: 149 testa thin, hard, smooth. 2n = 26. Only one sp., P.
(1851) & Icon. 3, t.169 (1851); Thompson, J. Agric. Tradit. stipitata N.E. Brown, SE Asia.
Bot. -Appl. 29: 185-203 (1982); Hay, Blumea 33: 427-457
(1988).
29. Lasia Loureiro
Sometimes gigantic, rhizome thick, condensed,
creeping; leaves several; petiole aculeate, geni- Lasia Loureiro, Fl. Cochinch. 64, 81 (1790); Hay, Blumea 33:
culate apically; leaf blade sagittate to hastate; 458-463 ( 1988).
inflorescence 1(-2); peduncle long; spathe erect,
upper part sometimes long-acuminate and spiral- Clump-forming; stem and petiole aculeate; leaf
ling, rarely fornicate; spadix usually much shorter blade pinnately to pedately lobed, rarely hastate-
than spathe; tepals 4-6; stamens 4-6; ovary 1- sagittate; inflorescence 1; peduncle long; spathe
locular, ovules 1-many, ovule campylotropous erect, linear-lanceolate, caducous, lowest part
to subamphitropous, placenta basal to parietal, gaping at anthesis, upper part spirally twisted,
style short or inconspicuous; berry obovoid, 1-7- thick, spongy; spadix short, sessile; tepals 4-6, sta-
seeded, usually red; seed reniform, orbicular or mens 4-6; ovary 1-locular, ovule 1, anatropous,
helically twisted, crested, warty or smooth. 2n = placenta apical, style inconspicuous; berry green,
26. About 12 spp., tropical SE Asia to Melanesia apex often minutely spiny; seed ± pyramidal,
and the Pacific, helophytic, alongside streams, campylotropous, testa thin, hard, sparsely spiny.
pond margins and other wet places; the rhizomes 2n = 26. Two spp., S and SE Asia, Indonesia,
of C. merkusii (Hassk.) Schott are an important Borneo, New Guinea, aquatic or swampy sites.
subsistence food in the Pacific (Thompson 1982);
four informal groups recognised by Hay (1988). 30. Urospatha Schott
Urospatha Schott, Aroideae 3 (1853).
27. Lasimorpha Schott Urospathella Bunting (1988).
Lasimorpha Schott, Bonplandia 5: 127 (1857); Hay, Blumea 33:
465-468 (1988). Rhizome hypogeal, horizontal to vertical; leaves
several; petiole smooth to scabrid-verrucose; leaf
Sometimes gigantic, forming large colonies, rhi- blade sagittate to hastate, rarely narrowly lan-
zome short, thick, hypogeal, stoloniferous; leaves ceolate; inflorescence 1; peduncle long; spathe
several; petiole 4-6-angled, aculeate along angles, convolute below, above long acuminate, usually
weakly geniculate apically, sheath short; inflores- spirally twisted; spadix sessile to shortly stipitate,
cence 1; peduncle long; spathe ovate, convolute short; tepals 4-6; stamens 4-6; ovary incompletely
basally, persistent; spadix sessile to shortly stipi- 1-2-locular, ovules (1-)2 or more per locule,
tate, obtuse; tepals 4-6; stamens 4-6, filaments anatropous, placenta axile or basal-parietal; seed
free or ± connate; ovary 1-locular, ovules 4-6, cat1lpylotropous, testa hard, thickish, smooth to
campylotropous, placenta basal and intrusive- verrucose. 2n = 52. About ten spp., Guatemala to
parietal; berry ± globose, red, 1-4-seeded; seed SE Brazil, open aquatic habitats, swamps, along
campylotropous, testa warty and spiny, en- rivers, in brackish water.
 Araceae 51

VI. Subfam. Calloideae Endl. (1837). to basal; berry and seed large, testa smooth, en-
dosperm absent.
Laticifers present, simple, articulated; rooted
aquatic; stem rhizomatous; leaves distichous;
32. Zamioculcas Schott Fig. 13
petiole sheath long; leaf blade cordate; venation
parallel-pinnate, uniformly fine; spathe fully ex- Zamioculcas Schott, Syn. Aroid.: 71 (1856); Mayo, Fl. Trop.
panded, elliptic- to ovate-lanceolate, persistent; East Africa, Araceae: 15-16 (1985).
flowers bisexual, 3-merous, perigone absent; sta-
mens 6 (or more); ovary 1-locular, ovules 6-9, Rhizome short, very thick; leaves few to several,
anatropous, placenta basal; endosperm copious. petiole thickened basally, geniculate distally; leaf
blade pinnately compound, leaflets oblong-
elliptic, thick, deciduous, capable of rooting to
31. Calla L. form new plants; inflorescences 1-2; peduncle
Calla L., Sp. Pl.: 968 (1753); Riedl in Hegi, Ill. Fl. Mitteleuropa, short; male flowers: filaments free, thecae introrse;
ed. 3, 2: 326-327 (1979). sterile flowers: with 4 tepals surrounding clavate
pistillode; female flowers: staminodes absent;
Stem green; petiole sheath with long, free apical berry 1-2-seeded, white; seed ellipsoid. 2n = 34.
ligule; inflorescence usually 1; spathe white within, Only one sp., Z. zamiifolia (Loddiges) Engler,
green without; spadix stipitate, apical flowers tropical and southern subtropical Africa (Kenya
often male; berry several-seeded, red; seed terete- to Natal); in moist forest and savannas, seasonally
oblong, testa thick. 2n = 36, 54, 72. Only one dormant or evergreen.
sp., C. palustris L., circumboreal, aquatic and
helophytic in streams, ponds, swamps and bogs. 33. Gonatopus J.D. Hooker ex Engler
Gonatopus J.D. Hooker ex Engler in A. & C. De Candolle,
VII. Subfam. Aroideae Engler (1876). Monogr. Phan. 2: 208 (1879); Mayo, Fl. Trop. East Africa,
Araceae: 9-15 (1985).
Heterolobium Peter (1930).
Laticifers present, rarely absent, usually simple, Microculcas Peter (1930).
more rarely anastomosing; stem most frequently
hypogeal, tuberous or rhizomatous, less often Rhizome or subglobose tuber; leaf 1; petiole
aerial, rarely hemiepiphytes, climbers or epi- geniculate basally or centrally; leaf blade pinnately
phytes, very rarely floating aquatics (Pistieae); compound, primary lateral pinnae pinnately to
petiole usually not geniculate apically (except tripinnately lobed; inflorescences 1-4, often
Anubiadeae, Bognera, Culcasieae, Zami.ocul- appearing before leaf; peduncle short to long;
cadeae, rarely in Philodendreae and Homalo- stamen filaments connate; female flowers:
meneae); spathe usually differentiated into lower, staminodes absent, rarely 1; berry ovoid-ellipsoid,
convolute tube and upper, gaping blade; flowers orange, red or :±: dirty white, large. 2n = 34, 68.
unisexual, perigone absent (except Stylochaet- Five spp., tropical and southern subtropical Africa
oneae, Zamioculcadeae). (Kenya to Natal), on forest floor, seasonally
dormant.
1. Tribe Zamioculcadeae Engler (1876).
2. Tribe Stylochaetoneae Schott (1856).
Laticifers absent; seasonally dormant, stem hypo-
geal; leaf blade compound; petiole geniculate; Laticifers absent; stem hypogeal, rhizomatous,
spathe convolute basally, blade reflexed at anthe- roots often very thick, fleshy; leaf primary lateral
sis; spadix subequalling spathe, female zone sepa- veins mostly arising basally, arcuate, finer vena-
rated from male by short, constricted zone of tion reticulate; spathe tube margins connate; spa-
sterile flowers, male zone cylindric to clavate, dix mostly hidden within spathe, fertile to apex;
fertile to apex; flowers unisexual with perigone of flowers with :±: urceolate, connate perigone; male
4 free, :±: prismatic tepals; male flowers: stamens 4, flower usually with central peglike pistillode, sta-
surrounding :±: clavate pistillode, filaments dis- men filaments filiform, anthers terminal, dehisc-
tinct (connate in Gonatopus), anther terminal, ing by longitudinal slit, connective slender; ovary
connective inconspicuous, pollen extruded in 1-4-locular, locules 1- to many-ovulate, ovules
strands; female flowers: ovary 2-locular, locules anatropous, placenta basal, parietal or axile; en-
1-ovulate, ovules hemianatropous, placenta axile dosperm copious.
52 Araceae

level; spathe tube ±ventricose at base, rarely con-

stricted between lower and upper swollen zones
or entirely narrow-cylindric; female flowers borne
spirally or in 1 whorl, sometimes contiguous with
male zone, often with ± naked zone or sterile
flowers in between; male flower 2-7 -androus;
perigone often with fleshy margins; infructescence
borne at or below ground level, berry fleshy, 1-
few-seeded. 2n = 28, 56. Seventeen spp., tropical
and southern subtropical Africa, in humid to dry
forest, woodland, savannas, seasonally dormant
or evergreen geophytes.

3. Tribe Dieffenbachieae Engler (1876).

Internodes distinct; spadix: female zone entirely
adnate to spathe, laxly flowered, sterile male flow-
ers (synandrodes) usually present between female
and male zones; male flowers: stamens connate
into a truncate, prismatic synandrium, thecae lat-
eral, fused connectives strongly thickened; female
flowers: distant, ovary 1-3-locular, locules 1-
ovulate, ovule anatropous, placenta basal or
basal-axile.

35. Dieffenbachia Schott

Dieffenbachia Schott, Wiener Z. Kunst 1829 (3): 803 (1829);
Young, Am. J. Bot. 73: 931-944 (1986).
Maguirea A.D. Hawkes (1948).

Sometimes robust; stem erect or decumbent; leaves

several per article; petiole not geniculate, sheath
relatively long; leaf blade ovate to lanceolate, often
variegated; primary lateral veins pinnate, forming
a marginal vein, finer venation parallel-pinnate;
inflorescences (1-)2-7; peduncle usually shorter
than spathe; spathe slightly constricted, tube usu-
ally much longer than blade, persistent; spadix:
female zone usually separated from male zone by
± naked axis bearing a few scattered sterile
Fig. 13A-G. Araceae-Aroideae. Zamioculcas zamiifolia. A flowers; male flowers: 4-5-androus; female flowers:
Habit; note geniculate leaves. B Inflorescence. C Staminate gynoecium surrounded by 4-5 clavate staminodes;
flower. D Sterile flower. E Stamen. F Pistillate flower, ovary ovary subglobose to ovoid, 1-3-locular, placenta
wall removed to show locules. G Rooted leaflet with young
axile-basal or basal; berry 1-3-seeded, globose,
tuberous rhizome and shoot. (Takhtajan 1982)
often scarlet; seeds globose to ovoid, testa smooth,
thickish, embryo large, endosperm absent. 2n =
34. Stylochaeton Leprieur 34, 68. About 30 spp., Mexico to northern Argen-
tina, evergreen, terrestrial or helophytic, in humid
Stylochaeton Leprieur, Ann. Sci. Nat. Bot. II, 2: 184 (1834); forests and damp sites.
Mayo, Fl. Trop. East Africa, Araceae: 42-54 (1985); Ntepe-
Nyame, Fl. Cameroun 31: 42-46 (1988).
36. Bognera Mayo & Nicolson
Leaves 1-few; leaf blade lanceolate to cordate, Bognera Mayo & Nicolson, Taxon 33: 690 (1984); Madison
hastate or sagittate; inflorescences l-4, often sub- in Aroideana 3: 101-102 (1980) (as Ulearum recondita
tended by cataphylls, borne at or below ground Madison).
 Araceae 53

Stem creeping, epigeal, rhizomatous; foliage leaf Often robust; leaf 1; leaf blade bipinnatifid to
1 per article, alternating with cataphylls; petiole tripinnatifid, ± dracontioid, rarely pinnatifid; in-
indistinctly geniculate at base and apex, shortly- florescence 1(-2), subtended by cataphylls; pe-
sheathed; leaf blade ovate-lanceolate; venation duncle shorter than petiole; spathe boat-shaped,
pinnate, secondary veins arcuate, ± parallel to tube convolute, blade gaping, marcescent and
primaries, finer veins distinctly reticulated; inflo- later deciduous; spadix sometimes much longer
rescence 1; peduncle shorter than petiole; spathe than spathe; female zone free, usually contiguous
boat-shaped, unconstricted; spadix: female zone with male, rarely with a few bisexual flowers in
separated from male zone by ± naked zone bear- between; thecae oblong to broadly ellipsoid; ovary
ing a few globose sterile flowers; male flowers: 3- 3-6(?-7)-locular, ovules anatropous, placenta
4-androus; female flowers: staminodes absent, axile; berry depressed-globose, slightly furrowed,
ovary 1-locular, placenta basal; berry and seed scarlet, 3-5-seeded; seed ellipsoid, testa smooth or
unknown. 2n = 34. Only one sp., B. recondita granulate. 2n = 34. Five spp., tropical and sub-
(Madison) Mayo & Nicolson, Brazil (western tropical S America.
Amazonia), on forest floor.

39. Asterostigma F.E.L. Fischer

4. Tribe Spathicarpeae Schott (1856) & C.A. Meyer Fig. 14
Seasonally dormant, stem tuberous, rarely
Asterostigma F.E.L. Fischer & C.A. Meyer, Bull. Cl. Phys.-
rhizomatous (Gearum), hypogeal; primary lat- Math. Acad. Imp. Sci. Saint Petersbourg II, 3: 148 (1845);
eral veins of leaf or leaf lobes forming single Bogner, Adansonia II, 9: 125-130 (1969).
marginal vein (except Spathantheum and Spa- Andromycia A. Rich. (1850).
thicarpa), finer venation reticulate; spathe usually
unconstricted; spadix fertile to apex (except Leaf usually 1; leaf blade pinnatisect or rarely
Mangonia); male flower a synandrium of ± con- entire; inflorescences 1-3, appearing before or
nate stamens (some partly or completely free in with leaf; peduncle relatively long; spathe ±
Gorgonidium); gynoecium surrounded by free unconstricted, persistent, tube narrow, blade
staminodes or urceolate synandrode; ovary gaping at anthesis; spadix: female zone free or
locules !-ovulate (except Mangonia); embryo partly adnate to spathe, laxly flowered, contig-
usually relatively small, axile, endosperm copious, uous with male zone; synandrium mushroom-
rarely absent (Gearum). shaped, umbonate; gynoecial staminodes 3-5,
free or connate, ovary 3-5-locular, ovules anatro-
pous, placenta axile-basal; berry ± depressed-
37. Mangonia Schott
globose; testa thin, smooth. 2n 34.
Mangonia Schott, Oesterr. Bot. Wochenbl. 7: 77 (1857); About seven spp., southern tropical S America,
Bogner, Darwiniana 18: 70-79 (1973). terrestrial.
Felipponia Hicken (1917), non Felipponea Brotherus (1912).
Felipponiella Hicken (1928).
40. Gorgonidium Schott
Small; leaves several; leaf blade narrowly elliptic
to oblong-sagittate; inflorescence 1, appearing Gorgonidium Schott, Ann. Mus. Bot. Lugd.-Bat. 1: 282 (1864);
before leaves, subtended by cataphylls; spathe: Bogner & Nicolson, Bot. Jahrb. 109: 529-554 (1988), rev.
tube subcylindric, blade lanceolate to oblong,
erect, gaping; spadix: female zone short, contigu- Leaf 1; leaf blade pinnatifid, pinnatisect or
ous with male zone or separated by short ± naked bipinnatifid; inflorescence 1(-2), appearing before
zone, male zone more laxly flowered, terminal leaf; peduncle usually shorter than spathe;
sterile appendix of sterile male flowers; spathe unconstricted, boat-shaped; spadix: female
synandrium 2-5-androus; ovary 2-3-locular, zone continguous with male; male flowers: 3-
ovules 2 per locule, anatropous, placenta axile; 7-androus, stamens free or ± connate; central
testa smooth. 2n = ? Two spp., S Brazil, Uruguay, pistillode often present and composed of 3-4
in seasonally wet places. stylodia and stigmatoids (G. vargasii Bogner &
Nicolson); gynoecial staminodes 6-8, ovary
(2-)4-5( -7)-locular, ovules atropous, placenta
38. Taccarum Brongn. ex Schott axile-basal; berry globose to depressed-globose;
Taccarum Brongn. ex Schott, Oesterr. Bot. Wochenbl. 7: 221 testa rough. 2n = 34. Three spp., N Argentina,
(1857); Bogner, Willdenowia 19: 191-198 (1989). Bolivia, Peru, 900-3000 m alt.
54 Araceae

ing before or with leaves; peduncle short; spathe

ovate-lanceolate, unconstricted; spadix shorter
than spathe, female zone partly adnate to spathe,
laxly flowered, separated from male by few bi-
sexual or sterile flowers; synandrium long-
stipitate, elongate-conoid, apical whorl of anthers
sometimes shortly overtopped by central
pistillode; gynoecial staminodes 3-5, ovary 3-5-
locular, ovules atropous, placenta axile-basal;
berry 3-5-seeded; seed rather large, testa thick,
succose. 2n = 34. Only one sp., S. vermitoxicus
(Griseb.) Engler, northern Argentina, Bolivia,
Paraguay, in dry thorn forest.

42. Gearum N.E. Brown

Gearum N.E. Brown, J. Bot. 20: 196 (1882); Bogner & Nicolson,
Bot. Jahrb. 109: 531 (1988); Mayo eta!., Kew Bull. 49: 785-
788 (1994).

Rhizome vertical; leaves 1-2, appearing soon

after inflorescence; petiole sheath about half the
petiole length; leaf blade pedatisect, coriaceous,
venation reticulate; inflorescence 1; peduncle
short; spathe constricted; spadix shorter than
spathe; female zone of spadix separated from
~ male by sterile zone bearing shallow, truncate
~~ c synandrodes; ovary depressed-globose, 3-4-

©
locular, ovules atropous, placenta axile-basal,
' stigma subsessile, weakly 3-4-lobed; berry
l
depressed-globose; seed broadly ellipsoid; testa
D
thin; embryo large, endosperm absent. 2n = 34
and ca. 68. Only one sp., G. brasiliense N.E. Brown,
C Brazil.

43. Spathantheum Schott

Spathantheum Schott, Bonplandia 7: 164 (1859); Crisci, Rev.
Mus. La Plata, Bot. 11:266-268 (1971).

Leaf 1, petiole sheath very short; leaf blade entire

to pinnately lobed, emarginate to subcordate at
Fig. 14A-F. Araceae-Aroideae. Asterostigma luschnathia- base, primary lateral veins forming submarginal
num. A Habit. B Spadix. C, D Synandrium seen from the side
and from above. E Pistil. F Same, longitudinal section.
collective vein; inflorescence 1-2; spathe oblong-
(Takhtajan 1982) elliptic, unconstricted; spadix entirely adnate to
spathe or male zone free (S. intermedium Bogner),
female zone either contiguous with male or with
an intermediate zone of mixed male and female
41. Synandrospadix Engler
flowers; stamens completely connate, anthers lat-
Synandrospadix Engler, Bot. Jahrb. Syst. 4: 61 (1883); Crisci, eral; gynoecial staminodes 5-8, ovary 5-8-locular,
Rev. Mus. La Plata, Bot. 11:255-260 (1971), Croat & Mount, ovules atropous, placenta axile-basal; berry 5-8-
Fl. Paraguay, Araceae: 34-37 (1988). seeded; testa smooth to slightly roughened.
Lilloa Spegazzini (1897).
2n = 34. Two spp., northern Argentina, Bolivia,
Peru.
Leaves several; petiole sheath long; leaf blade en-
tire, cordate-sagittate; inflorescences 1-2, appear-
 Araceae 55

44. Spathicarpa W.J. Hooker merous (subgen. Pteromischum) or solitary per

article; petiole rarely geniculate; leaf blade linear,
Spathicarpa W.J. Hooker, Bot. Misc. 2: 146 (1831); Crisci, Rev.
Mus. La Plata, Bot. 11: 260-266 (1971); Uhlarz, Beitr. Bioi. ovate, elliptic, oblong, cordate, sagittate, hastate,
Pflanz. 57: 389-429 (1982); Croat & Mount, Fl. Paraguay, trifid to almost trisect, or pinnatifid, bipinnatifid
Araceae: 29-34 (1988). or rarely pedatisect; inflorescences 1-11, secreting
Aropsis Rojas Acosta (1918). resin at anthesis from spathe and/or spadix; pe-
duncle usually short; spathe usually constricted
Small; tuber horizontal; leaves several; petiole between convolute tube and upper ± boat-
sheath rather long; leaf blade lanceolate to hastate, shaped, gaping blade; spadix sessile to stipitate,
sagittate; primary lateral veins mostly arising female and male zones usually separated by
basally, forming 1-2 submarginal collective veins; sterile zone of staminodial flowers, apical sterile
inflorescence 1, appearing with leaves; peduncle staminodial zone also rarely present; male flower
long; spathe unconstricted, persistent; spadix en- 2-6-androus; ovary (2-)4-8( -47)-locular, ovules
tirely adnate to spathe, often recurved at an thesis, 1-50 per locule, hemianatropous to hemi-
laxly flowered, female flowers forming 2 outer lon- atropous, placenta axile to basal; berry white, or-
gitudinal rows, enclosing 2 parallel rows of male ange or red; seed tiny to fairly large, ovoid-oblong
flowers; synandrium stipitate, peltate; gynoecial to ellipsoid, testa often costate; endosperm starch-
staminodes 3, situated only on one side of gyno- free. 2n = 28, 30, 32, 34, 36, 48 (26). Over 500 spp.,
ecium, ovary 1-locular, ovule atropous, placenta tropical and subtropical America (Mexico to
basal; berry ovoid; testa succose. 2n = 34. About northern Argentina); tropical humid forests,
five spp., NE Brazil to northern Argentina, in savannas, swamps, usually root-climbers, hemi-
semiarid and humid forests. epiphytes or epiphytes, less often lithophytic,
terrestrial or helophytic; important in horticulture
5. Tribe Philodendreae Schott (1856) as foliage ornamental plants, e.g. P. bipin-
natifidum Schott ex Endl., P. scandens C. Koch &
Laticifers present, simple; resin canals present in H. Sello; three subgenera and numerous sections
roots, stems, leaves and inflorescences; sclerotic recognised (Mayo 1990, Croat 1997).
hypodermis present in roots; terrestrial, climbing
or epiphytic, stem usually epigeal, intravaginal
6. Tribe Homalomeneae (Schott) M. Hotta
squamules present; petiole only rarely geniculate
apically; primary lateral veins pinnate (rarely (1970).
pedate) forming 1 marginal vein, finer venation Laticifers present, simple; resin canals present in
parallel-pinnate; spathe closing after anthesis and roots, stems and leaves; sclerotic hypodermis
persisting until fruit maturity, then deciduous at present in roots; terrestrial or rheophytic; stem
base; flowers unisexual, perigone absent; anther usually epigeal; petiole geniculum absent (very
thecae lacking cell wall thickenings in endo- rarely present in Homalomena); primary lateral
thecium (except P. goeldii, P. leal-costae); en- veins pinnate forming 1 marginal vein, higher-
dosperm copious. order venation parallel-pinnate; spathe often
boat-shaped, constricted or unconstricted, closing
after anthesis and persistent; flowers unisexual,
45. Philodendron Schott
perigone absent; anther thecae with cell-wall
Philodendron Schott, Wiener Z. Kunst 1829 (3): 780 (1829), thickenings in endothecium; endosperm copious.
nom. et orth. cons. ("Philodendrum"); Bunting, Acta Bot.
Venezuel. 10: 289-318 (1977, "1975") & Rev. Fac. Agron.
(Maracay) 10: 209-253 (1980); Mayo & Barroso, Aroideana 46. Furtadoa M. Hotta
2: 82-94 (1979); Mayo, Systematics of Philodendron Schott
(Araceae) with special reference to inflorescence characters, Furtadoa M. Hotta, Acta Phytotaxon. Geobot. 32: 142 (1981) &
Ph.D. thesis, Univ. Reading (1986) & Acta Bot. Brasilica 1 Gard. Bull. Singapore 38: 43-54 (1985).
(2) (Sup!.): 27-40 (1989, "1988") & Bot. J. Linn. Soc. 100:
139-172 (1989) & Kew Bull. 45: 37-71 (1990) & Kew Bull. Small, stem repent; leaf blade elliptic; inflores-
46: 601-681 (1991), rev. subg. Mecanostigma; Grayum,
Systematic Botany Monographs val. 43 (1995); Croat, Ann.
cences 1-3; peduncle relatively long; spathe boat-
Mo. Bot. Gar d. 84: 311-704 ( 1997). shaped, not constricted; spadix: female and male
Thaumatophyllum Schott (1859). zones with a zone of few sterile flowers in between;
male flowers: stamen 1, overtopped by flask-
Sometimes gigantic; internodes usually long, shaped pistillode, thecae dehiscing by longitudi-
intravaginal squamules present, foliage leaves nu- nal slit; female flowers: gynoecium with 1 truncate
56 Araceae

staminode, ovary 1-locular, ovules many, Rhizome thick, creeping; petiole relatively long,
hemianatropous, placenta basal; berry light green; rarely shortly and sparsely spiny; inflorescences
seed ellipsoid, testa smooth, thin. 2n = 40. Two 1-3; peduncle fairly long; spathe closing after an-
spp., Sumatra and Malay Peninsula, rheophytes thesis; spadix: female and male zones usually con-
on rocks in streams. The structure of the male tiguous, rarely with few sterile flowers in between;
flower is unique. male flowers: 3-8-androus; berry green to pale
green. 2n = 48, 72. Eight spp. Senegal to Angola,
helophytes, sometimes completely submerged, or
47. Homalomena Schott in forest swamps, rarely rheophytic.
Homalomena Schott in Schott & Endlicher, Melet. Bot.: 20
(1832); Furtado, Gard. Bull. Singapore 10: 183-238 (1939);
Hotta, Acta Phytotaxon. Geobot. 33: 127-139 (1982) & Gard. 8. Tribe Schismatoglottideae Nakai (1943).
Bull. Singapore 38: 43-54 (1985); Bogner, Aroideana 8: 73-
75 (1986).
Evergreen herbs; usually epigeal; leaves several
Diandriella Engler (1910). per article; petiole sheath usually with long,
marcescent, apical ligule; leaf blade apex with
Rarely robust; petiole rarely aculeate; leaf blade tubular mucro (except most Schismatoglottis
lanceolate to sagittate, rarely peltate; inflores- spp.), primary lateral veins pinnate, forming 1
cences 1-6; peduncle short; spathe boat-shaped; usually prominent marginal vein, finer venation
spadix stipitate or sessile, female and male zones parallel-pinnate; inflorescence usually 1; spathe
contiguous or with sterile male flowers or naked tube persistent, blade usually white (except
zone in between; male flower ( 1-)2-4-androus, Piptospatha), caduceus at anthesis, rarely
thecae usually opening by longitudinal (more marcescent (Schismatoglottis texturata (Schott)
rarely transverse) slit, pollen usually extruded Engl., Hottarum lucens Bogner); stamens usually
in strands; female flower usually with 1(-3) free, sometimes filaments ± connate, often ±
staminodes, ovary incompletely 2-4( -5)-locular, elongated; ovary 1-locular, ovules atropous
ovules many, hemianatropous, placentas parietal, to hemitropous (except Schismatoglottis); testa
or parietal above and axile below, or basal; seed ± usually costate, embryo axile, endosperm
ellipsoid, testa thick, costate. 2n = 38, 40, 42, 80. copious.
About 110 spp., S and SE Asia to Melanesia, tropi-
cal America (Costa Rica to Bolivia), terrestrial and 49. Schismatoglottis Zollinger & Moritzi
rheophytic; five sections recognised.
Schismatoglottis Zollinger & Moritzi in Moritzi, Syst.
Verzeichniss Zollinger: 83 (1846); Bunting, Ann. Mo. Bot.
7. Tribe Anubiadeae Engler (1876). Gard. 47: 69-71 (1960); Hotta, Mem. Coli. Sci. Univ. Kyoto,
ser. B, 32: 19-30 (1965) & ibid.: 223-238 (1966); Bunting &
Roots with sclerotic hypodermis; petiole geni- Steyermark, Brittonia 21: 187-190 (1969); Bogner & Hotta,
culate apically; leaf blade elliptic, lanceolate to Acta Phytotaxon. Geobot. 34: 48-50 (1983).
Philonotion Schott (1857).
hastate-trifid; primary lateral veins pinnate, form-
ing 1 marginal vein, finer veins parallel-pinnate
with ± distinct transverse veins; spathe boat- Stem rhizomatous or epigeal, shortly erect; petiole
shaped, unconstricted, persistent; stamens con- sheath less than half petiole length, sometimes
nate into ± prismatic synandrium; thecae lateral ligulate; leaf blade narrowly elliptic to cordate-
or marginal, rarely covering whole synandrium, sagittate, rarely with apical tubular mucro; inflo-
dehiscing by longitudinal slit, fused connectives rescences 1-3; spadix zonation variable: female
thickened, ± truncate; ovary (1-)2-3-locular, zone sometimes bearing sterile organs at the very
ovules many per locule, anatropous, placenta base, female and male zones sometimes with
axile; berry depressed-globose to obovoid; seed naked zone or zone of sterile flowers in between,
small, ± ovoid to subcylindric, endosperm upper part of spadix clavate to cylindric, terminal
copious. appendix of sterile flowers sometimes present;
male flowers: 2-3-androus, filaments usually dis-
tinct, sometimes connate basally; pollen extruded
48. Anubias Schott in strands; staminodes obpyramidal to clavate,
usually truncate; female flowers: sometimes with
Anubias Schott, Oesterr. Bot. Wochenbl. 7: 398 (1857); Crusio,
Meded. Landbouwhogesch. Wageningen 79 (14): 1-48 whorl of 1-4 staminodes, ovules (1-)few to nu-
(1979), rev. & Aqua-Planta, Sonderheft 1: 1-14 (1987). merous, anatropous to hemianatropous, placentas
Amauriella Rendle (1913). 1-4, parietal, usually extending from base to
 Araceae 57

apex of locule; berry oblong to globose, green or 52. Bucephalandra Schott

dull yellow or deep red, few- to many-seeded;
Bucephalandra Schott, Gen. Aroid.: t. 56 (1858); Bogner,
seed shortly ellipsoid, straight. 2n = 26, 39, 52. Aroideana 3: 134-143 (1980) & Plant. Syst. Evol. 145: 159-
About 120 spp., SE Asia to New Guinea, northern 164 (1984).
S America, on forest floor, terrestrial or Microcasia Beccari ( 1879).
rheophytic.
Stem rhizomatous, decumbent; leaf blade elliptic,
oblanceolate or obovate; spathe not constricted,
50. Piptospatha N.E. Brown
ellipsoid, tube funnel-shaped, blade gaping and
Piptospatha N.E. Brown, Gard. Chron. II, 11: 138 (1879); caducous; spadix shorter than spathe, with a few
Hotta, Mem. Coil. Sci. Univ. Kyoto, ser. B, 32: 19-30 (1965). sterile flowers at extreme base, female zone nar-
Gamogyne N.E. Brown (1882). rower than upper parts, separated from male zone
by about 2 rows of flattened, scalelike staminodes,
Stem decumbent or erect; leaf blade narrowly male zone with 2-5 rows of flowers, apical zone
lanceolate to elliptic; inflorescence ± nodding; globose, ellipsoid to subcylindr~c, c?mpos~d of
spathe not constricted, stoutly ellipsoid; spadix truncate, obpyramidal to subcylmdnc, papillose
subsessile, sometimes with a few sterile flowers at sterile male flowers, the uppermost ± connate;
extreme base, female and male zones contiguous male flower 1-androus; ovary depressed-globose,
or with short zone of sterile male flowers between, ovules many, placenta basal; berry with numerous
apex sometimes with a few sterile flowers; male seeds; seed narrow-ellipsoid, with long, curved
flower 1-2-androus; gynoecia free or superficially micropylar appendage. 2n = ca. 26. Three spp.,
connate (coherent), ovules many, placentas 2-4, Borneo, rheophytes in tropical humid forest.
parietal or parietal and basal; berry obovoid; seed
elongate-ellipsoid to cylindric with long micropy-
lar appendage. 2n = 26. Ten spp., Borneo, Malay 53. Phymatarum M. Hotta
Peninsula; terrestrial or rheophytic; closely related
to Hottarum. Phymata rum M. Hotta, Mem. Coli. Sci. Univ. Kyoto, ser. B, 32:
29 (1965); Bogner, Plant. Syst. Evol. 144: 62-66 (1984).

51. Hottarum Bogner & Nicolson Stem rhizomatous, decumbent; leaf blade nar-
rowly elliptic; spathe constricted, blade boat-
Hottarum Bogner & Nicolson, Aroideana 1: 72 (197~, :'1978");
Bogner, Plant. Syst. Evol. 142: 49-55 (1983) & tbtd., 145: shaped, gaping, caducous after anthesis; spadix
159-164 (1984); Bogner & Hotta, Adansonia IV, 5: 27-31 with extreme base usually bearing a few sterile
(1983). flowers, female zone adnate to spathe, conoid to
Microcasia sect. Truncatae M. Hotta, Mem. Coil. Sci. Univ. subcylindric, separated from male zone by cylin-
Kyoto, ser. B, Bioi. 32: 21 (1965).
dric to ellipsoid zone of sterile male flowers, male
zone very short and slightly narrower, terminal
Stem ± erect; leaf blade ± elliptic; spathe con- appendix much longer, elongate-conoid, com-
stricted or not; spadix sometimes adnate to spathe posed of sterile male flowers; male flower 1-
for two-thirds oflength (H. lucens Bogner), female androus; sterile flowers subprismatic, tuberculate,
zone sometimes with sterile flowers at base, flattened or excavated; ovary depressed-globose,
either contiguous with male, or with a few sterile ovules many, placenta basal; berry many-seeded,
male flowers in between, terminal appendix broadly obovate-oblong, greenish white; seed el-
of sterile flowers present; male flower (1- )2- lipsoid, with long micropylar appendage. 2n = 26.
3( -4)-androus, sterile male flowers composed of Three spp., Borneo, rheophytes in forest, ruderal
truncate, prismatic staminodes; female flower habitats, and disused rice paddi.
sometimes with 1 clavate staminode in basal
flowers, ovary depressed-globose, ovules 10-15,
placenta basal; berry ± globose, whitish green, 54. Aridarum Ridley
many-seeded; seed ellipsoid to elongate, with
long, micropylar appendage. 2n = 26. Six spp., Aridarum Ridley, J. Bot. 51: 201 (1913); Hotta, Mem. Coli. Sci.
Univ. Kyoto, ser. B, 32: 19 (1965); Bogner, Aroideana 2: 110-
Borneo; tropical humid forest, rheophytes on 121 (1979) & Aroideana 4: 57-63 (1981) & Blumea 28: 403-
rocks in and along streams; H. lucens is rather 405 (1983).
distinct.
Stem decumbent; leaves sometimes distichous;
leaf blade linear to elliptic; inflorescence often
58 Araceae

somewhat nodding; spathe not constricted, 56. Lagenandra Dalzell

stoutly ellipsoid, blade caducous; spadix normally
Lagenandra Dalzell, Hooker's J. Bot. Kew Gard. Misc. 4: 289
with a few sterile flowers at base, female zone cy- (1852); De Wit, Meded. Landbouwhogesch. Wageningen
lindric, separated from male by zone of sterile 78: 5-45 (1978), rev. & Aquariumplanten: 286-307 (1983);
flowers, short terminal appendix of sterile flowers Bogner & Jacobsen, Aqua-Planta 12: 43-50 {1987);
present; male flower 1-2-androus, thecae apically Nicolson, Fl. Ceylon 6: 75-85 (1988).
narrowed into long or short horn, dehiscing by
apical pore; sterile male flowers of truncate Leaf blade involute in bud, linear to ovate; pe-
staminodes often with small central cavity; ovary duncle usually short; spathe thick-walled, margins
laterally compressed, ovules many, placenta basal; free above kettle, mouth of spathe tube forming a
berry globose or ellipsoid to cylindric; seed ellip- narrow aperture, blade straight or twisted, open-
soid, elongate. 2n = 24. Seven spp., Borneo, ing by narrow slit or widely gaping; spadix: female
rheophytes in humid forest. zone subglobose, separated by slender naked axis
from male zone, appendix small, conoid; gynoecia
free, in spirals or rarely whorled, ovules 1-
55. Heteroaridarum M. Hotta 12( -15), placenta basal, style absent or distinct,
Heteroaridarum M. Hotta, Acta Phytotaxon. Geobot. 27: 63 stigma broad; infructescence usually globose, ber-
(1976). ries free, dehiscent basally with pericarp becoming
revolute; testa costate. 2n = 36, 72. Fourteen spp.,
Stem short; leaf blade elliptic, coriaceous; inflores- Indian subcontinent, Sri Lanka; freshwater aquat-
cence 1; spathe slightly constricted; spadix: ics, usually helophytes, rarely rheophytes.
female zone separated from male by short zone
of sterile flowers, male zone much longer; male 57. Cryptocoryne Fischer ex Wydler
flowers: 3-androus, 2 outermost stamens larger
with shortly horned thecae, central stamen Cryptocoryne Fischer ex Wydler, Linnaea 5: 428 (1830);
smaller with hornless thecae, all thecae opening by De Wit, Aquariumplanten: 141-261 (1983); Jacobsen,
Cryptocorynen, 112pp. (1982) & Nord. J. Bot. 5: 31-50
apical pore; sterile male flowers composed of (1985); Jacobsen & Bogner, Aqua-Planta 12: 96-103 (1987);
basally connate, truncate staminodes; ovary Jacobsen et al., Aqua-Planta 14: 127-132 (1989); Nicolson,
depressed-globose, ovules many, placentas 2, Fl. Ceylon 6: 85-99 (1988).
basal and apical, apical one bearing smaller and
apparently sterile ovules; berry and seed un- Leaf blade convolute in bud, linear to cordate; pe-
known. 2n = ? Only one sp., H. borneense M. duncle short, elongating in fruit; spathe margins
Hotta, Borneo, rheophyte. connate above kettle (except C. spiralis, C.
cognata), blade cordate to lanceolate, variously
coloured, ± spreading or twisted, often with a dis-
9. Tribe Cryptocoryneae Blume (1836).
tinct collar at mouth of spathe tube; spadix: fe-
Aquatic, evergreen, rhizomatous; leaves several; male zone a single whorl of 4-8 gynoecia, usually
primary lateral veins ascending, fusing distally separated from male zone by slender naked axis,
into an inconspicuous marginal vein, finer vena- appendix short, conoid to subclavate; ovaries
tion reticulate; inflorescence 1; spathe tube mar- connate, ovules 5-many, placenta subbasal to ob-
gins basally or entirely connate, usually slightly liquely parietal (morphologically basal); berries
swollen basally forming a "kettle", blade lan- connate forming ovoid syncarp which dehisces
ceolate to cordate, often twisted or long-spiralled apically as one unit to become star-shaped; seed
or long-caudate; spadix entirely enclosed in ellipsoid-oblong, testa rough to subcostate, em-
kettle of spathe tube, sterile terminal appendix bryo well developed. 2n = 20, 22, 28, 30, 33, 34, 36,
apically adnate to spathe at apex of kettle, female 42, 54, 66, 68, 72, 85, 88, 90, 102, 132. About 50 spp.,
zone with a whorl of pistillodial olfactory bodies S and SE Asia to New Guinea, submerged or emer-
situated above the fertile gynoecia (except some gent, freshwater tidal zone, forest streams and
Lagenandra spp.); male flowers 1(-2)-androus, pools; rarely seasonally dormant; Jacobsen (1982)
stamens free, anther sessile or with short filament, recognised 26 informal subgroups.
apex excavated with prominent margins, thecae
ending in prominent horn, dehiscing by apical
10. Tribe Zomicarpeae Schott (1856).
pore; pollen exine smooth; ovary 1-locular,
ovules atropous; embryo elongate, endosperm Laticifers anastomosing; small, evergreen or
copious. seasonally dormant; leaf blade cordate-sagittate
 Araceae 59

(except Zomicarpa); primary veins radiating Rhizomatous; leaves few; leaf blade usually varie-
from base, finer venation reticulate; spadix longer gated; inflorescence 1; peduncle subequalling or
than spathe (except Zomicarpa and 1 sp. of longer than leaves; spathe unconstricted, fully
Zomicarpella), sterile terminal appendix present, expanded, margins reflexed at anthesis; spadix:
female spadix zone adnate to spathe; stamens free; female zone separated from male by sterile zone
gynoecia few, ovules anatropous, placenta basal, which is basally either naked or with a few scat-
style short, stigma discoid. tered sterile flowers and apically bears a short
dense zone of staminodes, male zone short,
few-flowered, sterile terminal appendix present,
58. Zomicarpa Schott staminodial at base, otherwise smooth, sub-
Zomicarpa Schott, Syn. Aroid.: 33 (1856); Bogner, Aroideana cylindric; male flowers 2-3-androus; gynoecium
3: 10-11 (1980}. slender, ovule 1; testa thin, embryo relatively
large, undifferentiated, endosperm absent. 2n =
Stem tuberous; leaves few; leaf blade trisect to 14. Only one sp., U. sagittatum Engler, S America
pedatisect, leaflets with distinct submarginal (western Amazonia), on forest floor.
collective veins; inflorescences 3-5, subtended
by cataphylls, appearing before or with leaves;
61. Filarum Nicolson
peduncle subequal to leaf or longer; spathe ±
constricted, tube ± gaping at anthesis, blade Filarum Nicolson, Brittonia 18: 348 (1967}.
expanded at anthesis, sometimes fornicate, ±
lanceolate; spadix subequal to spathe, female Tuberous; leaves few; inflorescence 1-3, ap-
zone laxly flowered, contiguous with male, pearing with leaves; peduncle slender; spathe
appendix smooth or staminodial; male flower oblong-lanceolate, unconstricted, fully expanded,
1-2-androus; ovules 6-9, stigma broad, sub- persistent; spadix very slender, male zone elon-
sessile; berry depressed-globose, whitish, few- gate, sparsely flowered, a few sterile rudimentary
seeded; seed ovoid to oblong-ellipsoid with flowers above and below male zone, appendix
large, strophiolate funicle, testa smooth, thin, slender, long; male flowers 1-androus; ovary
transparent, embryo elongate, partly green, oblong, ovule 1; berry subglobose; testa thin,
endosperm copious, green and starchy. 2n = smooth, embryo large, endosperm absent. 2n = 28
20. Three spp., NE Brazil, geophytes, seasonally Only one sp., F. manserichense Nicolson, Amazo-
dormant. nian Peru, geophyte.

59. Zomicarpella N.E. Brown 11. Tribe Caladieae Schott (1832).

Laticifers anastomosing; terrestrial, geophytic,
Zomicarpella N.E. Brown, Gard. Chron. II, 16: 266 (1881}; ·
Bogner, Aroideana 1: 72 (1979, "1978"}. rarely aquatic (]asarum), climber or hemi-
epiphyte (Syngonium); stem tuberous, rhizoma-
Stem rhizomatous; leaves 1-few; leaf blade usually tous or aerial; leaf basal ribs well developed,
variegated; inflorescences 1-2, appearing with venation pinnate, forming submarginal collective
leaf; peduncle equalling petiole; spathe oblong- vein, 1 or more marginal veins also usually
lanceolate, completely expanded or lower part present, finer venation reticulate; spadix: female
partially convolute, not or slightly constricted; zone often separated from male by subconical
spadix longer than or subequal to spathe, female to attenuate zone of synandrodes, male zone
zone contiguous with male or with short sterile subcylindric to subclavate, sterile appendix absent
zone in between, appendix slender, long; male (except Hapaline appendiculata); flowers uni-
flowers 1-androus; pollen extruded in strands; sexual, perigone absent; stamens connate forming
ovary ellipsoid-oblong, ovules 1-6; berry white, 1- synandrium; pollen extruded in strands (except
few-seeded; testa thin, smooth, embryo small, en- Scaphispatha); ovules anatropous to hemi-
dosperm copious. 2n = 26. Two spp., northern S anatropous; endosperm copious (except ]asarum,
America (western Amazonia), geophytes, season- Syngonium, Hapaline).
ally dormant.
62. Scaphispatha Brongn. ex Schott
60. Ulearum Engler
Scaphispatha Brongn. ex Schott, Prodr. Syst. Aroid.: 214
Ulearum Engler, Bot. Jahrb. 37: 95 (1905}. (1860}; Bogner, Aroideana 3: 4-12 (1980).
60 Araceae

Stem a subglobose tuber; leaves 1-2; leaf blade 65. Xanthosoma Schott Fig. 15B-K
peltate, ovate-subcordate to -subsagittate; inflo-
Xanthosoma Schott in Schott & Endlicher, Melet. Bot.: 19
rescence 1, appearing before leaves; peduncle (1832); Madison, Phytologia 35: 103-107 (1976) & Selbyana
long, slender; spathe ovate, fully expanded at an- 5: 342-377 (1981); Grayum, Ann. Mo. Bot. Gard. 73: 462-
thesis, lower part later convolute and persistent, 474; Mayo & Bogner, Willdenowia 18: 231-242 (1988).
upper part marcescent; spadix shorter than
spathe, male and female zones contiguous; Sometimes gigantic and arborescent; tuberous;
synandrium 4-androus; ovary 1-locular, ovules leaf blade cordate-sagittate, hastate, tri- or
3-5, placenta basal; berry subglobose to obovoid, pedatisect, rarely linear-lanceolate or peltate;
1-seeded; seed subglobose, testa smooth and inflorescences 1-several, always appearing with
thin. 2n = 28. Only one sp. S. gracilis Brongn. leaves; peduncle usually rather short; spathe
ex Schott, Bolivia, Brazil, geophyte, seasonally tube usually ventricose; synandrium 4-6-androus;
dormant. pollen shed in tetrads; ovary incompletely ( 1-)2-
4-locular, ovules ca. 20-many, placentas (1- )2-4,
intrusive-parietal; berry many-seeded; seed ovoid,
63. Caladium Vent.
testa costate. 2n = 26, 39, 52. About 57 spp.,
Caladium Vent., Descript. Plant. Nouv. Jard. Cels: 30 (1801); Mexico to northern Argentina, on forest floor
Madison, Phytologia 35: 103-107 (1976) & Selbyana 5: 342- or seasonally flooded sites, sometimes weedy,
377 (1981), rev.; Bogner, Aroideana 7: 4-5 (1984); Grayum, evergreen or seasonally dormant. The nutritious
Ann. Mo. Bot. Gard. 73: 462-474 (1986); Mayo & Bogner,
Willdenowia 18: 231-242 (1988). tubers of the X. sagittifolium complex are an
Aphyllarum S. Moore (1895). important subsistence food throughout the humid
tropics.
Stem a subglobose tuber; leaf blade usually
peltate, often variegated, cordate-sagittate or sag-
66. Chlorospatha Engler
ittate, rarely trisect; inflorescences 1-2, appearing
with or before leaves; peduncle subequal to peti- Chlorospatha Engler, Gartenflora 27: 97 (1878); Madison,
ole; spathe tube usually ventricose-globose; spadix Selbyana 5: 342-377 (1981), rev.; Bogner, Aroideana 8: 48-
subequal to spathe; synandrium 3-5-androus, 54 (1985); Grayum, Ann. Mo. Bot. Gard. 73:462-474 (1986);
Mayo & Bogner, Willdenowia 18: 231-242 (1988); Grayum,
margins sinuous; ovary 1(-3)-locular, ovules l-ea. Novon 1: 12-14 (1991).
20, placentas 1-3, subbasal to intrusive-parietal; Caladiopsis Engler (1905).
berry white, 1- to many-seeded; seed ovoid to
ellipsoid, testa longitudinally costate. 2n = 22, Stem caulescent, decumbent to erect; leaves 1-few;
26, 28, 30, 32. About 12 spp., tropical America, petiole sheath relatively long; leaf blade cordate,
geophytes, seasonally dormant or evergreen; sagittate, hastate, trisect or pedatisect; inflores-
C. bicolor (Ait.) Vent. and its many cultivars are cences 3-8; peduncle slender, supported by peti-
horticulturally important worldwide and often ole sheath; spathe tube narrow, elongate; female
naturalised in tropical regions. zone of spadix free or adnate to spathe; female
and sterile flowers often distant; synandrium 3-5-
64. ]asarum G.S. Bunting androus; pollen shed in tetrads; ovary incom-
pletely (1- )2-4-locular, ovules few to several,
]asarum G.S. Bunting, Acta Bot. Venez. 10:264 (1977, "1975"); placenta intrusive-parietal, rarely basal; berry ::!::
Bogner, Aroideana 8: 55-63 (1985). 3-5-furrowed, white to yellowish; seed minute,
costate. 2n = 26. About 16 spp., Costa Rica to
Stem hypogeal, rhizomatous, erect; leaves several; Peru, on forest floor, evergreen.
leaf blade linear, ::!:: bullate; peduncle long;
spathe borne above water level, blade caudate;
female zone of spadix basally adnate to spa- 67. Syngonium Schott
the; synandrium 3-4-androus; ovary 1-locular, Syngonium Schott, Wiener Z. Kunst 1829 (3): 780 (1829);
( 1-)2-3( -4)-ovulate, ovule anatropous, placenta Croat, Ann. Mo. Bot. Gard. 68: 565-651 (1982), rev.
basal; berry 1(-2)-seeded; seed large, obovoid, Porphyrospatha Engler (1879).
testa smooth, embryo large, plumule well
developed, endosperm very sparse. 2n = 22. Only Climbers or hemiepiphytes with elongated inter-
one sp., f. steyermarkii G.S. Bunting, Venezuela, nodes; leaf blade sagittate, trisect, pedatisect,
Guyana, submerged aquatic in blackwater rarely pinnatifid; inflorescences several, pendent
streams. in fruit; peduncle short; spadix much shorter than
 Araceae 61

connate, syncarp stoutly ovoid, usually brown;

seed rather large, testa smooth, thin, embryo
large, endosperm absent. 2n = 28 (24, 26). Thirty
five spp., Mexico to SE Brazil, sometimes natu-
ralised elsewhere in the tropics; some species, e.g.
S. podophyllum Schott, are important foliage
ornamental plants in horticulture.

68. Hapaline Schott

Hapaline Schott, Gen. Aroid.: t. 44 ( 1858), nom. cons.; Bogner,
Plant. Syst. Evol. 144: 59-61 (1984); Boyce, Kew Bull. 51: 63-
82 (1996), rev.

Stem tuberous; leaf 1-few; leaf blade cordate to

sagittate; inflorescence appearing with leaves;
peduncle slender, longer than petiole; spathe not
distinctly constricted, tube short, blade linear-
lanceolate; spadix subequal or longer than spathe,
female zone of2-3 flowers, adnate to spathe, sepa-
rated from male by short zone of sterile flowers,
terminal appendix filiform (when present);
synandrium 3-androus, peltate, thecae almost
pendent from margin; ovary 1-locular, ovule 1,
anatropous, placenta parietal-subbasal; berry
white, 1-seeded; testa smooth, embryo large,
endosperm absent. 2n = 26, 28. Six spp., Borneo,
geophytes, seasonally dormant.

12. Tribe Nephthytideae Engler (1887)

Petiole geniculum present; leaf essentially tripar-
tite, main venation pinnate, fine venation reticu-
late; spathe boat-shaped or fully expanded, not
clearly differentiated into tube and blade; spadix
cylindric, female and male zones usually conti-
guous, sterile flowers ± absent (except Pseudo-
.; hydrosme); stamen filaments lacking or very short
(except Nephthytis); ovules 1 per locule, anatro-
.
pous, placenta basal (except Pseudohydrosme);
. berry and seed large, testa thin, endosperm
., .\~<r
absent. Chromosomes large to very large .
K

Fig. 15A-K. Araceae-Aroideae. A Colocasia esculenta, habit.

B-F Xanthosoma sagittifolium. B Inflorescence, spathe partly
69. Nephthytis Schott
removed. C Group of synandria. D Synandrium, longitudinal Nephthytis Schott, Oesterr. Bot. Wochenbl. 7: 406 (1857);
section. E Pistil, side view. F Same, longitudinal section. G-L Bogner, Aroideana 3: 75-85 (1980); Knecht, Contrib. Arac.
X. platylobum. G Habit. H Spadix. I Synandrium, side view. Cote d'Ivoire: 163-171 (1983); Namur & Bogner, Adansonia
J Pistillate flowers, seen from above. K Pistil, side view. IV, 16:71-74 (1994); Ntepe-Nyame, Fl. Cameroun 31:46- 56
(Takhtajan 1982) (1988); Hay eta!., Novon 4: 365-368 (1994).

Rhizome creeping, usually subepigeal, rarely hy-

spathe; synandrium 3-4-androus; gynoecia partly pogeal; leaves usually several; leaf blade cordate-
to completely connate, eventually forming syn- sagittate, deeply sagittate or tripartite, primary
carp, ovary (1-)2(-3)-locular, ovules 1(-2) per lateral veins ascending, joining into a submarginal
locule, anatropous, placenta axile-basal; berries collective vein; inflorescence 1; peduncle relatively
62 Araceae

long; spathe fully expanded, ovate-oblong-elliptic, than ovary, stigma broad, discoid, concave; berry
often reflexed at anthesis; spadix stipitate; male large; seed large, ellipsoid, testa smooth, thin,
flowers 2-4-androus; ovary 1-locular; berry sub- endosperm absent.
globose to ellipsoid, orange; seed obovoid or
subglobose, testa absent at maturity. 2n = 40, 60.
72. Aglaonema Schott
Ten spp., W and WC Africa (Liberia to Gabon),
one sp. in Borneo. Aglaonema Schott, Wiener Z. Kunst 1829 (3): 892 (1829);
Nicolson, Smithson. Contrib. Bot. 1: 1-69 (1969), rev.

70. Anchomanes Schott Stem creeping to erect, internodes green, smooth,

Anchomanes Schott, Oesterr. Bot. Wochenbl. 3: 314 (1853);
distinct; leaf blade often variegated; inflorescences
Knecht, Contrib. Arac. Cote d'Ivoire: 145-151 (1983); Mayo 1-9; peduncle shorter or longer than petiole,
& Bogner, Fl. Trop. East Africa, Araceae: 24-28 (1985); deflexing in fruit; spadix cylindric to clavate,
Ntepe-Nyame, Fl. Cameroun 31: 22-26 (1988). subsessile to stipitate; female zone ± few-
flowered, male zone longer; male flower 1-
Robust to gigantic; rhizome hypogeal, erect or androus, filament ± distinct; ovary subglobose,
horizontal; leaf 1; petiole very long, aculeate placenta basal; berry red at maturity. 2n = 40,
(except A. boehmii), geniculate at main blade 60, 80, 100, 120 (70, 110). Twenty one spp., S and
divisions; leaf blade dracontioid, ultimate lobes SE Asia to New Guinea. Important ornamental
trapezoid, truncate or shallowly bifid; inflores- foliage plants.
cence 1; peduncle long; spathe boat-shaped; spa-
dix short or subequal to spathe; ovary 1-locular,
style sometimes deflexed; berry oblong-ellipsoid, 73. Aglaodorum Schott
red, purplish or whitish; seed obovoid to oblong- Aglaodorum Schott, Gen. Aroid.: t. 58 (1858); Nicolson,
ovoid. 2n = 40. Four to five spp., tropical Africa Smithson. Contrib. Bot. 1: 3-4 (1969).
(Gambia to Zambia); tropical moist forests, savan-
nas, swamps, seasonally dormant. Rhizome much-branched, creeping; leaves several
per article; leaf blade thick, midrib very thick;
inflorescence 1; peduncle long; spadix stipitate,
71. Pseudohydrosme Engler female zone with 1 whorl of flowers; male
Pseudohydrosme Engler, Bot. Jahrb. 15: 455 (1892, "1893"); flowers 3-4-androus; gynoecium with whorl of
Bogner, Aroideana 4: 31-37 (1981). 1-3 short, prismatic staminodes, ovary 1-2-
Zyganthera N.E. Brown (1901). locular, placenta parietal; berry green with very
thick pericarp; seed plumule well developed.
Robust; tuber subglobose, hypogeal; leaf 1; petiole 2n = 40. Only one sp., A. griffithii (Schott) Schott,
and blade as for Anchomanes; inflorescence 1; pe- Borneo, Malay Peninsula, Sumatra, in open
duncle very short; spathe ovate-elliptic-oblong, ± swamps, especially freshwater tidal zones and
boat-shaped, convolute basally into obconic tube, brackish water.
upper part very broad, fornicate with auriculate
margins; spadix short, sessile, male zone with
apical appendix in P. buettneri Engler; stamens 14. Tribe Culcasieae Engler (1887).
2-5; staminodes of appendix (when present) Usually climbers or hemiepiphytes with slender
subprismatic, free; ovary 2(-3)-locular, placenta stems, sometimes erect or prostrate terrestrial
axile-basal, style short; berry and seed unknown. plants; roots, stems and leaves containing resin
2n = ca. 40. Two spp., Central-W Africa (Gabon), canals; sclerotic hypodermis present in roots;
seasonally dormant. petiole geniculate apically; primary veins pinnate,
finer venation reticulate; spadix: female zone con-
13. Tribe Aglaonemateae Engler (1876). tiguous with male, or with zone of sterile male
flowers in between; male flower: 2-4 androus, an-
Leaf blade (linear-)ovate to elliptic-oblong, base thers sessile, lateral, lacking endothecial thicken-
sometimes subcordate; primary veins pinnate, ings, thecae dehiscing by short apical slit or pore,
finer veins parallel-pinnate; spathe boat-shaped, connective usually strongly thickened; ovary
unconstricted; spadix: female and male zones con- locules 1-ovulate; stigma sessile, large, discoid;
tiguous; thecae dehiscing by apical pores; ovary 1- berry ± globose, usually red; testa thin, endo-
locular, ovule 1, anatropous, style short, narrower sperm absent.
 Araceae 63

74. Culcasia Palisot de Beauvais ovules 1-2, anatropous, placenta subbasal; berry
and seed large, plumule well developed, en-
Culcasia Palisot de Beauvais, Fl. Oware 1: 3 (1805), nom. cons.;
Bogner, Misc. Pap. Landbouwhogesch. Wageningen 19:57- dosperm absent.
63 (1980); Knecht, Contrib. Arac. Cote d'Ivoire: 188-227
(1983); Mayo, Fl. Trop. East Africa, Araceae: 16-22 (1985);
Ntepe-Nyame, Fl. Cameroun 31: 76-117 (1988). 76. Montrichardia H. Cruger Fig. 16
Montrichardia H. Cruger, Bot. Zeitung (Berlin) 12: 25 (1854),
Laticifers absent; sometimes erect, terrestrial; leaf nom. cons.; Jonker-Verhoef & Jonker, Fl. Suriname 1: 75-79
blade simple; inflorescences 1-12(-20 or more); (1953).
spathe boat-shaped, convolute basally, deciduous
or persistent; spadix sessile or stipitate, cylindric- Robust to gigantic; rhizomatous; leaves several
clavate, equalling or exceeding spathe; ovary 1-3- in apical crown; petiole sheath apically ligulate;
locular, locules 1-ovulate, ovule anatropous or inflorescence 1(-2); peduncle relatively short;
campylotropous, placenta axile, basal or basal- spathe thick; spadix subequal to spathe; stigma
parietal, berry red to orange, sometimes greenish appearing sessile on stylar region; berry sub-
yellow; seed ovoid to ellipsoid. 2n = 42, 84. About cylindric, pericarp spongiose; seed obovoid to
27 spp., tropical Africa (Senegal to Tanzania), ellipsoid, testa smooth to rough. 2n = 48. Two
root-climbers, hemiepiphytes, or erect and spp., Belize to eastern Brazil, freshwater
terrestrial. helophytes forming extensive stands along river
margins.
75. Cercestis Schott
16. Tribe Zantedeschieae Engler (1887).
Cercestis Schott, Oesterr. Bot. Wochenbl. 7: 414 (1857);
Knecht, Contrib. Arac. Cote d'Ivoire: 172-187 (1983 ); Stem a depressed-globose tuber or short rhizome
Bogner, Aroideana 8: 73-75 (1986); Ntepe-Nyame, Fl. (Z. aethiopica); leaf blade lanceolate to hastate-
Cameroun 31: 64-75 (1988); Bogner & Knecht, Adansonia sagittate; primary lateral veins pinnate, forming 1
IV, 16: 331-335 (1994).
Rhektophyllum N.E. Brown (1882); Mayo, Fl. Trop. East marginal vein, finer venation parallel-pinnate;
Africa, Araceae: 22-24 (1985); Ntepe-Nyame, Fl. Cameroun peduncle long; spathe :±:: obconic, unconstricted;
31: 58-64 (1988). spadix shorter than spathe, male and female zones
contiguous; stamens free, thecae dehiscing by
Laticifers present; sometimes robust, producing apical pore, connective thickened apically; pollen
flagelliform shoots, internodes short in flowering
zone of stem; leaf blade oblong-lanceolate to cor-
date, sagittate, hastate or tripartite, sometimes
laciniate-pinnatifid; inflorescences 1-16; peduncle
short; spathe boat-shaped to subcylindric, convo-
lute basally, persistent; spadix sessile, shorter than
spathe; ovary 1-locular, ovule hemianatropous
to anatropous, rarely campylotropous, placenta
basal-parietal; seed obovoid. 2n = 42. Thirteen
spp., Senegal to western Uganda, root -climbers or
hemiepiphytes, rarely creeping on forest floor.

15. Tribe Montrichardieae Engler (1876).

Arborescent helophytes; sclerotic hypodermis
present in roots; petiole geniculum absent; leaf
blade sagittate, hastate to trifid (rarely trisect),
primary lateral veins pinnate; spathe deciduous
after anthesis, slightly constricted, differentiated
into lower tube and upper, :±:: boat-shaped blade;
spadix: female and male zone contiguous, sterile
flowers absent; male flower: 3-6-androus, stamens Fig. 16A-E. Araceae-Aroideae. Montrichardia arborescens. A
:±:: sessile, connective overtopping thecae, thecae Flowering shoot. B Spadix. C Stamens. D Pistil, side view. E
dehiscing by short, apical slit; ovary 1-locular, Same, longitudinal section. F Seed. (Takhtajan 1982)
64 Araceae

extruded in strands; ovary 1-5-locular, locules certain species of Nephthytis and to tribe
(1-)4( -B)-ovulate, ovules anatropous, placenta ax- Zomicarpeae.
ile to subapical; testa striate, endosperm copious.
18. Tribe Thomsonieae Blume (1837).
77. Zantedeschia K. Sprengel Seasonally dormant; stem tuberous, rarely rhi-
Zantedeschia K. Sprengel, Syst. Veg. 3: 756, 765 (1826), nom. zomatous; petiole long, terete, sheath very short;
cons.; Letty, Bothalia 11: 5-26 (1973), rev.; Perry, S. Afr. J. leaf blade dracontioid, first foliage leaf of seedling
Bot. 55: 447-451 (1989). divided, all divisions upright in bud; ultimate
Pseudohomalomena A.D. Hawkes (1951). lobes usually ovate-elliptic, acuminate, primary
lateral veins forming submarginal collective veins,
Leaves several; leaf blade often variegated; inflo- finer venation reticulate; inflorescence 1, sub-
rescences 1-2; spathe white, cream, yellow, pink tended by cataphylls; spadix with sterile terminal
or rosy purple; spadix sessile or stipitate; male appendix, female zone usually contiguous with
flowers 2-3-androus; gynoecium rarely (only male; pollen extruded in strands; ovary locules 1-
in Z. aethiopica (L.) Sprengel) with whorl of about ovulate, ovules anatropous, testa smooth, embryo
3 ± spathulate staminodes, otherwise staminodes large, endosperm absent.
absent, ovary shortly ovoid; berry obovoid or
subglobose, usually green or orange, rarely yellow
and mucilaginous within (Z. aethiopica); seed 79. Amorphophallus Blume ex Decaisne
subglobose to ovoid, embryo axile. 2n = 32. Eight Amorphophallus Blume ex Decaisne, Nouv. Ann. Mus. Hist.
spp., Angola to S Africa, helophytic or terrestrial, Nat. 3: 366 (1834), nom. cons.; Hetterscheid & Ittenbach,
seasonally dormant, sometimes evergreen (Z. Aroideana 19: 7-131 (1996).
aethiopica); Z. aethiopica is a familiar garden Thomsonia Wallich (1830), nom. rej.; Bogner, Pl. Syst. Evol.
125: 15-20 (1976).
plant with many hybrids and is of considerable Plesmonium Schott (1856); Bogner, Adansonia II, 20: 305-308
importance in the cut flower trade. (1980).

Sometimes gigantic; tuber depressed-globose to

17. Tribe Callopsideae Engler (1906). cylindric, rarely rhizomatous; leaf usually 1; inflo-
Small, evergreen, rhizome slender; leaf blade rescence usually appearing before leaf; peduncle
cordate-ovate, primary lateral veins forming very short to long; spathe either ± undifferenti-
single marginal vein, finer venation reticulate; ated into tube and blade, or with campanulate,
spathe completely expanded at anthesis, persis- cylindric or ventricose, convolute tube and erect
tent, partially closing in fruit; spadix fertile to to spreading blade, rarely connate basally (A.
apex, female zone entirely adnate to spathe, con- pusillus Hetterscheid & Serebryanyi); spadix male
tiguous with male zone or with short naked zone zone rarely separated from female by zone of
in between; stamens free, anthers sessile, thecae sterile flowers, appendix rarely reduced or absent,
subopposite, dehiscing by apical pore; pistils few, smooth to staminodial; male flowers 1-6-androus,
slender, ovary 1-locular, ovule 1, anatropous, pla- filaments sometimes connate, connective rarely
centa basal; embryo elongate, endosperm copious. prominulent (A. longiconnectivus Bogner), thecae
dehiscing apically by pore or transverse slit;
gynoecia sometimes± remote; ovary 1-4-locular,
78. Callopsis Engler locules 1-ovulate, placenta axile to basal; berry
Callopsis Engler, Notizbl. Konigl. Bot. Gart. Berlin 1:27 (1895); orange to red, rarely blue or white. 2n = 26, 28,
Mayo, Fl. Trop. East Africa, Araceae: 54-56 (1985). 39. About 150 spp., tropical Africa and Madagas-
car, S and SE Asia to Melanesia, the Pacific, and
Leaves 2 per article; petiole sheath very short; N Australia, in humid to dry forests and savannas.
inflorescences 2, appearing with leaves; peduncle A. paeoniifolius (Dennstedt) Nicolson is an im-
slender, equalling or exceeding petiole; spathe portant food plant in the Old World tropics,
ovate-elliptic to subcircular, white; spadix: especially in India where it is widely cultivated;
male zone cylindric; gynoecium flask-shaped, A. konjac C. Koch is cultivated in southern Japan
falcate; berry ellipsoid, green; testa ± smooth, as a source of food gels. About 14 sections or
thin. 2n = 36. Only one sp., C. volkensii Engler, E informal sectional groups are recognised, but
Africa (Kenya, Tanzania); terrestrial. Systematic many may be unnatural (W. Hetterscheid, pers.
position problematic; C. shows similarities to comm.).
 Araceae 65

80. Pseudodracontium N.E. Brown Leaf 1(-2); leaf blade cordate; inflorescences
1-2( -3), appearing with or before leaf; spathe not
Pseudodracontium N.E. Brown, J. Bot. 20: 193 (1882);
Gagnepain, Fl. gen. Indo-Chine 6: 1152-1157 (1942); constricted; spadix: lower part of female zone
Serebryanyi, Blumea 40: 217-235 (1995), rev. adnate to spathe, contiguous with male or with
few bisexual flowers in between, sometimes (C.
Tuberous; leaves often more than 1; inflorescence glaucophyllum Bogner) flowers of "female" zone
often appearing with leaf; peduncle long; spathe almost entirely bisexual; synandrium 2-6( -1 )-
boat-shaped, convolute basally; spadix shorter androus, elongate, stamens either completely con-
than spathe, male zone laxly flowered, appendix nate with thecae marginal, inverted and dehiscing
composed of clavate, irregularly connate on lower surface, or incompletely connate with
staminodes, separated from male zone by naked filament apices ± free and thecae dehiscing
axis; male flowers 3-6-androus, filaments free or apically, sometimes remote; berry orange-red.
± connate; ovary 1-locular, placenta basal; berry 2n = 54, 108. Three spp., NW Madagascar, on
ellipsoid. 2n 26; Seven spp., SE Asia limestone in crevices or holes with leaf litter.
(Indochina), in moist forest.
83. Colletogyne Buchet
19. Tribe Arophyteae Bogner (1972). Colletogyne Buchet, Bull. Soc. Bot. Fr. 86: 23 (1939); Bogner,
Seasonally dormant; stem tuberous (except most Bot. Jahrb. 92: 21-24 (1972).
Arophyton spp.); primary lateral veins pinnate,
forming 1 submarginal and 1-2 marginal veins, Leaf 1(-2); leaf blade cordate; inflorescences 1-3,
finer venation reticulate; spadix fertile to apex appearing before or with leaf; peduncle longer
(except some Arophyton spp.); male flower a than spathe; spathe obovate, unconstricted, fully
synandrium of partially or completely connate expanded at anthesis, later closing, persistent;
stamens (sometimes reduced to 1 stamen); gyno- spadix: female zone entirely adnate to spathe, con-
ecium surrounded by urceolate synandrodium, tiguous with male zone; synandrium reduced to 1
ovary 1-locular, ovule 1, atropous, placenta basal; stamen; berry fusiform. 2n = 54. Only one sp., C.
testa thin, smooth, embryo large, endosperm perrieri Buchet, NW Madagascar, on limestone, in
absent. holes or crevices with leaf litter.

81. Arophyton Jumelle 20. Tribe Peltandreae Engler (1876).

Arophyton Jumelle, Ann. Mus. Colon. Marseille 36 annee, IV, Rhizomatous; primary lateral veins pinnate, form-
6: 23 (1928); Bogner, Bot. Jahrb. 92: 24-45 (1972), rev. ing submarginal collective vein, 1-2 marginal
Synandrogyne S. Buchet (1939). veins also present, finer veins ± parallel-pinnate;
Humbertina S. Buchet (1941). inflorescences usually 2, infructescence pendent;
spathe ± constricted centrally, tube ellipsoid to
Stem usually rhizomatous, rarely tuberous; leaves oblong, persistent, blade gaping at anthesis, later
1-few; leaf blade cordate, hastate, tripartite or marcescent; male flower a truncate, prismatic
pedatisect; inflorescences 2-3; spathe not or synandrium of connate stamens, thecae lateral,
slightly constricted; female zone of spadix free dehiscing by apical pore or short slit; ovary 1-
or partly adnate to spathe, contiguous with male locular, ovules atropous; berry and seeds large,
or separated by zone of sterile or bisexual flowers, testa smooth, embryo large, plumule well devel-
terminal appendix sometimes present; synan- oped, endosperm vestigial to absent.
drium 2-7-androus, thecae marginal, subglobose,
dehiscing on upper surface; berry red or green;
seed ellipsoid to globular. 2n = 38, 54, 76 (40). 84. Peltandra Raf.
Seven spp., Madagascar, on forest floor in ever-
Peltandra Raf., J. Phys. Chim. Hist. Nat. Arts 89: 103 (1819),
green and deciduous forest, epiphytic or in lime- nom. cons.; Huttleston, Temp. N. Am. Arac.: 65-86 (1953);
stone crevices. Blackwell & Blackwell, J. Elisha Mitchell Sci. Soc. 90: 137-
140 (1975).

82. Carlephyton Jumelle Rhizome short, erect; petiole and sheath relatively
Carlephyton Jumelle, Ann. Mus. Colon. Marseille 27 annee, III, long; leaf blade sagittate-hastate; peduncle
7: 187 (1919); Bogner, Bot. Jahrb. Syst. 92: 10-21 (1972). subequalling petiole; spadix: female and male
66 Araceae

zones contiguous or with zone of sterile flowers margins connate, blade fornicate, gaping; spadix:
in between, terminal appendix of sterile flowers female zone adnate to spathe, very short, contigu-
sometimes present; synandrium 4-5-androus; gy- ous with male, male zone laxly flowered, appendix
noecium surrounded by cuplike synandrodium, slender to massive and knob like, hidden or shortly
ovules 1-few, atropous to hemiatropous, placenta exserted; berry flattened at apex, several-seeded;
parietal to basal; berry green or orange, 1-3- seed longitudinally rugose, strophiolate. 2n = 28,
seeded; seed ovoid. 2n = 56, 112. Three spp., tem- 42, 56. Three spp., Mediterranean, Macaronesia.
perate to subtropical eastern N America (Maine to
Florida), helophytes, seasonally dormant.
22. Tribe Ambrosineae Schott (1832).
Stem tuberous; leaves few; leaf blade ovate-
85. Typhonodorum Schott
elliptic, primary lateral veins arising basally, arcu-
Typhonodorum Schott, Oesterr. Bot. Wochenbl. 7: 70 (1857); ate, running into apex, finer veins reticulate;
Bogner, Fl. Madagascar 31: 22-26 (1975); Mayo, Fl. Trop. inflorescence 1, on recurved peduncle; spathe
East Africa, Araceae: 38-40 (1985). boat-shaped, unconstricted externally, basal tube
convolute, apical part narrowly gaping, rostrate,
Often gigantic, rhizome thick, aerial shoots form- internally divided into 2 chambers, an upper,
ing massive false stems; leaves forming terminal morphologically ventral one and a lower, morpho-
crown; petiole and sheath very long; leaf blade logically dorsal one; spadix shortly appendiculate,
subtriangular to sagittate; inflorescences 2-3, pe- adnate to spathe by longitudinal, septumlike lat-
duncle shorter than petiole; spadix with 4 zones eral dilations, ventral chamber containing female
(base to apex): female zone, narrower zone of ster- zone consisting of only 1 gynoecium, dorsal cham-
ile male flowers, subcylindric-conoid male zone, ber containing 8-10 biseriate male flowers; male
terminal appendix of sterile male flowers; flower a 2-androus, sessile synandrium, thecae
synandria 4-6-androus; ovule 1( -2), atropous, confluent, dehiscing by single continuous slit;
placenta basal, stigma broad, shallowly 3-6-lobed; female flower 1, ovary 1-locular, ovules many,
berry very large, yellow; seed compressed- atropous, placenta basal, style long, curved,
obovoid. 2n = 112. Only one sp., T. lindleyanum, stigma discoid; berry many-seeded; seed ellipsoid,
Madagascar, Comores, Pemba, Zanzibar, testa costate, with large white conical strophiole,
Mauritius (?naturalized), a helophyte in fresh- endosperm copious, starch abundant.
water swamps, also in brackish water, forming
huge stands.
87. Ambrosina Bassi
21. Tribe Arisareae Dumortier (1827). Ambrosina Bassi, Ambrosina Nov. Plant Gen.: 3 (1763) &
Bononiensi Sci. Inst. Acad. Comment. 5: 82-86 (1766);
Laticifers simple; stem a tuber or rhizome; leaves Killian, Bull. Soc. Hist. Nat. Afr. N. 20: 257-278 (1929) & 24:
1-3; primary lateral veins of leaf blade forming 259-294 (1933).
submarginal collective vein, finer venation reticu-
late; inflorescence 1, appearing with leaves; spathe Very small, seasonally dormant; inflorescence
not constricted, tube margins connate, blade for- borne at ground level; peduncle relatively long;
nicate, gaping; spadix with usually smooth, sterile, spathe ± horizontal; berry depressed-globose.
terminal appendix; flowers unisexual, perigone 2n = 22. Only one sp., A. bassii L., western
absent; male flower 1-androus, thecae apically Mediterranean.
confluent, dehiscing by 1 continuous slit; female
flowers 2-5, ovary 1-locular, ovules several,
23. Tribe Areae
atropous, placenta basal; endosperm copious,
starch abundant. Seasonally dormant geophytes; stem usually
tuberous; leaves several (often 1 in Sauromatum);
primary lateral veins of leaf blade or lobes or
86. Arisarum P. Miller leaflets forming submarginal collective veins,
Arisarum P. Miller, Gard. Diet. abr. ed., [121] (1754); Boyce, finer venation reticulate; inflorescence usually 1,
Kew Mag. 7: 14-21 (1990). usually appearing with or after leaves (except
Sauromatum and Biarum); spathe usually
Seasonally dormant; leaf blade cordate to sub- strongly constricted, with convolute (connate in
hastate; spathe: tube ± cylindric or subventricose, Sauromatum, some Typhonium spp. and Biarum)
 Araceae 67

tube and gaping, erect to reflexed blade; spadix:

terminal sterile appendix always present, usually
smooth; stamens usually free; ovary 1-locular,
ovules atropous; berry subglobose; seed usually
with rather large, fleshy strophiole (aril), testa
rugose-reticulate (except Sauromatum and some
Biarum spp.), embryo ± small, endosperm
copious, starch abundant.

88. Arum L. Fig. 17F-I

Arum L., Sp. Pl.: 964 (1753); Boyce, The genus A rum: 1-196
(1993) (rev.).

Tubers elongated in some species; leaf blade sagit-

tate to hastate; peduncle short to long; spadix: fe-
male and male zones usually separated by zone of
filamentous pistillodes, rarely contiguous or inter-
mediate zone naked, male zone rather short,
usually separated from appendix by zone of fila-
mentous staminodes, appendix often stipitate;
male flowers 3-4-androus, anthers subsessile;
ovules 6 or more, placenta parietal-basal, stigma
exuding nectar droplet at anthesis; berry many-
seeded, red, pericarp juicy. 2n = 28, 42, 56, 70, 84.
Twenty six spp., temperate Europe, Macaronesia,
Mediterranean, Middle East to Central Asia, in
forests, orchards, open ground, rock crevices.
Boyce (1993) recognised two subgenera.

89. Eminium (Blume) Schott Fig. 17A-E

Eminium (Blume) Schott, Aroideae: 16 (1855, "1853"); Riedl,
Fl. Iranica, Araceae: 5- 7 (1963) & Ann. Naturh. Mus. Wien
73: 103-121 (1969) & Fl. Iraq 8: 187-203 (1985); Lobin &
Boyce, Willdenowia 20: 43-51 (1991).

Leaf blade thick, hastate to pedatifid or pedatisect,

rarely oblong-elliptic, posterior lobes of pedatifid
and pedatisect leaves ± erect and spirally twisted;
spadix: female zone separated from male by zone
of subulate sterile flowers; male flowers 2-
androus, anthers sessile; ovules 2, placenta basal Fig. 17A-1. Araceae-Aroideae. A-E Eminium alberti. A Habit.
to subbasal; infructescence subglobose, borne at B Spadix. C Stamen. D Pistil. E Same, longitudinal section. F-
or below soil level, berry white to pale lilac, peri- 1 Arum korolkowii. F Habit. G Spadix. H Stamen. I Pistil,
carp firm, not juicy. 2n = 28. Eight spp., Turkey, longitudinal section. (Takhtajan 1982)
Middle East, Central Asia.

pedatifid to pedatisect; peduncle longer than peti-

90. Dracunculus P. Miller ole; spadix: female zone contiguous with male,
Dracunculus P. Miller, Gard. Diet., Abr. ed., [455] (1754); sterile flowers absent, appendix very long, long-
Boyce, Thaiszia 4: 175-182 (1994). stipitate, fistulose; male flower 3-4-androus, fila-
ments connate basally; pollen extruded in strands;
Large; tuber stoloniferous in D. canariensis; peti- ovules few, placentas 2, apical and basal; berry
ole sheath very long, forming false stem; leaf blade orange-yellow to red, few-seeded. 2n = 28. Two
68 Araceae

spp., Mediterranean, Macaronesia, macchie, olive florescence 1 or more; spathe tube sometimes
groves, woodland. with connate margins, rarely septate internally
forming two chambers; spadix: female zone
91. Helicodiceros Schott ex K. Koch separated from male by sterile zone covered en-
tirely or only basally with sterile flowers, appendix
Helicodiceros Schott ex K. Koch, Index Sem. Hort. Berol. 1855, usually stipitate, rarely with spoonlike insertion
Appendix: 2 (1856), nom. cons.; Arrigoni et a!., Le piante (T. cochleare A. Hay), conic to filiform-subulate;
endemiche della Sardegna: 416-420 (1982); Boyce, Thaiszia
4: 175-182 (1994), rev.
male flower 1-3-androus; ovules 1-2, placenta
basal; infructescence subglobose to subcylindric,
Leaf blade pedatifid to pedatisect, posterior lobes borne above soil level, berry dark red to purple.
2n = 16, 18, 20, 26, 36, 52, 54, 65, > 100 (14). About
± erect and spirally twisted; peduncle short;
spathe held obliquely, deflexed at constriction; 40 spp., S and SE Asia to N and E Australia, in
spadix: female zone separated from male by zone humid and seasonal forests, some species weedy
of long, curved, subulate sterile flowers, appendix and naturalised outside Asia in tropical, sub-
stipitate, long, lying on spathe blade, covered with tropical and temperate regions; usually seasonally
curved sterile flowers varying from large and dormant, sometimes evergreen; two sections
subulate at the appendix base to setiform at the recognised.
apex; male flower 2-3-androus, anthers sessile;
ovules ca. 6, placentas 2, apical one with 3-4 94. Sauromatum Schott
ovules and basal one with 1-2 ovules; berry
Sauromatum Schott in Schott & Endlicher, Melet. Bot.: 17
orange-coloured, (1- )2-3-seeded. 2n = 56. Only (1832); Mayo, Fl. Trop. East Africa, Araceae: 56-60 (1985).
one sp., H. muscivorus (L.) Engler, western ]aimenostia Guinea & Gomez Moreno (1946).
Mediterranean, on stony ground, in rock crevices.
Leaves 1-4; leafblade pedatifid or pedatisect; inflo-
92. Theriophonum Blume rescence appearing before leaf; spathe only slightly
constricted, tube with connate margins; spadix:
Theriophonum Blume, Rumphia 1: 127 (1837, "1835"); female zone separated from male by rather long
Sivadasan & Nicolson, Kew Bull. 37: 277-290 (1982), rev. sterile zone bearing remote, patent, filiform to
Pauella Ramamurthy & Sebastine (1967).
clavate staminodes at base and naked above, ap-
pendix rather slender; male flower few-androus;
Small; leaf blade ovate, sagittate to hastate, some-
times linear-lanceolate; peduncle usually short, ovules 1-4, placenta basal; infructescence sub-
globose, partly hypogeal, berry purplish red,
deflexing in fruit; spathe tube marcescent; spadix:
darker at apex; testa smooth. 2n = 26, 52, 104. Two
female zone few-flowered, separated from male by
rather long sterile zone partly or completely cov- spp., tropical Africa, Saudi Arabia, Yemen, S Asia,
Myanmar, SW China, in upland forest and grass-
ered by filiform to subulate sterile flowers situated
land, occasionally epiphytic.
either basally, centrally or apically, male zone
cylindric, sometimes with zone of sterile flowers
above, appendix stipitate or not, cylindric to long- 95. Biarum Schott
subulate; male flower 1-2-androus; ovules 3-9 (or Biarum Schott in Schott & Endlicher, Melet. Bot.: 17 (1832),
more), placentas 2, apical and basal; berry few- nom. cons.; Bogner & Boyce, Willdenowia 18: 409-417
seeded. 2n = 16 (14, 18). Five spp., India, Sri (1989).
Lanka, on forest floor, grassy places, damp sites.
Petiole sometimes hardly distinct from blade; leaf
93. Typhonium Schott blade linear to obovate or elliptic-oblong; inflores-
cence usually appearing before leaves, sometimes
Typhonium Schott, Wiener Z. Kunst 1829 (3): 732 (1829); borne partly below ground level; spathe tube mar-
Nicolson & Sivadasan, Blumea 27: 483-497 (1981); Murata, gins partially to almost completely connate, rarely
Am. J. Bot. 77: 1475-1481 (1990); Hay, Blumea 37: 345-376
(1993); Sriboonma et al., J. Fac. Sci. Univ. Tokyo, sect. III, free (B. aleppicum J. Thiebaut); spadix: female
15: 255-313 (1994), rev. zone subhemispherical, separated from male by
Lazarum A. Hay (1992). sterile zone bearing filiform to conic, suberect to
decurved sterile flowers at base only or through-
Sometimes minute; tuber, rarely stoloniferous or out (rarely naked), male zone sometimes sepa-
creeping rhizome; leaf blade cordate, sagittate, rated from appendix by short zone of subulate
hastate, trifid, trisect, pedatifid or pedatisect; in- sterile flowers, terminal appendix slender, elon-
 Araceae 69

gate to conoid, sometimes stipitate; male flowers

1-2-androus; ovule 1, placenta basal; infructe-
scence subglobose borne at or slightly below soil
level, berry white to pale lilac, pericarp firm, not
juicy; testa rugose-reticulate, rarely smooth. 2n =
16, 20, 22, 24, 26, 32, 36, 74, 96, 98. Twenty two
spp., Mediterranean, Middle East, on stony
ground, under trees and in rock crevices.

24. Tribe Arisaemateae Nakai (1943).

Laticifers simple; tuberous, rarely rhizomatous;
leaf blade usually compound: trisect, pedatisect or
radiatisect, rarely cordate or trifid; spathe tube
usually cylindric, blade usually fornicate; spadix
with terminal appendix, appendix often cylin-
dric, clavate or elongate-tapering; flowers uni-
sexual, perigone absent; ovary 1-locular, ovules
atropous, placenta basal; testa rough, endosperm
copious.

96. Pinellia Tenore

Pinellia Tenore, Atti Acad. Sci. Napoli 4: 69 (1839), nom. cons.;
Li Heng, Fl. Reip. Pop. Sin. 13:200-206 (1979); Ulharz, Beitr.
Bioi. Pflanz. 60: 277-291 (1986) & ibid. 61: 241-282 (1986);
Boyce, Kew Mag. 5: 18-21 (1988).

Petiole sometimes bearing tubercles; leaf blade

usually trisect or pedatisect, rarely entire and
ovate-oblong to cordate; peduncle equal or longer
than petiole; spathe weakly constricted, tube nar-
row, persistent, almost closed at mouth by trans-
verse septum, blade ± fornicate; spadix: female
zone adnate to spathe, separated from male by
spathe septum and short, naked zone, appendix
smooth, subulate, slender, long-exserted; male
flower 1-2-androus, stamens free or sometimes
irregularly connate; ovule 1, ± atropous; testa
rugulose. 2n ± 26, 52. Six spp., temperate E Asia,
P. ternata often naturalised elsewhere, in forests
or on rocks, also weedy in fields.
Fig. 18A-M. Araceae-Aroideae. A-E Arisaema amurense. A
Habit. B Staminate spadix. C Synandrium. D Pistillate spadix.
E Pistil. F, G A. thunbergii. F Inflorescence with long exserted
97. Arisaema Mart. Fig. 18 appendix. G Infructescence. H-N A. griffithii. H Habit. I Inflo-
Arisaema Mart., Flora 14: 459 (1831); Huttleston, Temp. N. rescence with flexuose appendix. J Staminate spadix. K
Am. Arac.: 87-154 (1953) (N Amer. spp.); Li Heng, Fl. Reip. Synandrium. L Pistillate spadix. M Pistil, longitudinal seciton.
Pop. Sinicae 13: 116- 194 (1979); Ohashi & Murata, J. Fac. (Takhtajan 1982)
Sci. Univ. Tokyo, Bot. 12: 281-336 (1980); Murata, J. Fac.
Sci. Univ. Tokyo, Bot. 13: 431-482 (1984); Mayo & Gilbert,
Kew Bull. 41: 261-278 (1986) (African spp.); Murata, Pl. Sometimes large; tuber depressed-globose, rarely
Sp. Bioi. 2: 57-66 (1987) & Bot. Mag. Tokyo 103: 371-382 rhizomatous, sometimes stoloniferous; petioles
(1990).
Flagellarisaema Nakai (1950). usually long, often forming false stem; leaf blade
Heteroarisaema Nakai (1950). trisect, pedatisect or radiatisect, rarely cordate or
Pleuriarum Nakai (1950). trifid; peduncle short to long; spathe tube usually
Ringentiarum Nakai (1950). cylindric, mouth often flaring with revolute to
70 Araceae

grotesquely auriculate margins, blade usually for- 99. Protarum Engler

nicate, often with elongate or filamentous apex;
Protarum Engler, Bot. Jahrb. 30, Beibl. 67: 42 (1901); Bogner,
spadix unisexual (paradioecious) or monoecious, Palmengarten 37: 40 (1973); Robertson, Fl. Pl. Seychelles:
male zone usually laxly flowered, contiguous with 259 (1989).
female, sometimes separated from appendix by
short zone of filamentous or subulate staminodes, Tuber globose; leaf 1; leaf blade subpedatisect to
appendix hidden or exserted, rarely absent (A. almost radiatisect, leaflets shortly stalked, lan-
exappendiculatum H. Hara), often cylindric, clav- ceolate, acuminate to acute; inflorescence 1; pe-
ate or elongate-tapering, sometimes drawn out duncle short; spadix shorter than spathe, female
into an extremely long flexuose thread, rarely zone conoid, male zone obconic, female and male
composed of several long threads; male flowers zones separated by slender sterile zone, terminal
remote, 2-5-androus, stamens connate, syn- appendix thick, obtuse; synandrium 3-androus, ±
andrium ± sessile to long-stipitate; ovules 1-9 per 6-lobed; female flower with whorl of 4-6 free
ovary; berry red, few-seeded; testa rough. 2n = 20, staminodes surrounding gynoecium, ovary 1-
22, 24, 26, 28, 39, 42, 48, 52, 56, 72, 112, 140, 168 locular, ovules 4, hemiatropous, placenta basal;
(64). About 170 spp., eastern and southeastern berry 1-seeded; seed ellipsoid, testa costate, em-
USA, northern Mexico, E and NE Africa, Arabian bryo discoid, endosperm copious. 2n = 28. Only
Peninsula, temperate E Asia, tropical S, SE and one sp., P. sechellarum, Seychelles Is., seasonally
E Asia, Indonesia, in the tropics only at high dormant.
altitudes. Murata (1990) recognized 11 sections.

100. Steudnera K. Koch

25. Tribe Colocasieae Engler (1876).
Steudnera K. Koch, Wochenschr. Giirtnerei Pflanzenk. 5: 114
Laticifers anastomosing (except Ariopsis); pri- (1862).
mary lateral veins of leaf blade or leaflets
(Protarum) pinnate, forming submarginal collec- Sometimes robust; stem epigeal, stout, erect or
tor vein, at least 1-2 other marginal veins also sometimes creeping; leaves 1-several; leaf blade
present; spathe usually constricted centrally (ex- peltate, ovate-subcordate; inflorescences 1-2;
cept Ariopsis, sometimes constricted twice in spa-the ovate or ovate-lanceolate, not constricted,
Remusatia) with lower part forming a convolute, barely convolute at base, ± fully expanded at an-
persistent tube and upper part a gaping, ± boat- thesis, upper part later reflexed; spadix: female
shaped, marcescent to caducous blade; flowers zone mostly adnate to spathe, contiguous with and
unisexual, perigone absent; male flower a synan- longer than male zone, terminal appendix absent;
drium of several connate stamens; endosperm synandrium 3-6-androus; gynoecium usually sur-
copious, with abundant starch. rounded by 2-5 short staminodes, ovary !-locular,
ovules many, hemiatropous, placentas 2-5, pari-
98. Ariopsis Nimmo etal; berry many-seeded; testa costate. 2n = 28, 56.
Eight spp., tropical S to SE Asia (Assam to
Ariopsis Nimmo in Graham, Cat. Pl. Bombay: 252 (1839). Vietnam), on forest floor, sometimes seasonally
dormant.
Laticifers simple; plant small; stem a partially hy-
pogeal tuber; leaves !-several; leaf blade peltate,
ovate-cordate to -emarginate; inflorescences 1-3, 101. Remusatia Schott
appearing either before or with the leaves; pe-
duncle slender, longer than spathe; spathe ovate, Remusatia Schott in Schott & Endlicher, Melet. Bot.: 18 (1832);
Mayo, Fl. Trop. East Africa, Araceae: 40-42 (1985); Li Heng,
boat-shaped, fornicate, not constricted, persis- Aroideana 10: 23-26 (1988); Li Heng & Hay, Acta Bot.
tent; spadix: female zone adnate to spathe, few- Yunnanica, Suppl. 5: 27-33 (1992).
flowered, separated from male by short naked Gonatanthus Klotzsch (1841).
axis, terminal appendix absent; synandria con-
nate, peltate, stipitate, 6-thecate; ovary 1-locular, Stem tuberous, subglobose, producing shoots
ovules many, atropous, placentas 4-6, parietal; bearing small cataphylls and axillary, clustered
berry many-seeded; testa costate, endosperm bulbils covered by recurved scales; leaves 1-2, leaf
copious. 2n = 28, 84. Two spp., tropical S Asia blade peltate, lanceolate to ovate-cordate; inflo-
(Myanmar to Western Ghats); in evergreen forest, rescence 1, appearing before leaf; peduncle short;
seasonally dormant. spathe with 1 or 2 constrictions, blade broad or
 Araceae 71

narrow and very long, reflexed at anthesis or ovate-cordate to sagittate, sometimes peltate, mar-
erect-sigmoid; spadix: female zone separated from gin rarely sinuate to deeply pinnatifid; inflores-
male by much narrower zone of sterile synand- cences several; spathe constricted, blade usually
rodes, terminal appendix absent; synandrium 2- boat -shaped and erect, rarely fornicate, sometimes
3-androus; pollen extruded in strands; female reflexed, rareJy persistent; spadix: female zone
flowers lacking staminodes, ovary 1-locular or separated from male by much narrower zone of
partially 2-3-locular apically, ovules several to sterile synandrodes, terminal appendix always
many, hemiatropous, placentas 2-4, parietal or present; synandrium 3-8-androus; pollen ex-
basal or both (R. yunnanensis H. Li & A. Hay); truded in strands; female flowers lacking stami-
berry many-seeded; testa thick, succulent, ver- nodes, ovary 1-locular or partially 3-4-locular at
ruculose to irregularly furrowed. 2n = 28, 42, 56. apex, ovules hemiatropous or anatropous, few,
Four spp., tropical Africa, Madagascar, S and SE placenta basal, style usually distinct, narrower
Asia, Indonesia, N Australia, in seasonal or ever- than ovary; berry 1-few-seeded, usually reddish;
green forest, terrestrial, rupicolous or epiphytic, testa smooth. 2n = 28, 42, 56, 70, 84. About 70 spp.,
seasonally dormant. S and SE Asia to northern Australia, Melanesia,
Pacific [A. macrorrhizos (L.) Schott widely
naturalised throughout the tropics], seasonally
102. Colocasia Schott Fig. 15A dormant to evergreen. Two sections recognised by
Colocasia Schott in Schott & Endlicher, Melet. Bot.: 18 (1832), Krause (in Engler and Krause 1920); Xenophya is
nom. cons.; Plucknett in Wang, Taro: 14-19 (1983); Shaw in reduced here following Hay and Wise (1991) but
Chandra, Edible Aroids: 160-168 (1984). merits recognition as a separate section.

Sometimes gigantic and arborescent; stem either

hypogeal and tuberous or mostly epigeal and mas- 26. Tribe Pistieae Blume (1836).
sive; leaves several; leaf blade peltate, ovate-
cordate; inflorescences usually many; peduncle Laticifers absent; acaulescent; leaves rosulate,
much shorter than petiole; spathe constricted; spa- densely pubescent; petiole very short; leaf blade
dix: female zone separated from male by much obovate-cuneate to oblong, midrib absent; inflo-
narrower zone of sterile synandrodes, terminal rescence 1, very small; spathe ± constricted, lower
appendix erect, ± smooth, sometimes reduced margins connate forming tube, free margins be-
to tiny stub; synandrium 3-6-androus; pollen tween tube and blade folded to form a partition
extruded in strands; female flowers lacking separating upper male chamber from lower fe-
staminodes, ovary 1-locular, ovules many, hemi- male one, blade gaping; spadix: female zone with a
atropous, placentas 2-4, parietal; berry greenish, single basal gynoecium and thin pouch-shaped
many-seeded; testa thickish, costate. 2n = 28, 42. flap below spathe partition, male zone with 2-8
Eight spp., NE India, SE Asia, Indonesia, seasonally flowers in a whorl, appendix naked, very short;
dormant or evergreen. C. esculenta (L.) Schott is synandrium sessile, of 2 connate stamens; ovary
cultivated and naturalised throughout the tropics obliquely adnate to spadix axis, 1-locular, ovules
with hundreds of cultivars; it is the world's most numerous, atropous, placenta parietal (morpho-
economically important aroid - for details see re- logically basal), style distinct, curved; berry
cent reviews by Wang (1983) and Chandra (1984) thin-walled, several-seeded, pericarp eventually
cited above. Two sections are recognised. decaying; seed cylindric, testa thick, reticulate-
alveolate, with micropylar operculum formed by
both integuments, embryo minute, endosperm
103. Alocasia (Schott) G. Don copious.
Alocasia (Schott) G. Don in Sweet, Hart. Brit., ed. 3:
631 (1839), nom. cons.; Burnett, Aroideana 7: 67-162
(1984); Hay, Aroids of Papua New Guinea (1990); Hay & 104. Pistia L.
Wise, Blumea 35: 499-545 (1991) (reg. rev. New Guinea and
Austral.) Pistia L., Sp. Pl.: 963 (1753); Buzgo, Bot. Jahrb. Syst. 115:557-
Xenophya Schott (1863); Nicolson, Blumea 16: 115-118 (1968). 570 (1994).
Schizocasia Schott ex Engler (1880).
Panzhuyuia Z.Y. Zhu (1985). Plant small, stoloniferous; roots much-branched,
feathery; leaf blade somewhat spongy; peduncle
Sometimes gigantic and arborescent; stem hypo- very short. 2n = 28. Only one sp., P. stratiotes L.,
geal to epigeal and erect; leaves several; leaf blade pantropical, free-floating evergreen aquatic.
72 Araceae

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single species in W Africa.

CHARACTERS OccuRRING IN FEw GENERA AND

SPECIES. Rheophytic plants in Pitcairnia and
Guzmania; climbing ones in Pitcairnia; distichous
phyllotaxis in Tillandsia, mainly in subgenus
Diaphoranthema; petiolate leaves in Pitcairnia,
Ronnbergia, Bromelia, and Cryptanthus, cuticular

1 Deceased.