Croat T. B., 1990, A comparison of aroid classification systems, Aroideana 13(1-4)

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44 AROIDEANA Vol. 13, No. 1-4
A Comparison of Aroid ClassiftcationSystems
Thomas B. Croat
Missouri Botanical Garden
p. o. Box 299
St. Louis, MO 63166-0299
USA
Abstract Hooker (1883) which in turn was based
on the first monograph of the family by
The paper compares four systems of
H. Schott (1860).
classification of the Araceae: Engler's
Like the Schott system, Hutchinson
original (1905-1920), M. Hotta's (1970), J.
based his classification primarily on floral
Bogner and D. Nicolson's On press) and
morphology. However, he divided the
M. Grayum's (1990). All are compared
genera not into subfamilies but into 18
against the backdrop of the traditional
tribes. Although Hutchinson's system has
system of classification by Adolf Engler
been used by some workers in general
and against each other.
review papers (e.g., Marchant, 1970,
1971a, 1971b, 1972, 1974; Raven and
Introduction
Axelrod, 1974; Li, 1979, 1980), it has been
A review of the systems of classifica deemed quite unnatural by modern
tion in the Araceae between the time of workers of the Araceae and it will not be
Linnaeus and the modern era was pre dealt with further.
sented by Nicolson (1960,1987). The last Since no other work has been so
thorough systematic treatment of the widely accepted, it is important that a
Araceae was published by Adolf Engler synopsis of Engler's original classifica
in Das Pjlanzenreich (Engler, 1905, tion be presented here. The modified
1911, 1912, 1915, 1920a, 1920b; Engler & Englerian system which follows differs
Krause, 1908, 1920; Krause 1908, 1913). from the original version only by the
This subfamilial classification has served inclusion of 14 accepted new genera
as the basis for virtually all non-taxo published since the appearance of
nomic studies of aroid morphology to Engler's revision. Originally, Engler had
this date. Although major subfamilial included 107 genera arranged in eight
revisions of the Araceae have been subfamilies. These retain their original
published (Hotta, 1970 and Hutchinson, numbers in the modification presented
1973), they have not gained wide accep while those added later into the system
tance. The system by Hotta, based pri are assigned lower case letters, as in 54a,
marily on the Englerian model, is proba Amauriella. Genera added since Engler
bly the more natural of the two. A are assigned upper case letters and also
synopsis of the Hotta system will be bear an asterisk. With minor exceptions
presented and discussed later. The only (see placement of Heteroaridarnm, Hot
other important treatment of the Araceae tarnm and jasarnm, more recently
in this century was that of Lemee (1941) switched by Bogner [pers. comm.]),
in his Dictionaire ....p hanerogams. This placement of new taxa within Engler's
work, published in French, is merely a system is based on their assignment in "A
version of Engler's treatment which in Critical List of Aroid Genera" (Bogner,
cluded four genera published since 1978). Three published genera have
Engler's treatment was published. In been added subsequent to the appear
contrast, Hutchinson's system is an ex ance of Bogner's list: Furtadoa (Hotta,
tension of the one devised by J. D. 1981) and Bognera (Nicolson, 1984),
THOMAS B. CROAT, 1990 45
both in the subfamily Philodendroideae dospatha [Croat], 18. Monstera
(Calloideae in Grayum, 1990) and An Adans., 19. Alloschemone
aphyllopsis Hay in the Lasioideae. Schott, 20. Amydrium Schott)
Lasiomorpha was resurrected to the ge Tribe 2. SPATHIPHYLLEAE Engl. (21.
neric level by Hay. These have been Spathiphyllum Schott, 22. Hol
included, using the same lettering system ochlamys Engl.)
as for genera previously added. Subfamily m. CALLOIDEAE Schott
Genera accepted by Engler but subse Tribe 1. SYMPLOCARPEAE Engl. (Ap
quently placed into synonymy are also propriately now ORONTIEAE)
indicated. The author who placed it into (23. Lysichiton Schott (as
synonymy is added in brackets. Some of Lyschitum) 24. Symplocarpus
the authors of Engler's generic names Salisb., 25. Orontium L.)
have been changed to reflect proper Tribe 2. CALLEAE Schott (26. Calla L.)
nomenclature.
Subfamily Iv. LASIOIDEAE Engl.
Tribe 1. LASIEAE Engl. (27. Cyrto
The SubfamiJial Classification of the sperma Griff., *27A. Lasiomor
Araceae by Engler (1905-1920) pha Schott [Hay] (989), 28.
Lasia Lour. 29. Anaphyllum
Subfamily I. POTHOIDEAE Engl.
Schott, 29A. Anaphyllopsis Hay,
(Properly ACOROIDEAE, according to
30. Podolasia N. E. Brown, 31.
the rules of nomenclature when Acarus
Urospatha Schott [Bogner],
is included.)
(1988, 1989) 32. Dracontioides
Tribe 1. POTHEAE Engl. (1. Pothos L., 2 ..
Pothoidium Schott, *2A. Engl., 33. Echidnium Schott =
Dracontium [Bogner], 34. Dra
Pedicellarum M. Hotta, 3.
Anadendrum Schott (as Anad contium L., *34A. Pycnospatha
endron), 3a. Epipremnopsis Thorel ex Gagnep.)
Engl. = Amydrium [Nicolson]) Tribe 2. AMORPHOPHALLEAE Eng!.
(Now correctly THOM
Tribe 2. HETEROPSIDEAE Engl, (4.
Heteropsis Kunth) SONIEAE) 05. Pseudohy
Tribe 3. ANTHURIEAE Engl. (5. An drosme Engl., 36. Plesmonium
thurium Schott) Schott = A morphophallus
Tribe 4. CULCASlEAE Engl. (6. Culcasia [Bogner], 37. Anchomanes
P. Beauv.) Schott, 38. 1bomsonia Wall. =
Tribe 5. ZAMIOCULCADEAE Engl. (7. Amorphophallus [Bogner,
Zamioculcas Schott, 8. Gona Mayo & Sivadasan], 39. Pseu
topus Hook. f.) dodracontium N. E. Brown, 40.
Tribe 6. ACOREAE Engl. (9. Acorus L. = Amorphophallus Blume)
Acoraceae, 10. Gymnostachys Tribe 3. NEPHTHYTIDEAE Engl. (41.
R. Br.) Nephthytis Schott, 42. Cercestis
Subfamily II. MONSTEROIDEAE Schott, 43. Rhektophyllum N. E.
Engl. Brown = Cercestis [Bogner])
Tribe 1. MONSTEREAE Eng!. (ll. Tribe 4. MONTRICHARDIEAE Eng!.
Rhaphidophora Hassk. (as Ra (44. Montrichardia Cruger)
phidophora) Engl. 12. Afrora Subfamily V. PHll.ODENDROIDEAE
phidophora Engl. = Rhaphido Engl.
phora [Hepper], 13. Epi Tribe 1. PHILODENDREAE Schott
premnum Schott, 14. Scin SubTribe 1. HOMALOM
dapsus Schott, 15. Stenosper ENINAE Schott, (*45A. Furta
mation Schott (as Stenosperma doa M. Hotta 45. Homalomena
tium), 16. Rhodospatha Poepp., Schott, 46. Diandriella Engl. =
17. Anepsias Schott = Rho- Homalomena [Bogner])
AROIDEANA Vol. 13, No. 1-4
Fig. 1. Cercestis kamerunianus N.E. Br. Fig. 2. Mangonia uruguaya (Hicken)
in Dyer, Croat 53498. Nigeria. Photo by Bogner, F Felippone s.n. (type), Uru
T B. Croat. guay. Photo by F. Felippone.
Fig. 3. Calla palustris L., de GraCi! 508. Fig. 4. Chlorospatha croatiana Grayum,
Photo by A. de Graaf. Croat 67109, Panama. Photo by T B.
Croat.
maMAS B. CROAT, 1990 47
Subtribe 2. SCHISMATOG jasarum Bunting, 67. Apbyl
LOTTIDINAE Schott (47. Scbis larum S. Moore, = Caladium
matoglottis Zoll. & Mar., 48. [Bogner & Mayo) 68. Cblorospa
Bueepbalandra Schott, *48A. tba Eng!., 69. Xantbosoma
Pbymatarum M. Hotta, 49. Ari Schott)
darum Ridley, *49A. Heteroari Subtribe 4. COLOCASII
darum M. Hotta, *49B. Hotta NAE Schott (70. Coloeasia
rum Bogner & Nicolson, 50. Schott)
Piptospatba N. E. Brown, 51. Subtribe 5. ALOCASIINAE
Mieroeasia Beccari = Bueepb Schott (71. Aloeasia (Schott) G.
alandra [Bogner]) Don, 72. Sebizoeasia Engler =
Subtribe 3. PHILO DEN Xenopbya [Nicolson) = Alocasia
DRINAE Schott (52. Pbiloden [A. Hay])
dron Schott [Krause (913) rec
Tribe 2. SYNGONIEAE Eng!. (73. Por
ognized Tbaumatopbyllum
pbyrospatba Eng!. = Syngo
Schott, now = Pbilodendron nium [Croat), 74. Syngonium
[Bunting), without numbering it
Schott)
or putting it in a key), 53.
Tribe 3. ARIOPSIDEAE Eng!. (75. Ariop
Pbilonotion Schott = Sebisma sis Nimmo ex J. Graham)
tog lottis [Bunting])
Subfamily VII. AROIDEAE Engl.
Tribe 2. ANUBIADEAE Eng!. (54A.
Tribe 1. STYLOCHAETONIEAE Schott
Amauriella Rendle = Anubias (76. Styloebaeton l.epr. as Sty
[Bogner), 54B. Anubias Schott)
loebiton)
Tribe 2A. BOGNEREAE Mayo & Ni
Tribe 1A. AROPHYTEAE Bogner (*76A.
colson (*54A. Bognera Mayo
Carlepbyton]um., *76B. Colle
& Nicolson)
togyne S. Buchet *76c. Aro
Tribe 3. AGLAONEMATEAE Engl. (55.
pbyton ]um.)
Aglaonema Schott, 56.
Tribe 2. ASTEROSTIGMATEAE Schott
Aglaodorum Schott)
(77. Mangonia Schott, 78. An
Tribe 4. DIEFFENBACHIEAE Engl. (57.
Dieffenbaebia Schott) dromyeia A. Rich. = Aster
ostigma [Bogner), 79. Tae
Tribe 5. ZANTEDESCHIEAE Eng!. (58.
earum Brongn. ex Schott, 80.
Zantedesebia Spreng.)
Asterostigma Fisch. & Mey., 81.
Tribe 6. TYPHONODOREAE Eng!. (59.
Synandrospadix Eng!., 82 .
Typbonodorum Lind!.)
Tribe 7. PELTA NDREAE Eng!. (60. Pel Spatbantbeum Schott, 83.
tandra Raf.) Gorgonidium Schott, 84.
Subfamily VI. COLOCASIOIDEAE Gearum N. E. Brown, 85. Spatb
Engl. iearpa Hook.)
Tribe 1. COLOCASIEAE Eng!. Tribe 3. PROTAREAE Eng!. (86. Prota
Subtribe 1. STEUDNERINAE rum Eng!.)
Eng!. & K. Kr. (61. Steudnera Tribe 4. CALLOPSIDEAE Eng!. (87. Cal
K. Koch, 62. Remusatia Schott, lopsis Eng!.)
63. Gonatantbus Klotzsch Tribe 5. ZOMICARPEAE Eng!. (88.
Subtribe 2. HAPALININAE Seapbispatba Brongn. ex
Eng!. & K. Kr. (64. Hapaline Schott, 89. Xenopbya Schott =
Schott) Aloeasia [Hay), 90. Zomiearpa
Subtribe 3. CALADIINAE Schott, 91. Zomiearpella N. E.
Eng!. & K. Kr. (65. Caladiopsis Brown, *91A. Filarum Nicolson,
Eng!. = Cblorospatba [Madi 92. Ulearum Eng!.)
son), 66. Caladium Vent., *66A. Tribe 6. AREAE Engl.
48 AROIDEANA Vol. 13, No. 1-4
Subtribe 1. ARINAE Schott systems will be presented here in synop
(93. Arnm 1., 94. Dracunculus tic form. These, as well as the system of
Schott, 95. HelicodicerosSchott, Hotta, will be compared with Engler's
96. Tberiophonum Blume, 97. classification. All of these systems have
Typhonium Schott, 98. Sauro benefited from a substantial amount of
matum Schott, 99. Eminium information not available to Engler. This
(Blume) Schott, 100. Biarnm included extensive surveys of the anat
Schott) omy (Solereder & Meyer, 1928; Cheadle,
Subtribe 2. ARISARINAE 1942; Metcalfe, 1967), including the ex
Schott (101. Arisarnm Targ. tensive surveys of vascular stem patterns
Tozz.) by French and Tomlinson 0980, 1981a,
Subtribe 3. ARISAEMATI 1981b, 1981c, 1981d, 1983), and floral
NAE Engl. 002. Arisaema anatomy (Eyde et aI., 1967) as well as leaf
Mart.) blade nervature (Ertl, 1932), embryology
Subtribe 4. PlNELLIINAE Qiissen, 1928), and of seedling morphol
Schott 003. Pinellia Ten.) ogy (Tillich, 1985).
Subtribe 5. AMBRO- Recent investigations by J. c. French
SININAE Schott 004. Am
and his associates have provided surveys
brosina Bassi)
on patterns of anther endothecial wall
Subtribe 6. CRYPTOCO
thickenings (French, 1986a), ovular vas
RYNINAE Schott 005. Lage
culature (French, 1986b), stamen vascu
nandra Dalzell, 106. Crypto
lature (French, 1986c), structure of ovu
coryne Fisch. ex Wydl.)
lar and placental trichomes (French,
1987a), the occurrence of sclerotic hypo
Subfamily vm. PISTIOIDEAE Engl. dermis in roots (French, 1987b), the
007. Pistia 1.) occurrence of resin canals in roots
(French, 1987c), the presence of anasto
mosing laticifers (French, 1988), and the
Recent Revisions of the Subfamilial presence of latex particles (Fox &
Classification of the Araceae French,in prep. ). A single surveyor the
study of a single character rarely pro
It is important to give recognition to
vides conclusive evidence for the cor
the work of a small group of active
rectness of the placement of any member
researchers working in different parts of
the world. These include J. Bogner in the suprageneric system of classifica
(Munich), J. c. French & P. B. Tomlinson tion; however, the accumulation of evi
(Rutgers and Harvard Forest, respec dence from these broad surveys often
tively), M. H. Grayum (formerly of Univ. suggests certain evolutionary trends
of Massachusetts, now Missouri Botani which swing the evidence toward adding
cal Garden), A. Hay, (Royal Botanic or removing elements of any group of
Gardens, Sydney), W. Hetterscheid (Hol plants.
land), M. Madison (formerly of Selby Plant chemistry, poorly known in
Gardens), S. J. Mayo & P. Boyce(Kew), Engler's time, has been surveyed by a
D. H. Nicolson (Smithsonian) and M. number of workers including Hegnauer
Serebryanyi (Moscow). Bogner, Mayo & (963), Gibbs (974), Fairbrothers et al.
Boyce are currently completing the Ar (975), Harris & Hartley (980), Williams
aceae treatment for K. Kubitzki's Fami et al. (981), Dahlgren & Clifford (982),
lies and Genera of Flowering Plants. Harborne (982) and Fox & French
Recently two major subfamilial classifica (988), as well as others. A great deal has
tions were completed (Grayum, 1990; been learned about the cytology of the
Bogner & Nicolson, in press). With the family including extensive surveys by
permission of the authors, both of these Jones (957), Marchant 0970, 1971a,
mOMAS B. CROAT, 1990 49
1971b, 1972, 1974), and especially Pe though out of his study area))
tersen (989). In addition, the important Subfamily n. POmOIDEAE
subject of continental drift (Raven & Tribe 1. POTHEAE (Potbos, Potboid
Axelrod, 1974; Schuster, 1976) has been ium)
helpful in dealing with the intricate Tribe 2. MONSTEREAE (Rbapbido
phytogeographical problems posed by pbora, Anadendrum, A my
this wide-ranging family. drium, Scindapsus)
Other important work includes sur Tribe 3. SPATHIPHYLLEAE (Spatbi
veys of palynology by Thanikaimoni pbyllum, Holocblamys)
(969) and by Grayum (1984). Important Subfamily m. LASIOIDEAE
surveys of molec;:ular systematics in the Tribe 1. SYMPLOCARPEAE (Lysicbiton,
Araceae concentrating on restriction site Symplocarpus)
variation in chloroplast DNA are cur Tribe 2. LASIEAE (Cyrtosperma, Lasia,
rently being carried out by J. C. French. Pycnospatba)
There is already preliminary evidence Tribe 3. AMORPHOPHALLEAE [Thom
that these surveys will be rich in informa sonieae] (1bomsonia = Amor
tion concerning the evolution of the pbopballus, Pseudodracon
Araceae. All of these listed above, cou tium, Amorpbopballus)
pled with many modem revisions of Subfamily Iv. PHITODENDROIDEAE
aroid taxa have combined to yield an Tribe 1. CALLEAE (Calla)
important body of useful knowledge to Tribe 2. AGLAONEMATEAE (Agla-
modem-day revisionists of the Araceae. onema, Aglaodorum)
All three of the modem systems dem Tribe 3. HOMALOMENEAE (Homalom
onstrate Significant differences from that ena, Diandriella = Homalom
of Engler. The system by Hotta will be ena [fide Bogner & Nicolson))
presented first, followed by the system of Tribe 4. SCHISMATOGLOTTIDEAE
Bogner & Nicolson, and finally by that of (Scbismatoglottis, Pbymata
Grayum. rum, Bucepbalandra, Micro
casia = Bucepbalandra, Ari
The Subfamilial Classification of the darum, Piptospatba)
Araceae by Hotta (1970) Tribe 5. COLOCASIEAE (Rem usa tia,
Gonatantbus, Hapaline, Col
The following system by Hotta is
based solely on genera in Eastern Asia ocasia, Alocasia, Scbizocasia =
and Malesia. It deals with members of all Xenopbya = Alocasia)
Subfamlly V. AROIDEAE
of Engler's subfamilies of Araceae but
Tribe 1. AREAE (Typbonium, Ari
includes only 39 genera (fewer if one
saema, Pinellia)
accepts the synonymization of those
Tribe 2. CRYPTOCORYNEAE (Crypto
genera indicated). It cannot thus be
coryne)
considered a thorough revision of the
Subfamily VI. PISTIOIDEAE
family because there are many tribes
(Pistia)
which are restricted to Africa or the
Americas which were not considered
(though some were included in synon Discussion
ymy). The system contains six subfamil
Although Hotta's arrangement of sub
ies, but only 14 tribes, because he was
families is in some ways more radical
not dealing with the entire family.
than that of Bogner & Nicolson, he does
not separate Acorns from the family, but
A Synopsis of Hotta's System
places it in its own subfamily with
Subfamlly I. ACOROIDEAE Gymnostacbys. However, the latter is out
Tribe 1. ACOREAE (Acorns [also Gym of the range of Hotta's study. Hotta
nostacbys in this subfamily, departs radically from Bogner & Ni-
50 AROIDEANA Vol. 13, No. 1-4
colson in merging the subfamily Mon ryninae to tribal level. He makes no
steroideae into the Pothoideae. Six char change in the subfamily Pistioideae.
acteristics are included as justification for Hotta was well ahead of most aroid
this, including: 1) a tendency toward a taxonomists in proposing major changes
climbing habit; 2) the presence of vessels in Engler's system. Some of these pro
in the stems; 3) reticulated leaf blade posed changes have been adopted by
venation; 4) the common occurrence of Grayum.
geniculate petioles; 5) the presence of
bisexual flowers; and 6) the unreliability
The Subfamilial Classification of the
of the character involving the presence
Araceae by Bogner & Nicolson (in
or absence of trichosclereids which has
press)
been used by Engler to separate the two
subfamilies. He also specifically states This system was first presented at the
that the tribe Zamioculcadeae is not Aroid Workshop at Harvard Forest in
closely related to the Pothos and May 1984 and was submitted as a chapter
Rhaphidophora group and places it with (as was the present paper) of "The
the subfamily Lasioideae (not actually Biology of the Araceae," a much-cited,
treating it, but including it in synonymy). but now defunct work to have been
In addition, Hotta also departs further published by Cornell University Press.
from the typical Englerian system in: After the collapse of this proposed book,
1) eliminating the subfamily Calloideae; the paper was accepted for publication
2) submerging Lysichiton, Symplocarpus in Willdenowia, where it will soon ap
and Orontium (though out of the range pear. The classification system is based
of his study), in his tribe Symplocarpeae on more than two decades of critical
in the subfamily Lasioideae; and 3) by observations by both authors, but espe
placing Calla in the tribe Calleae at the cially on the long-standing and intense
head of the subfamily Philodendroideae. interest of the first author. Bogner has
He also placed Dracontium from the cultivated and observed most aroid gen
Lasioideae into synonymy under his tribe era at the Botanical Garden in Munich.
Amorphophalleae (now Thomsonieae). His persevering interest in obtaining live
The Philodendroideae in Hotta's treat material to study has carried him to most
ment remains substantially intact with parts of the world (at his own expense)
respect to the Asian genera. He does not and has given him not only the world's
deal with many of the American and best generic collection of Araceae, but
African tribes of the subfamily. Neverthe also an insight into the taxonomy of the
less, the subfamily is radically altered by family not afforded many of his
the inclusion of Calla as tribe Calleae, the predecessors or contemporaries. Ni
tribe Asterostigmateae (from Engler's colson, owing to his long career with
subfamily Aroideae) and the entire sub Araceae (beginning 25 years ago in Asia),
family Colocasioideae as tribe Coloca his intense bibliographic interest cou
sieae. Though Hotta treats only the tribe pled with language translation skills and
Colocasieae, he synonymizes the entire a keen interest in nomenclatural prob
subfamily Colocasiodeae under the sub lems, makes him a unique addition to the
family Philodendroideae. He also syn team. The system contains nine subfami
onymizes Engler's tribe Philodendreae. lies, 35 tribes and 13 subtribes.
Hotta's subfamily Aroideae deals with
only a few genera. Aside from removing
A Synopsis of Bogner & Nicolson's
the Asterostigmateae (as mentioned
System
above), he basically follows Engler's
classification for the included genera he Subfamily 1. GYMNOSTACHYDOI
treats but raises the subtribe Cryptoco- DEAE
51
THOMAS B. CROAT, 1990
Tribe 1. Gymnostachydeae (1. Gym Hottarum, 46. Bucephalandra,
nostachys) 47. Phymatarum, 48. Ari
Subfamily n. POTHOIDEAE darum, 49. Heteroaridarum)
Tribe 1. Potheae (2. Pothos, 3. Pedicel Subtribe 3. PHILODEN-
larum, 4. Pothoidium) DRINAE (50. Philodendron)
Subfamily ill. MONSTEROIDEAE Tribe 2. ANUBIADEAE (51. Anubias)
Tribe 1. ANADENDREAE (5. Anaden Tribe 3. BOGNEREAE (52. Bognera)
drum) Tribe 4. AGLAONEMATEAE (53.
Tribe 2. MONSTEREAE (6. Amydrium, Aglaonema, 54. Aglaodorum)
7. Rhaphidophora, 8. Epi Tribe 5. DIEFFENBACHIEAE (55. Die/
premnum, 9. Scindapsus, fenbachia)
10. Alloschemone, 11. Steno Tribe 6. ZANTEDESCHIEAE (56.
spermation, 12. Rhodospatha, Zantedeschia)
13. Monstera) Tribe 7. TYPHONODOREAE (57. Ty-
Tribe 3. HETEROPSIDEAE (14. Heter phonodorum)
opsis) Tribe 8. PELTANDREAE (58. Peltandra)
Tribe 4. SPATHIPHYLLEAE 05. Spa- Subfamily VII. COLOCASIOIDEAE
thiphyllum, 16. Holochlamys) Tribe 1. CALADIEAE (59. Xanthosoma,
Subfamily Iv. CAU..OIDEAE 60. Chlorospatha, 61. Cala
Tribe 1. Calleae (17. Calla) dium, 62. Scaphispatha, 63.
Subfamily V. LASIOIDEAE jasarum)
Tribe 1. ORONTIEAE 08. Lysichiton, Tribe 2. STEUDNEREAE
19. Symplocarpus, 20. Oron SubTribe 1. STEUDNERI
tium) NAE (64. Steudnera, 65. Re
Tribe 2. ANTHURIEAE (21. Anthurium) musatia, 66. Gonatanthus)
Tribe 3. LASIEAE SubTribe 2. HAPALIN-
Subtribe 1. DRACONTIINAE INAE (67. Hapaline)
(22. Cyrtosperma, 23. Lasiomor Tribe 3. PROTAREAE (68. Protarum)
pha, 24. Lasia, 25. Anaphyllum, Tribe 4. COLOCASIEAE (69. Colocasia,
26. Anaphyllopsis, 27. Podola 70. Alocasia)
sia, 28. Urospatha, 29. Dracon Tribe 5. SYNGONIEAE (71. Syngo-
tioides, 30. Dracontium nium)
Subtribe 2. PYCNOSPA Tribe 6. ARIOPSIDEAE (72. Ariopsis)
THINAE C31. Pycnospatha) Subfamily VIII.AROIDEAE
Tribe 4. ZAMIOCULCADEAE (32. Zami Tribe 1. STYLOCHAETONIEAE (73. Sty
oculcas, 33. Gonatopus) lochaeton)
Tribe 5. CALLOPSIDEAE (34. Callopsis) Tribe 2. AROPHYTEAE 04. Carle
Tribe 6. NEPHTHYTIDEAE C35. Pseu phyton, 75. Colletogyne, 76.
dohydrosme, 36. Anchomanes, Arophyton)
37. Nephthytis, 38. Cercestis) Tribe 3. SPATHICARPEAE (77. Man
Tribe 7. CULCASIEAE (39. Culcasia) gonia,78. Taccarum, 79. Aster
Tribe 8. MONTRICHARDIEAE (40. Mon ostigma, 80. Gorgonidium, 81.
trichardia) Synandrospadix, 82. Gearum,
Subfamily VI. PHILODENDROIDEAE 83. Spathantheum, 84. Spathi
Tribe 1. PHILODENDREAE carpa)
Subtribe 1. HOMALOM Tribe 4. ZOMICARPEAE (85. Zomi
ENINAE (41. Furtadoa, 42. carpa, 86. Filarum, 87. Zomi
Homalomena) carpella, 88. Ulearum)
Subtribe 2. SCHISMATOG Tribe 5. THOMSONIEAE (89. Amor
LOTTIDINAE (43. Schisma phophallus, 90. Pseudodra
tog lottis, 44. Piptospatha, 45. contium)
52 AROIDEANA Vol. 13, No. 1-4
Tribe 6. AREAE the separation of Gymnostachys from the
Subtribe 1. ARINAE (91. Pothoideae into the subfamily Gym
Arum, 92. Dracunculus, 93. nostachydoideae; 3) the transfer of the
Helicodiceros, 94. Therio Anthurieae from the Pothoideae to the
phonum, 95. Typhonium, 96. Lasioideae; 4) the Heteropsidae from
Sauromatum, 97. Eminium, Pothoideae to Monsteroideae; 5) the
98. Biarum) Orontieae from the Calloideae to Lasioi
Subtribe 2. ARISARINAE deae; 6) the Thomsonieae from the
(99. Arisarum) Lasioideae to Aroideae; and 7) the Cal
Sub tribe 3. ARISAEMATI lopsideae from Aroideae to Lasioideae.
NAE (100. Arisaema) The suggestion that Acorus was not a
Subtribe 4. ATHERURINAE good member of the Pothoideae was
001. Pinellia) already accepted by several other au
Subtribe 5. AMBROSI thors (Eyde et al., 1967; Hotta, 1970;
NINAE 002. Ambrosina) Thorne, 1976, 1983), but its exclusion
Subtribe 6. CRYPTOCO from the Araceae was first suggested by
RYNINAE 003. Lagenandra, Deyl (955) and later by Grayum 0984,
104. Cryptocoryne) 1987) and Tillich (985). Other signifi
Subfamily IX. PISTIOIDEAE cant changes involved major realign
005. Pistia)
ments in the Pothoideae and Lasioideae.
Many alterations involved changes in
Discussion rank within the subfamilies or the move
ment of a few genera from one estab
Bogner & Nicolson's system, with 105
lished subfamily to another. Anaden
genera, reduces the number of genera
drum was moved from tribe Potheae in
from the 110 recognized by Bogner
the subfamily Pothoideae to its own
(978). Those genera reduced to synon
tribe, the Anadendreae in the Monsteroi
ymy since the 1978 paper are: 1bom
deae. Also transferred from the Pothoi
sonia and Plesmonium = Amorphophal
deae were the tribes Zamioculcadeae
Ius, Echidnium = Dracontium, Rhekto
and Culcasieae, which were placed in
phyllum = Cercestis, Diandriella =
the subfamily Lasioideae. Other genera
Homalomena. Two other genera re
transferred were Protarum (tribe Protar
duced to synonymy in recent years are:
eae) to the Colocasioideae, and
Caladiopsis = Chlorospatha (Madison,
1981), and Porphyrospatha = Syngonium Scaphispatha (tribe Zomicarpeae) from
the subfamily Aroideae to the tribe
(Croat, 1981).
Five genera have been added since Caladieae in the subfamily Colocasioi
Bogner's 1978 list was published in deae (Bogner, 1980). Earlier (Bogner
Aroideana. These include the reinstate 1980a), jasarum was moved from the
ment of Alloschemone next to Scindap subtribe Alocasiinae to the tribe
sus in the Monstereae, the incorporation Caladieae.
of Bognera following the tribe Anubia Some subtribes were merged, such as
deae in the subfamily Philodendroideae, the Alocasiinae into Colocasiinae. In
Furtadoa in the subtribe Homalom other cases, new subtribes were created,
eninae of tribe Philodendreae, and La such as in the Lasioideae, with the
siomorpha and Anaphyllopsis in the subtribe Dracontiinae of the Lasieae
subtribe Dracontiinae of the tribe La accomodating all genera in the tribe
sieae. Lasieae except Pycnospatha. The new
The most Significant changes in the subtribe Pycnospathinae contains only
revised system by Bogner & Nicolson, as the latter genus (Bogner, 1973).
outlined above, include the following: 1) The subtribe Steudnerinae in the Colo
the removal of Acorus from the family; 2) casioideae was elevated to tribal status
THOMAS B. CROAT, 1990 53
(Steudnereae) and split into two sub A Synopsis of Grayum's System
tribes which are the Steudnerinae with
I. Subfamily P011IOIDEAE
Steudnera, Remusatia and Gonatanthus
1. Tribe G YMNOSTACHYDEAE
and the Hapalininae with Hapaline. Also
( Gymnostachys)
elevated was the subtribe Caladiinae in
2. Tribe SPATHIPHYLLEAE (Spa
the subfamily Colocasioideae, which be
thiphyllum, Holochlamys)
came the tribe Caladieae. It was also
3. Tribe ANTHURIEAE (Anthurium)
shifted to the first tribe of the subfamily.
The remainder of the changes incorpo 4. Tribe POTHEAE (Pothos, Pedicel
rated by Bogner & Nicolson involved larnm, Pothoidium)
changing positions of genera within 5. Tribe ANADENDREAE (Anad
existing tribes or subtribes. These in endrum)
clude movement of 1) Amydrium closer 6. Tribe MONSTEREAE
to Rhaphidophora in the tribe Mon a. Subtribe HETEROPSIDINAE
stereaej 2) Piptospatha from the last (Heteropsis)
position in the subtribe Schismatoglottid b. Subtribe MONSTERINAE
inae (subfamily Philodendroideae) to the (Rhaphidophora, Monstera,
second position following Schismatoglot Amydrium, Epipremnum,
tis, and 3) moving Hottarnm from the Sc in daps us, Alloschemone,
next to the last position in subtribe Stenospermation, Rhodospa
Schismatoglottidinae to the third position tha)
following Piptospatha. 7. Tribe ZAMIOCULCADEAE (Zami-
Overall the proposed changes in the oculcas, Gonatopus)
Englerian system were generally conser II. Subfamily CALLOIDEAE (Referred
vative ones which have definitely re to later in this paper for comparative
sulted in an improvement in the subfa purposes only as "Philodendroideae")
milial classification. A. Galla Alliance
8. Tribe CALLEAE (Galla)
The Subfamilial Classification of the B. Nephthytis Alliance
Araceae by Grayum (1990) 9. Tribe NEPHTHYTIDEAE (Neph
thyt is, Anchomanes, Pseudohy
Working concurrently with Bogner &
drosme)
Nicolson yet independently from them,
M. H. Grayum, under the direction of J. 10. Tribe CALLOPSIDEAE (Gallopsis,
Ulearnm, Pilarnm, Zomicarpella)
Walker at the University of Massachusetts
(Amherst), prepared his own subfamilial 11. Tribe MONTRICHARDIEAE (Mon
trichardia)
classification of the family. His research,
though concentrating on the first rigor C. Aglaonema Alliance
ous survey of pollen using scanning 12. Tribe ANUBIADEAE (Anubias)
electron microscopy, also involved the 13. Tribe ZANTEDESCHIEAE (Zante
most thorough analysis of all morpholog deschia)
ical character states since the time of 14. Tribe AGLAONEMATEAE (Aglao
Engler and of the three systems analyzed, nema, Aglaodornm)
his is the only one which is accompanied 15. Tribe SPATHICARPEAE (Man
by a complete explanation of the ration gonia, Asterostigma, Synan
ale behInd the placement of the taxa drospadix, Taccarnm, Gorgo
involved. nidium, Gearnm, Spathantheum,
Grayum has proposed the most radical Spathicarpa)
alterations in the subfamilial classifica 16. Tribe DIEFFENBACHIEAE (Die!
tion of the Araceae to date. The system jenbachia)
contains five subfamilies, 40 tribes and 13 17. Tribe BOGNEREAE (Bognera)
subtribes. D. Peltandra Alliance