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 : 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 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 ....phanerogams. 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: (Hotta, tension of the one devised by J. D. 1981) and (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 . Schott, 20. Amydrium Schott) Lasiomorpha was resurrected to the ge­ 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. 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. Lour. 29. Subfamily I. POTHOIDEAE Engl. Schott, 29A. Hay, (Properly ACOROIDEAE, according to the rules of nomenclature when Acarus 30. N. E. Brown, 31. is included.) Schott [Bogner], Tribe 1. POTHEAE Engl. (1. Pothos L., 2 .. (1988, 1989) 32. Pothoidium Schott, *2A. Engl., 33. Echidnium Schott = Pedicellarum M. Hotta, 3. [Bogner], 34. Dra­ Anadendrum Schott (as Anad­ contium L., *34A. endron), 3a. Epipremnopsis Thorel ex Gagnep.) Engl. = Amydrium [Nicolson]) Tribe 2. AMORPHOPHALLEAE Eng!. Tribe 2. HETEROPSIDEAE Engl, (4. (Now correctly THOM­ 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. [Bogner], 37. P. Beauv.) Schott, 38. 1bomsonia Wall. = Tribe 5. ZAMIOCULCADEAE Engl. (7. [Bogner, 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.) Schott, 42. Subfamily II. Schott, 43. Rhektophyllum N. E. Engl. Brown = Cercestis [Bogner]) Tribe 1. MONSTEREAE Eng!. (ll. Tribe 4. MONTRICHARDIEAE Eng!. Rhaphidophora Hassk. (as Ra­ (44. 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. 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. 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. 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, = 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) = 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. [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) toglottis [Bunting]) Subfamily VII. Engl. Tribe 2. ANUBIADEAE Eng!. (54A. Tribe 1. STYLOCHAETONIEAE Schott Amauriella Rendle = (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) S. Buchet *76c. Tribe 3. AGLAONEMATEAE Engl. (55. togyne Aro­ pbyton ]um.) Schott, 56. Schott) Tribe 2. ASTEROSTIGMATEAE Schott Tribe 4. DIEFFENBACHIEAE Engl. (57. (77. Mangonia Schott, 78. An­ Dieffenbaebia Schott) dromyeia A. Rich. = Aster­ Tribe 5. ZANTEDESCHIEAE Eng!. (58. ostigma [Bogner), 79. Tae­ Zantedesebia Spreng.) earum Brongn. ex Schott, 80. Tribe 6. TYPHONODOREAE Eng!. (59. Fisch. & Mey., 81. Typbonodorum Lind!.) Eng!., 82 . Tribe 7. PELTANDREAE Eng!. (60. Pel­ Spatbantbeum Schott, 83. tandra Raf.) Schott, 84. Subfamily VI. COLOCASIOIDEAE 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. Tribe 4. CALLOPSIDEAE Eng!. (87. Cal­ K. Koch, 62. Schott, lopsis Eng!.) 63. Gonatantbus Klotzsch Tribe 5. ZOMICARPEAE Eng!. (88. Subtribe 2. HAPALININAE Seapbispatba Brongn. ex Eng!. & K. Kr. (64. Schott, 89. Xenopbya Schott = Schott) Aloeasia [Hay), 90. Zomiearpa Subtribe 3. CALADIINAE Schott, 91. Zomiearpella N. E. Eng!. & K. Kr. (65. Caladiopsis Brown, *91A. Nicolson, Eng!. = Cblorospatba [Madi­ 92. Eng!.) son), 66. Caladium Vent., *66A. Tribe 6. Engl. 48 AROIDEANA Vol. 13, No. 1-4

Subtribe 1. ARINAE Schott systems will be presented here in synop­ (93. Arnm 1., 94. 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 Schott, 98. Sauro­ benefited from a substantial amount of matum Schott, 99. 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. anatomy (Eyde et aI., 1967) as well as Mart.) blade nervature (Ertl, 1932), embryology Subtribe 4. PlNELLIINAE Qiissen, 1928), and of seedling morphol­ Schott 003. 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), 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. 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 It is important to give recognition to study of a single character rarely pro­ the work of a small group of active vides conclusive evidence for the cor­ researchers working in different parts of rectness of the placement of any member 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 . land), M. Madison (formerly of Selby 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 (, 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, The following system by Hotta is Gonatantbus, Hapaline, Col­ based solely on genera in Eastern Asia ocasia, Alocasia, Scbizocasia = and Malesia. It deals with members of all Xenopbya = Alocasia) of Engler's subfamilies of Araceae but Subfamlly V. AROIDEAE includes only 39 genera (fewer if one Tribe 1. AREAE (Typbonium, Ari­ accepts the synonymization of those saema, Pinellia) genera indicated). It cannot thus be Tribe 2. CRYPTOCORYNEAE (Crypto­ considered a thorough revision of the coryne) family because there are many tribes Subfamily VI. PISTIOIDEAE which are restricted to or the (Pistia) 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 A Synopsis of Hotta's System not separate Acorns from the family, but 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 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 THOMAS B. CROAT, 1990 51

Tribe 1. Gymnostachydeae (1. Gym­ Hottarum, 46. , nostachys) 47. , 48. Ari­ Subfamily n. POTHOIDEAE darum, 49. Heteroaridarum) Tribe 1. Potheae (2. Pothos, 3. Pedicel­ Subtribe 3. PHILODEN- larum, 4. Pothoidium) DRINAE (50. ) 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, ) 13. Monstera) Tribe 7. TYPHONODOREAE (57. Ty- Tribe 3. HETEROPSIDEAE (14. Heter­ phonodorum) opsis) Tribe 8. PELTANDREAE (58. ) Tribe 4. SPATHIPHYLLEAE 05. Spa- Subfamily VII. COLOCASIOIDEAE thiphyllum, 16. Holochlamys) Tribe 1. CALADIEAE (59. , 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. ) pha, 24. Lasia, 25. Anaphyllum, Tribe 4. COLOCASIEAE (69. , 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. ) THINAE C31. Pycnospatha) Subfamily VIII.AROIDEAE Tribe 4. ZAMIOCULCADEAE (32. Zami­ Tribe 1. STYLOCHAETONIEAE (73. Sty­ oculcas, 33. Gonatopus) lochaeton) Tribe 5. CALLOPSIDEAE (34. ) 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. , 79. Aster­ Tribe 8. MONTRICHARDIEAE (40. Mon­ ostigma, 80. Gorgonidium, 81. trichardia) Synandrospadix, 82. Gearum, Subfamily VI. PHILODENDROIDEAE 83. , 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 toglottis, 44. , 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 , 92. Dracunculus, 93. nostachydoideae; 3) the transfer of the , 94. Therio­ Anthurieae from the Pothoideae to the phonum, 95. Typhonium, 96. Lasioideae; 4) the Heteropsidae from , 97. Eminium, Pothoideae to Monsteroideae; 5) the 98. ) Orontieae from the Calloideae to Lasioi­ Subtribe 2. ARISARINAE deae; 6) the Thomsonieae from the (99. ) 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. ) Thorne, 1976, 1983), but its exclusion Subtribe 6. CRYPTOCO­ from the Araceae was first suggested by RYNINAE 003. , Deyl (955) and later by Grayum 0984, 104. ) 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 from the 110 recognized by Bogner drum was moved from tribe Potheae in (978). Those genera reduced to synon­ the subfamily Pothoideae to its own ymy since the 1978 paper are: 1bom­ tribe, the Anadendreae in the Monsteroi­ deae. Also transferred from the Pothoi­ sonia and Plesmonium = Amorphophal­ deae were the tribes Zamioculcadeae Ius, Echidnium = Dracontium, Rhekto­ phyllum = Cercestis, Diandriella = and Culcasieae, which were placed in Homalomena. Two other genera re­ the subfamily Lasioideae. Other genera duced to synonymy in recent years are: transferred were Protarum (tribe Protar­ Caladiopsis = Chlorospatha (Madison, eae) to the Colocasioideae, and 1981), and Porphyrospatha = Syngonium Scaphispatha (tribe Zomicarpeae) from (Croat, 1981). the subfamily Aroideae to the tribe 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 (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 Steudnera, Remusatia and Gonatanthus I. Subfamily P011IOIDEAE and the Hapalininae with Hapaline. Also 1. Tribe G YMNOSTACHYDEAE elevated was the subtribe Caladiinae in ( Gymnostachys) the subfamily Colocasioideae, which be­ 2. Tribe SPATHIPHYLLEAE (Spa­ came the tribe Caladieae. It was also thiphyllum, Holochlamys) shifted to the first tribe of the subfamily. 3. Tribe ANTHURIEAE (Anthurium) 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, , 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 Working concurrently with Bogner & thytis, Anchomanes, Pseudohy­ Nicolson yet independently from them, drosme) M. H. Grayum, under the direction of J. 10. Tribe CALLOPSIDEAE (Gallopsis, Walker at the University of Massachusetts Ulearnm, Pilarnm, Zomicarpella) (Amherst), prepared his own subfamilial 11. Tribe MONTRICHARDIEAE (Mon­ classification of the family. His research, trichardia) though concentrating on the first rigor­ C. Aglaonema Alliance ous survey of 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 ) 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 54 AROIDEANA Vol. 13, No. 1-4

18. Tribe PELTANDREAE (Peltandra, 31. Tribe STYLOCHAETONIEAE (Sty­ ) lochaeton) 19. Tribe AROPHYTEAE (Arophyton, V. Subfamily AROIDEAE Carlephyton, Colletogyne) 32. Tribe THOMSONIEAE (Pseu- 20. Tribe SCHISMATOGLOTTIDEAE dodracontium, Amorphophallus) (, Piptospatha, 33. Tribe ARISAREAE (Arisarum) Bucephalandra, Phymata rum, 34. Tribe PINELLIEAE (Pinellia) , Heteroaridarum, Hot­ 35. Tribe PISTIEAE (Pistia) tarum) 36. Tribe CRYPTOCORYNEAE (Cryp­ E. Philodendron Alliance tocoryne, Lagenandra) 21. Tribe CULCASIEAE (Culcasia) 37. Tribe AMBROSINEAE (Am­ 22. Tribe CERCESTIDEAE (Cercestis) brosina) 23. Tribe HOMALOMENEAE (Fur­ 38. Tribe ARIOPSIDEAE (Ariopsis) tadoa, Homalomena) 39. Tribe ARISAEMATEAE (Ari­ 24. Tribe PHILODENDREAE (Phil­ saema) odendron) 40. Tribe AREAE (Arum, Dracun­ m. Subfamily COLOCASIOIDEAE culus, Helicodiceros, Therio­ 25. Tribe ZOMICARPEAE (Zomi­ phonum, Typhonium, Sauro­ carpa) matum, Eminium, Biarum) 26. Tribe COLOCASIEAE * These genera added since 1990 paper a. Subtribe PROTARINAE (Prota­ (Grayum, pers. comm.). rum) b. Subtribe STEUDNERINAE (Ste­ Discussion udnera) In agreement with Hotta (1970), c. Subtribe REMUSATIINAE (Re­ Grayum also merges the Pothoideae and musatia, Gonatanthus) Monsteroideae and places the tribes d. Subtribe COLOCASIINAE (Col­ Calleae, Spathicarpeae and ocasia, Alocasia) Zantedeschieae in the Calloideae (previ­ 27. Tribe CALADIEAE Schott ously named Philodendroideae). Unlike a. Subtribe JASARINAE (Jasarum) Hotta, he the tribe Zamiocul­ b. Subtribe SCAPHISPATHINAE cadeae in the Pothoideae. Similarities (Scaphispatha) with the system of Hotta end here, but c. Subtribe CALADIINAE (Ca­ Grayum makes other radical departures ladium, Xanthosoma, Chloro­ from the Englerian system. Because the spatha, Aphyllarum) changes made by Grayum in Engler's d. Subtribe SYNGONIINAE (Syn­ system are many, the discussion will gonium) proceed section by section. e. Subtribe HAPALININAE (Hapaline) POTHOIDEAE Iv. Subfamily LASIOIDEAE 28. Tribe SYMPLOCARPEAE (Symplo­ Grayum's system appeared originally carpus, Lysichiton) in his Ph.D. thesis (Grayum, 1984) and 29. Tribe ORONTIEAE (Orontium) was subsequently published with some 30. Tribe LASIEAE modifications (Grayum, 1990). This a. Subtribe DRACONTIINAE (An­ paper will deal primarily with the latest aphyllopsis*, Cyrtosperma, Lasia, revision. Grayum considers the linear Lasiomorpha*, Anaphyllum, sequencing of his taxa of no significance. Podolasia, Urospatha, Dracontioi­ Although Grayum (984) removed Gym­ des, Dracontium) nostachys from Engler's Acoreae to its b. Subtribe PYCNOSPATHINAE own subfamily, the Gymnostachydoi­ (Pycnospatha) deae, his 1990 revision retains it in the THOMAS B. CROAT, 1990 55 subfamily Pothoideae and the tribe Gym­ 5. The movement of the tribe Mon­ nostachydeae. With the subfamily Mon­ trichardieae from the Lasioideae to steroideae merged with the Pothoideae, the Nephthytis Alliance. Anadendrnm is retained in the tribe 6. The movement of the tribe Spathi­ Potheae and placed in its own tribe carpeae (Engler's Asterostig­ (Anadendreae). The tribe Heteropsideae mateae) to Calloideae in the is reduced to a subtribe of the Mon­ Aglaonema Alliance. stereae. The Aglaonema Alliance is otherwise made up completely of Englerian phil­ odendroid members with the tribes Anu­ CALLOIDEAE sensu Engler biadeae, Zantedeschieae and Aglaone­ The subfamily Calloideae of Schott is mateae clustered there. disbanded, with Symplocarpus, Lysichi­ The Peltandra Alliance finds tribes ton and Orontium comprising two sub­ Peltandreae and Typhonodoreae merged tribes in the tribe Orontieae. Philoden­ into the former. A major change in the droideae, for reasons of priority, is re­ Peltandra Alliance is the transfer of tribe named as subfamily Calloideae. Arophyteae from the Aroideae. Finally the Aglaonema Alliance differs radically from the Englerian system in PHILODENDROIDEAE sensu Engler that the entire tribe Asterostigmateae CALLOIDEAE from the subfamily Aroideae is trans­ As redefined, the subfamily Calloideae ferred to subfamily Calloideae. Grayum's is arranged in five major "alliances": Aglaonema Alliance underwent changes Calla, Nephthyis, Aglaonema, Peltandra, from his 1984 treatment. These include: and Philodendron. 1. The removal of the Homalomeneae The main changes within the Calloi­ to the Philodendron Alliance. deae include a number of taxa that are 2. The inclusion of tribe Dieffen­ added to the subfamily from other sub­ bachieae, with Diriffenbachia only, families. These include: and Bognereae, with Bognera only. 1. The addition of the tribe Culcasieae Grayum (pers. comm.), based on to the Nephthytis Alliance from the unpublished investigations by Pothoideae. Bogner, now believes that Bognera 2. The movement of the tribe Neph­ should be included in tribe Dieffen­ thytideae from Lasioideae to the bachieae. Nephthytis Alliance. 3. The placement of the genera An­ COLOCASIOIDEAE chomanes and of the tribe Thomsonieae (Amor­ Substantial changes were also made in phophalleae) of the Lasioideae into the subfamily Colocasioideae, the princi­ the tribe Nephthytideae in the pal change being the transfer of Zomicar­ Nephthytis Alliance. Grayum placed peae (now to include both Zomicarpella Zomicarpella in the tribe Callop­ and Zomicarpa according to Grayum, sideae, but now believes (pers. pers. comm.) from Engler's Aroideae and comm.), as a result of Jim French's the recognition of two coordinate tribes, recent discovery of laticifers in Colocasieae and Caladieae, the first in Zomicarpella, that it really belongs the Old World, the second in the New in Zomicarpeae of the Colocasioi­ World (except Hapalininae). Grayum deae. accepts two of Engler's three tribes, e.g., 4. The transfer of tribe Callopsideae Colocasieae (much altered) and Syngo­ from the Aroideae to the Nephthytis nieae (as a subtribe), but he creates more Alliance. subtribes in the Colocasioideae. The 56 AROIDEANA Vol. 13, No. 1-4 following alterations occur in his treat­ AROIDEAE ment of the subfamily. 1. The tribe Ariopsideae is moved to Perhaps the most important change in the subfamily Aroideae. the subfamily Aroideae is the inclusion of 2. The tribe Protareae is moved from the genus Pistia, which previously was the subfamily Aroideae and placed generally included in its own subfamily. in the tribe Colocasieae as subtribe­ Other important changes in the sub­ Protarinae. (Bogner & Nicolson family Aroideae include: agree with its placement in the 1. The transfer of tribe Thomsonieae Colocasioideae.) (Amorphophalleae) from Lasioi­ 3. The subtribe Steudnerinae is not deae. moved, but the subtribe Remusati­ 2. The transfer of tribe Ariopsideae inae is segregated out with Re­ from the Colocasioideae to the musatia and Gonatanthus (consid­ Aroideae. ered probably synomymous). 3. The movement of tribe Stylochae­ 4. The subtribe Colocasiinae absorbs tonieae to Lasioideae. the subtribe Alocasiinae, while the 4. A shift of Zomicarpeae, Spathicar­ Hapalininae is moved to tribe peae, and Callopsideae to the Caladieae. "Philodendroideae. » 5. jasarum, added to the system since 5. Transfer of Protareae, which be­ Engler's revision, is placed in its comes subtribe Protarinae, to the own subtribe Jasarinae at the head Colocasioideae and breaking up of of the tribe Caladieae. the Zomicarpeae with Ulearum, 6. Scaphispatha is transferred (follow­ Pilarum or Zomicarpella being in­ ing Bogner) into the Colocasioi­ corporated in the tribe Callopsidae deae from the tribe Zomicarpeae of subfamily Calloideae and Zomi­ (subfamily Aroideae) and is placed carpa being placed in the Coloca­ in the Caladieae in its own subtribe. sioideae (as tribe Zomicarpeae). Grayum now (pers. comm.) would also place Zomicarpella here. LASIOIDEAE These changes radically alter the Major changes within the Lasioideae make-up of the subfamily Aroideae. The include: remainder of this subfamily, including all 1. The movement of the tribe Thom­ of the tribe Areae, survives intact. How­ sonieae (Amorphophalleae) to the ever, the subtribe Arineae is elevated to subfamily Aroideae. tribal status. 2. The placement of tribes Neph­ It is evident that Grayum has made thytideae and Montrichardieae in major alterations in Engler's system of the Philodendroideae, substantially classification. Though it is not the pur­ reducing the size of the subfamily pose of this discourse to explain the Lasioideae. As already mentioned, rationale behind the many changes pro­ however, three temperate genera posed, Grayum has made a convincing from the tribe Orontieae are added argument in most cases for the changes to the Lasioideae. made. His system will no doubt be 3. The division of Engler's tribe La­ carefully considered within the next few sieae (following Bogner) into the years. Now that there are two newly two subtribes Dracontiinae and proposed suprageneric classification sys­ Pycnospathinae, with the latter tems, research efforts have been directed containing only Pycnospatha. at a resolution of the differences between 4. The transfer of the tribe Stylochae­ these two systems in an attempt to come tonieae () from the Ar­ up with a system that can be agreed upon oideae to the Lasioideae. by most aroid workers. THOMAS B. CROAT, 1990 57

General Comparisons of Three drum and Heteropsis are transitional Recent Systems of Classification of genera as well. tbeAraceae While Grayum sees little relationship between the Pothoideae and the subfam­ One of the most difficult things about ily Lasioideae, Bogner & Nicolson place making the comparison of the three two of Engler's original pothoid tribes in systems involved the subfamilies Phil­ odendroideae and Calloideae. Because the Lasioideae, namely the Anthurieae and Zamioculcadeae. Grayum leaves Grayum included Calla in the Philoden­ droideae and because the name Calloi­ both in the subfamily Pothoideae. This deae has priority, the subfamily name would appear to be one of the several had to be changed. In order to avoid more marked differences between the confusion between the Calloideae of system of Bogner & Nicolson and that of Bogner & Nicolson (containing only Grayum. Both Hotta and Grayum dis­ Calla) and the Calloideae of Grayum, I band the Calloideae altogether and place will refer to Calloideae of Grayum for the the Orontieae in the Lasioideae while purposes of this final comparison as placing the tribe Calleae in the "Phil­ "Philodendroideae. " odendroideae." Bogner & Nicolson re­ All three systems outlined here indi­ tain Calla in the monotypic subfamily cate that at least the genus Acorus does Calloideae. They agree with the other not fit well into the main body of the two authors in placing Lysichiton, Pothoideae. Hotta removes it, along with Symplocarpus, and Orontium in the Gymnostachys, to the subfamily Acoroi­ Lasioideae. While Bogner & Nicolson deae. Bogner & Nicolson, as well as include all three of these temperate Grayum, go further in rejecting it from genera in the same tribe, Grayum sepa­ the family all together. rates Orontium into its own tribe. Following Deyl (955), Grayum 0984, Aside from the placement of the 1987) proposed to remove Acorus from Symplocarpeae in the Lasioideae, Hotta the Araceae into its own family. He cites leaves the Lasioideae intact. Bogner & 15 major characters in which Acorus Nicolson agree with the transfer of differs from all other Araceae (including Symplocarpeae, but also add to the Gymnostachys). Tillich (1985) independ­ Lasioideae the Anthurieae, Culcasieae, ently arrived at the same conclusion and the Zamioculcadeae from the Pot­ based on his study of aroid seedling hoideae. In addition, they transfer the morphology. Callopsideae from the Aroideae to the All three systems reflect a close rela­ Lasioideae. Hotta places Cu/casia here as tionship between Pothoideae and the well. Grayum, on the other hand, places Monsteroideae. Both Hotta and Grayum the Culcasieae in the "Philoden­ treat the two subfamilies as one, the droldeae," moving Anchomanes and Pothoideae. Bogner & Nicolson include Pseudohydrosme from the Lasioideae to Anadendrum and Heteropsis in the Mon­ "Philodendroideae" as tribe Neph­ steroideae and further suggest that thyideae. He also moves the Montrichar­ Amydrium is a transitional genus (partly dieae and Nephthytideae to "Philoden­ in that it lacks trichosclereids) which droideae." could be placed in either subfamily Despite what appears to be a lot of "depending on what characters one em­ disagreement with the placement of phasizes in delimiting the two subfami­ many of the above-mentioned genera, lies." Grayum states (pers. comm.) that Grayum, as well as Bogner & Nicolson, Pothos and Pothoidium are also transi­ agree that the Culcasieae, Neph­ tional in the same sense in that they thytideae, Callopsidae, Montrichardia possess trichosclereids and that Anaden- and Anchomanes are all related. They 58 AROIDEANA Vol. 13, No. 1-4 just do not agree to which subfamily they Alocasia in their tribe Colocasieae. belong. Grayum includes the Old World genera In the subfamily "Philodendroideae, " Colocasia and Alocasia in the subtribe Hotta and Grayum agree on the inclusion Colocasiinae of tribe Colocasieae and of Calia, but Hotta broadens the subfam­ places the other four New World genera ily substantially by including all of the (Caladium, Xanthosoma, Chlorospatha Colocasioideae (Le., at least the Asian and Aphyllarum) in his subtribe Cala­ genera he studied). The "Philodendroi­ diinae of tribe Caladieae along with deae" ()f Grayum is substantially larger Hapalininae. than that of Bogner & Nicolson, with 40 Except for moving the tribe Asterostig­ genera arranged in 17 tribes and five mateae to the subfamily Philodendroi­ alliances. This contrasts with the 18 deae, Hotta appears to have left the genera in eight tribes according to subfamily Aroideae intact, although he Bogner & Nicolson. The size of Grayum's did not deal with it in great detail. The "Philodendroideae" has resulted largely other two systems agree on the move­ from the inclusion of Spathicarpeae from ment of Protareae to Colocasioideae as the subfamily Aroideae as well as the well as on the removal of the Callop­ other tribes from Pothoideae and Lasioi­ sideae. Grayum places the latter in his deae already mentioned. The latter in­ "Philodendroideae" while Bogner & Ni­ clude Calleae, Culcasieae, Neph­ colson placed it in the Lasioideae. In thytideae, Callopsideae, and Montrichar­ addition Grayum transfers the tribe Th­ dieae. omsonieae (Amorphophalleae) from the Other major departures of Grayum Lasioideae to the Aroideae, and also the Zomicarpeae (which Bogner & Nicolson from Bogner & Nicolson include the maintain) from the Aroideae partly to the incorporatlon of Arophyteae and Schis­ Colocasioideae and partly to the Neph­ matoglottideae into the "Philodendroi­ thytis Alliance. More importantly, he deae." The latter is placed in the Peltan­ adds Pistia, thus eliminating the subfam­ dra Alliance. Bogner & Nicolson, on the ily Pistioideae. The two systems other­ other hand, retain Arophyteae in the wise agree except for the order of the Aroideae (Bogner, 1978) along with the taxa within the subfamily, which Grayum Cryptocoryninae. Hotta also leaves Cryp­ regards as of no significance. tocoryninae in the Aroideae, but does not deal with Arophyteae. Hotta incorporates the Colocasioideae Conclusion into the Philodendroideae while Bogner It is clear from the comparisons above & Nicolson leave it essentially intact, that despite recent studies (or perhaps except for the inclusion of Protareae even because of them), disagreement from the Aroideae. Grayum's Colocasioi­ still exists concerning the placement of deae differs from Bogner & Nicolson's taxa at all levels within the family but largely in the arrangement of the tribes especially on the composition of the within the subfamily. Both treatments subfamilies. It is for this reason that no include the same genera with the excep­ attempt will be made here to construct a tion of Ariopsis which Grayum places in description of the respective subfamilial the Aroideae. Grayum also moves groupings. Further work in a variety of jasarum and Scaphispatha to their own disciplines will be needed on the more subtribes and reduces the tribe Protareae poorly known genera. It is expected that to a subtribe of his tribe Colocasieae. the new molecular systematics studies Bogner & Nicolson include Xanthosoma, being carried out by James French and Aphyllarum, Chlorospatha, Caladium, his colleagues from Rutgers will provide Scaphispatha, and jasarum in their tribe the next important source of information Caladieae while including Colocasia and for a realignment of the Araceae. I hope THOMAS B. CROAT, 1990 59 that a consensus will emerge regarding ____ & P. F. Yeo. 1985. The Families the classification of the Araceae. of the . Structure, Evolution and Taxonomy. Springer, , New York, Tokyo. ACKNOWLEDGMENTS Deyl, M. 1955. The evolution of the I wish to thank the following persons plants and the taxonomy of the who reviewed this chapter and made monocotyledons. Sborn. Nar. Mus. v. significant contributions; Josef Bogner, Praze, Rada B, Pdr Vedy 11(6), Munich Botanic Garden; Michael H. Botanica 3:1-143. Grayum, Missouri Botanical Garden; Engler, A. 1905. Pothoideae. In A. Engler Simon Mayo, Royal Botanic Gardens; (ed.), Das Pflanzenreich IV 23B Dan H. Nicolson, Smithsonian Institu­ (Heft 21):1-330. tion. ____. 1911. Lasioideae. In A. Engler I am also grateful to Ann B. Ruger, (ed.), Das Pflanzenreich IV 23C Frances K. Mazanec, and Petra (Heft 48):1-130. Malesevich for the invaluable assistance ____. 1912. Homalomeninae und given in researching, editing, and proof­ Schismatoglottidinae. In A. Engler reading this difficult manuscript. (ed.), Das Pflanzenreich IV 23Da Finally, I thank the National Science (Heft 55):1-134. Foundation (Grant number BSR89- ____. 1915. Anubiadeae, Aglaone­ 05890) for their continued support of my mateae, Dieffenbachieae, research. 0 Zantedeschieae, Typhonodoreae, Peltandreae. In A. Engler (ed.), Das liTERATURE CITED Pflanzenreich IV 23Dc (Heft 64):1- 78. Bogner,]. 1973. Die gattung Pycnospa­ ____. 1920a. Aroideae und Pistioi­ tha Thorel ex Gagnep. (Araceae) deae. In A. Engler (ed.), Das Pflan­ Oesten: Bot. Z. 122:199-216. zenreich IV 23F (Heft 73):1-274. ___.1978 [1979]. A critical list of the ____ . 1920b. Araceae pars generalis aroid genera. Aroideana 1:63-73. et index familiae generalis. In A. ___. 1980. The genus Scaphispatha Engler (ed.), Das Pflanzenreich IV Brongn. ex Schott. Aroideana 3:4- 12. 23A (Heft 74):1-71. ____. 1980a. Berichtigungen und ____ & K. Krause. 1908. Monsteroi­ Ergazungen zu Jasarum steyer­ deae. In A. Engler (ed.), Das Pflan­ markii Bunt. und Lagenandra insig­ zen reich IV 23B CHeft 37):4-138. nis Trim. Aqua-Planta 3-80:2. ____ & . 1920. Colocasioi- ____ & D. H. Nicolson. A revised deae. In A. Engler (ed.), Das Pflan­ classification of Araceae with dichot­ zenreich IV 23E (Heft 71):1-139. omous keys. Willdenowia. (in Ertl, P. o. 1932. Vergleichende Unter­ press). suchungen ilber die Entwicklung der Cheadle, V. 1. 1942. The occurrence and Blattnervatur der Araceen. types of vessels in the various organs 126:115-248. of the plant in the Monocotyledonae. Eyde, R. H., D. H. Nicolson & P. Sherwin. Amer.] Bot. 29:441-450. 1967. A survey of floral anatomy in Croat, T B. 1981. A revision of Syngo­ Araceae. Amer. j. Bot. 54:47S-497. nium (Araceae). Ann. Missouri Bot. Fairbrothers, D. E., ]. ]. Mabry, R. 1. Card. 68:565-651. Scogin & B. 1. Turner. 1975. The Dahlgren, R. M. T & H. T Clifford. 1982. bases of angiosperm phylogeny: The Monocotyledons: A Comparative chemotaxonomy. Ann. Missouri Bot. Study. Academic Press, London. Card. 62:765-800. 60 AROIDEANA Vol. 13, No. 1-4

Fox, M. G. & J. C. French. 1988. System­ ___ & . 1981d. Vascular atic occurrence of sterols in Araceae: patterns in stems of Araceae: sub­ subfamily Colocasioideae. Amer. ]. family Philodendroideae. Bot. Gaz. Bot. 75: 132-137. (Crawfordsville) 142:550-563. ____ & . Systematic survey & . 1983. Vascular of four types of latex particles in patterns in stems of Araceae: sub­ Araceae: subfamily Colocasioideae. families Colocasioideae, Aroideae (in prep.). and Pistioideae. Amer.J Bot. 70:756- French, J. C. 1986a. Patterns of endothe­ 771. cial wall thickenings in Araceae: ___ & . 1984. Patterns of subfamilies Colocasioideae, Aroi­ Stem Vasculature in Philodendron. deae, and Pistioideae. Bot. Gaz. 147: Amer.]. Bot. 71:1432-1443. 166-179. Gibbs, R. D. 1974. Cbemotaxonomy of ____. 1986b. Ovular vasculature in flowering plants, Vol. III. McGill­ Araceae. Bot. Gaz. 147: 478-495. Queen's Univ. Press, Montreal. ____. 1986c. Patterns of stamen Grayum, M. H. 1984. Palynology and vasculature in Araceae. Amer. ]. Bot. phylogeny of the Araceae. Ph.D. 73: 434-449. Thesis, University of Massachusetts, ____. 1987a. Structure of ovular and Amherst. placental trichomes in Araceae. Bot. ____. 1987. A summary of evidence Gaz. 148: 198-208. and arguments supporting the re­ ____. 1987b. Systematic occurrence moval of Acorus from the Araceae. of a sclerotic hypodermis in roots of Taxon 36:723-729. Araceae. Amer.]. Bot. 74: 891-903. ___. 1990. Evolution and phylo­ ____ . 1987c. Systematic occurrence geny of the Araceae. Ann. Missouri of resin canals in roots of Araceae. Bot. Gard. 77:628-697. Bot. Gaz. 148: 360-371. ____ . 1988. Systematic occurrence Harborne, J. B. 1982. Flavonoid com­ of anastomosing laticifers in Ar­ pounds. In R. M. T. Dahlgren & H. T. aceae. Bot. Gaz. 149: 71-81. Clifford, The Monocotyledons: A ___. & P. B. Tomlinson. 1980. Comparative Study. Academic Press, Preliminary observations on the vas­ london, pp. 264-274. cular systems in stems of certain Harris, P. J. & R. D. Hartley. 1980. Araceae. In C. D. Brickell, D. F. Phenolic constituents of the cell Cutler & M. Gregory (eds.), Petaloid walls of monocotyledons. Biocbem. Monocotyledons. Academic Press, Syst. Bcol. 8:153-160. lDndon, pp. 105-116. Hegnauer, R. 1963. Cbemotaxonomie der ____ & . 1981a. Vascular Pflanzen. 2. Monocotyledoneae. patterns in stems of Araceae: sub­ Birkhouser, Basel. families Calloideae and Lasioideae. Hooker, J. D. 1883. Aroideae. In G. Bot. Gaz. (Crawfordsville) 142:366- Bentham & J. D. Hooker, Genera 381. Plantarum, Vol. 3. lDndon, pp. & . 1981b. Vascular 955-1000. patterns in stems of Araceae: sub­ Hotta, M. 1970. A system of the family family pothoideae. A mer. ]. Bot. Araceae in]apan and adjacent areas. 68:713-729. Mem. Fac. Sci. Kyoto Imp. Univ., Ser. & . 1981c. Vascular BioI. 4: 72-96. patterns in stems of Araceae: sub­ ____. 1981. A new genus of the family Monsteroideae. A mer. ]. Bot. family Araceae from West Sumatra. 68:1115-1129. Acta Phytotax. Geobot. 32: 142-145. THOMAS B. CROAT, 1990 61

Hutchinson, J. 1973. The families of Kew Bull. 28:199-210. jlowering plants. Clarendon Press, Metcalfe, C. R. 1967. Distribution of latex london. in the plant kingdom. Econ. Bot. 21: Jones, G. E. 1957. Chromosome numbers 115-127. and phylogenetic relationships in Nicolson, D. H. 1960. A brief review of the Araceae. Ph. D. Thesis, Univer­ classifications in the Araceae. sity of , Charlottesville. Baileya 8: 62-67. Jiissen, F. J. 1928. Die Haploidgeneration ___. 1984. Suprageneric names at­ der Araceen und ihre Verwertung fur tributable to Araceae. Taxon 33: die System. Bot. jahrb. Syst. 62: 680-690. 155-283. ___'. 1987. Derivation of aroid ge­ Krause, K. 1908. Calloideae. In A. Engler neric names. Aroideana 10:15-25. (ed. ), Das Pjlanzenreich W. 23B Petersen, G. 1989. Cytology and sys­ (Heft 37) :140-155. tematics of Araceae. Nordic J Bot. ___'. 1913. Philodendrinae. In A. 119-166. Engler (ed. ), Das Pjlanzenreich W. Raven, P. H. & D. I. Axelrod. 1974. 23Db (Heft 60): 1-143. Angiosperm biogeography and past Li, H. 1979. Angiospermae, Monocotyle­ continental movements. Ann. Mis­ doneae, Araceae, Lemnaceae. In C. souri Bot. Gard. 61:539-673. y. Wu & H. Li (editors), Flora Schott, H. 1860. Prodromus Systematis Reipublicae Popularis Sinicae, Vol. Aroidearum. Congregationis 13(2). Mechitharisticae, Vindobonae. ___. 1980. Himalayas-Hengduan Schuster, R. M. 1976. Plate tectonics and Mountains-the centre of distribu­ its bearing on the geographical ori­ tion and differentiation of the genus gin and dispersal of angiosperms. In Arisaema; to discuss the problems C. B. Beck (ed.), Origin and Early about the origin and migration of this Evolution ofAngiosperms. Columbia genus. Acta Bot. Yunnanica 2: 402- Univ. Press, New York, pp. 48-138. 416. Solereder, H. & F. J. Meyer. 1928. Sys­ Lemee, A. 1941. Aracees. In Dictionaire tematische Anatomie der Monokoty­ Descriptif et Synonymique des Gen­ ledonen. Heft 3. Principes-Synan­ res de Plantes Phanerogames 8:88- thae-Spathiflorae. Gebriider 101. Brest. Bomtraeger, Berlin. Madison, M. T. 1981. Notes on Caladium Thanikaimoni, G. 1969. Esquisse paly­ (Araceae) and its allies. Selbyana 5: nologique des Aracees. Inst. Fran~. 342-377. Pondichery, Trav. Sect. Sci. Tech. Marchant, C. J. 1970. Chromosome varia­ 5(5):1-31. tion in Araceae: I. Pothoeae to Sty­ Thome, R. F. 1976. A phylogenetic lochitoneae. Kew Bull. 24:315-322. classification of the Angiospermae. ___. 1971a. Chromosome variation Evol. Bioi. 9:35-106. in Araceae: II. Richardieae to Coloca­ ___. 1983. Proposed new realign­ sieae. Kew Bull. 25:47-56. ments in the angiosperms. Nordic]. ___. 1971b. Chromosome variation Bot. 3:85-117. in Araceae: III. Philodendreae to Tillich, H. J. 1985. Keimlingsbau und Pythonieae. Kew Bull. 25: 323-329. verwandtschaftliche Beziehungen ___. 1972. Chromosome variation der Araceae. Gleditschia 13:63-73. in Araceae: Iv. Areae. Kew Bull. 26: Williams, C. A., J. B. Harbome & S. J. 395-404. Mayo. 1981. Anthocyanin pigments ___. 1974. Chromosome variation and leaf flavonoids in the family in Araceae: V. Acoreae to Lasieae. Araceae. Phytochemistry 20:217-234. 62 AROIDEANA Vol. 13, No. 1-4

Fig. 5. Typhonium pedatisectum Gage, Fig. 6. Hapaline brownii Hook. f, Hay Philippines. Photo by unknown. 2036, Malaysia, cultivated at Kew. Photo by T B. Croat.

Fig. 7. Gorgonidium vermicidum (Speg.) Fig. 8. Bu ndra motleyana, Bogner & Nicolson, Croat 68462, Argen­ Bogner 1366, Sarawak (spathe artificiall y tina. Photo by T B. Croat. opened). Photo by J. Bogner. Fig. 9. T7.?eriopbol1u mjlscberi Sivadasan, Fig. 10. Pbymatarum horneense M. Ta milnadu, Ind ia. Photo by G. J. Thiyaga­ Horta, Bogner 1506, Sa rawak. Photo by raj. A . Tangerin i.

Fig. 11 . Culcasia striolata Eng!. Croat Fig. 12. Gonatantbus pumilus Eng!. & K. 53499, Nigeria. Photo by T B. Croat. Krause, Thailand. Photo by L. Birk.