SMITHSONIAN CONTRIBUTIONS TO 

BOTANY 

NUMBER 1 

Dan Henry NicoZson A Revision 

of the 

Genus Aglaonema 

(Araceae) 

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CITY OF WASHINGTON 

I 969 
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Contents 

Page 

Abstract ............................. 1 

Introduction ............................ 1 

Aglaonema within the Family Araceae ................. 3 

Taxonomic History of Aglaonema ................... 4 

Embryology ............................ 8 

Morphology. ........................... 11 

Plant Diseases ........................... 16 

Aglaonema Schott .......................... 17 

Dubious Binomials ......................... 59 

Excluded Species .......................... 60 

Identification List. ......................... 61 

Literature Cited .......................... 63 

Cytology ............................. 6 

ABSTRACT 

Nicolson, Dan Henry. A Revision of the Genus Aglaonema (Araceae) . Smithsonian 

Contributions to Botany, 1 : 1-66. 1969.-This revision of Aglaonema (ranging from 

northeastern India through New Guinea) is based on study of over 1500 herbarium 

specimens in 33 herbaria, supplemented by field studies and over 100 personal collections made in southeastern Asia over a period of 18 months. Much of the literature 

of the genus is reviewed, particularly on cytology, embryology, and morphology. The 

taxonomic portion of the text includes keys, synonymies, descriptions, notes on range, 

habitat, typification as well as discussion of taxonomic and nomenclatural problems. 

The sections Aglaonema and Chamaecaulon are recognized for the first time. 

Although 76 binomials have been published for the genus, only 21 are recognized 

and accepted herein as species. No new species have been recognized. 

Introduction 

Aglaonema is a genus of tropical aroids of 21 species 

native to southeastern Asia from northeastern India 

across southern China and Indonesia through New 

Guinea. All species are herbaceous. The generic name 

means “shining stamen” in Greek. 

This paper is a condensation of a doctoral dissertation accepted by Cornell University in 1964. The 

thesis is available through University Microfilms, Inc., 

of Ann Arbor, Michigan, as negative microfilm number 65-3347. Thanks are due to my special committee: 

Drs. W. J. Dress (Chairman), R. E. Lee, H. P. Banks, 

and J. F. Cornman. I owe a particular debt to Dr. 

G. H. M. Lawrence, who was chairman of my special 

committee in the initial stages. Mr. C. V. Morton of 

the Smithsonian Department of Botany also critically 

read the manuscript and many of his suggestions were 

gratefully adopted. 

I am also deeply grateful to Dean D. Umali and 

Mr. J. V. Pancho of the Philippine College of Agriculture, Mr. A. Dilmy (Bogor), Mr. J. Womersley 

(Lae), Dr. H. M. Burkill (Singapore), Mr. Tem 

Smitinand (Bangkok), and Mr. J. A. R. Anderson 

(Kuching) . These, plus others I have not mentioned, 

deserve special thanks for arranging field trips with 

and for me, as well as housing for my family. 

In my visits to herbaria Dr. C. G. G. J. van Steenis 

and the entire staff of the Rijksherbarium (Leiden) 

were especially generous, as were, also, Dr. Alicia 

Lourteig (Paris), Dr. K. Rechinger (Wien), Dr. G. 

Schultze (Berlin), Dr. W. T. Stearn (British Museum), 

Dr. C. Bonner (Geneva), Mr. L. L. Forman (Kew) , 

Dan Henry Nicolson, Department of Botany, Smithsonian 

Institution, Washington, D.C. 20560. 

Dr. A. Kostermans (Bogor), Dr. H. E. Moore 

(Ithaca) , and Dr. G. S. Bunting (Ithaca) , who took 

special interest in my work and problems. 

I thank the National Science Foundation for 

Graduate Fellowships numbers 20006 ( 1960-1961 ) , 21204 (1961-1962) and 32118 (1962-1963). Finally 

I wish to thank my wife, who never faltered in her 

enthusiasm and shouldered much of the routine work. 

study emphasized visiting the type-localities of as many 

described taxa as possible and the herbaria known to 

have major collections of tropical Asiatic plants. Typelocalities were emphasized for two reasons : to collect 

topotype material to help offset loss of type material 

destroyed during World War I1 and to obtain some 

idea of variability in type-localities. Other considerations (political situations, contacts, time, money) 

largely determined where and how long I could stay 

and study within the natural distribution of the genus 

Aglaonema. The following rCsumC indicates how many 

months were spent in each country and in what provinces (districts or divisions) collections were made: 

Sept. 60-Jan. 6 1 Philippines Agusan, Albay, Bukidnon, 

Catanduanes, Davao, 

Laguna, Leyte, Misamis 

Oriental, Negros 

Oriental, Quezon 

MATERIALS AND METHODS.-The first Stage Of this 

Dates Country Provinces 

Jan. 61-June 61 Indonesia West Java 

June 61-Aug. 61 Malaysia Johore, Penang, Perak, 

Sarawak, Selangor, 

Singapore 

Sept. 61-Nov. 61 Territory of Morobe District, BougainNew Guinea, ville, New Britain, 

Papua Manokwari 

Dec. 61-Feb. 62 Thailand Bangkok, Chanthaburi, 

Chieng Mai, Nakhon 

Nayok, Trat, Trang 

1 
2 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

The number of my collections (total=103) of 

Aglaonema by country are as follows: Indonesia, 17; 

Malaysia, 29; New Guinea, 3; Philippines, 34; Thailand, 16; U.S.A., 4. 

Duplicates (an average of 4 per collection) were 

distributed in the following order: 1, Washington 

(US) ; 2, national herbarium of county of origin; 

3, Ithaca (BH) ; 4, Leiden (L) ; 5, Kew (K) ; 6, Berlin 

(B) ; 7, Vienna (W) ; 8, Paris (P) ; 9, Manila (PNH) ; 

10, Bogor (BO) ; 11, Singapore (SING) ; 12, Calcutta 

(CAL); 13, London (BM); 14, Geneva (G); 15, 

Cambridge (A) ; 16, New York (NY) ; 17, Brisbane 

(BRI) ; 18, Bangkok (BK) . The abbreviations are the 

standard ones of Lanjouw and Stafleu ( 1964). 

I have studied the available material of Aglaonema 

in the following herbaria: 

Bogor (BO) 

Kew (K) 

Singapore (SING) 

Leiden (L) 

Paris (P) 

Calcutta (CAL) 

London (BM) 

Washington (US) 

Manila (PNH) 

New York (NY) 

Ithaca (BH) 

Bangkok (BK) 

Cambridge (A) 

Geneva (G) 

Berlin (B) 

Cambridge (GH) 

Copenhagen (C) 

Florence (FI) 

276 

242 

233 

133 

108 

73 

70 

52 

46 

42 

38 

33 

25 

22 

21 

20 

20 

19 

Kepong (KEP) 

Kuching (SAR) 

Stockholm (S) 

Utrecht (U) 

Munich (M) 

Edinburgh (E) 

Zurich (Z) 

Bangkok (BKF) 

Vienna ( W) 

Lae (LAE) 

Dehra Dun (DD) 

Brisbane (BRI) 

Brussels (BR) 

Manokwari (MAN) 

Rangoon (RANG) 

Total 

14 

12 

11 

10 

10 

7 

7 

5 

3 

3 

3 

2 

2 

0 

0 

1562 

- 

These sheets were recorded on three-by-five inch 

filing cards with the collector’s name, numbers, herbarium, determination, date of collection, locality, field 

notes, and measurements. By this method duplicate 

sheets could be recognized in different herbaria. Since 

the 1,562 sheets include 461 duplicates, the sheets 

actually represent 1,101 collections. Thus, this study 

is based on 103 personal collections and 1,101 earlier 

collections, a total of 1,204 collections, more extensive 

than the most recent monograph of Aglaonerna 

(Engler, 1915), which cited only 160 collections. 

According to the card file, Ridley collected Aglaonema the largest number of times (86). Other collectors who collected considerable numbers of Aglaonema 

were Kerr (42), Bunnemeijer (38), and Koorders 

(32). Engler probably saw most of Ridley’s and 

Koorders’ collections but none of Bunnemeijer’s and 

Kerr’s since they are more recent. 

Data were accumulated from each specimen. The 

habit of the stem was noted (erect, repent, or decumbent), its length measured or taken from field notes, 

and its largest diameter measured. Although experiments were not performed, it is my impression that the 

width of a pressed and dried stem is approximately that 

of a fresh stem since the shrinkage in drying appears to 

be matched by the increase due to flattening during 

pressing. Internode length was noted and was commonly found to be about equal to the stem thickness; 

i.e., internodes were longest on the thickest basal portion of the stem and shortest on the uppermost portion 

of the stem. Petiolar length, petiolar sheath length, 

blade length, and blade width were measured on the 

most accessible and complete leaf on a specimen in order to ensure that ratios comparing these measurements 

would be real. If other complete leaves were available, 

one or two were measured to obtain an indication of 

variability in the actual measurements and ratios between leaves on a given specimen. The petiolar sheath 

was noted as being scarious or membranous. Leaf shape 

and the shape of the leaf bases and apices were recorded. The terminology for leaf shape follows that of 

the Committee on Terminology (1962) except that I 

regard their numbers 36 and 37 (q.v.) as lanceolate 

(not narrowly ovate), and I use “broadly” when the 

margin of the leaf base or apex forms an angle between 

70” and 89” and “narrowly” when the margin of the 

angle formed is less than 45”. For terminology of leaf 

apices and bases I followed Stearn (1956) with’a few 

minor exceptions; i.e., my “obtuse” is pointed (not 

rounded) with margins diverging at more than 90”. 

The presence of variegation was noted and described. 

The relative distinctiveness of primary venation from 

secondary venation was noted and, if the primary venation was strongly or weakly differentiated, then the 

number and angle of divergence of the veins from the 

midrib was noted. The presence, location, and appearance of cataphylls were noted. The number and length 

of peduncles were noted. Width of the spathe was established by measuring the circumference with a string. 

The lengths of the spathe, spathe decurrency, stipe, 

spadix, pistillate and staminate portions of the spadix 

were measured from the most accessible side of the 

spadix, the front, with the exception of the stipe length, 

which was measured along the back of the spadix. The 

color, length, and width of the fruits were noted. 
NUMBER 1 3 

Aglaonema within the Family Araceae 

I have previously discussed (1960) the various classifications proposed for the Araceae. There are two contending classifications of the family, Hutchinson’s 

(1959) and Engler’s (1920). I have concluded that 

the latter system seems to achieve more realistic groupings of genera than the former. 

There have been two other classifications of Araceae, 

both of which are progenitors of Hutchinson’s system. 

Hooker’s (1883) system was the basis for Hutchinson’s, 

and Schott’s (1860) was the basis of Hooker’s 

treatment. 

Engler (1920) and Hutchinson (1959) divided the 

family into rather different groupings. Hutchinson used 

2 series and 18 tribes while Engler used 8 subfamilies and 31 tribes. Hutchinson placed Aglaonema 

in the tribe Richardieae (comprising 20 genera) and 

Engler placed Aglaonema in the subfamily Philodendroideae (comprising 17 genera). If we eliminate the 

eleven genera common to Engler’s Philodendroideae 

and Hutchinson’s Richardieae, the three genera that 

were reduced by Engler but were accepted by Hutchinson, and the one genus that was published subsequent 

to Engler’s work, there remain eleven problematic 

genera on which the two authors disagreed. Six of these 

genera (Amauriella, Anubias, Dieffenbachia, Peltandra, Philodendron, Typhonodorum) were included in 

Engler’s Philodendroideae but were placed in other 

tribes ( Colocasieae, Dieffenbachieae, Philodendreae) 

by Hutchinson. Five of the genera (Anchomanes, 

Montrichardia, Nephth ytis, Plesmonium, Pseudoh ydroAme) were included in Hutchinson’s Richardieae 

but were placed in the subfamily Lasioideae by Engler. 

With regard to these eleven controversial genera, I 

find Engler’s Philodendroideae more natural than Hutchinson’s Richardieae. Within the monocotyledons, the 

Araceae are rather distinctive in having reticulate as 

well as striate (“parallel”) venation. All the genera in 

Engler’s Philodendroideae have striate venation. Hutchinson’s Richardieae are mostly genera with striate 

venation, but the five controversial genera included by 

Hutchinson all have reticulate venation. Engler’s classification is followed in this present discussion. 

Although Engler (1920, p. 63) regarded one subfamily, Pothoideae, as the root of the family and giving 

rise to the other subfamilies, in practice he treated the 

family as polyphyletic, with separate and overlapping 

sequences in each subfamily. The resulting system is 

complex, and discussion of it is beyond the scope of this 

paper. The members of the Philodendroideae, however, are relatively easy to recognize, for only the 

Philodendroideae combine the characteristics of being 

terrestrial (not free-floating aquatics like Pistia) , having unisexual flowers, and having striate (or parallel) 

venation. 

Engler ( 1912) divided the subfamily Philodendroideae into seven tribes. His “systema subfamiliae” 

has so many errors (when compared to his generic 

descriptions) and is so confusing (because of contrasted leads which are not strictly mutually exclusive) 

that I prefer to present an original dichotomous key, 

retaining the order of tribes as given by Engler and 

using the characteristics emphasized in his key but 

corrected in accordance with his descriptions : 

1. Caudex epigeal. 

2. Ovules many in each locule. 

3. Stamensfree ......................... 1. PHILODENDREAE 

3. Stamens fused into synandria [or free] ................ 2. ANUBIADEAE 

4. Locule solitary. Stamens free .................. 3. ACLAONEMATEAE 

2. Ovule solitary in each locule. 

4. Locules several. Stamens fused into synandria .......... 4. DIEFFENBACHIEAE 

5. Stamens free .......................... 5. ZANTEDESCHIEAE 

5. Stamens fused into synandria. 

6. Staminodia free ....................... 6. TYPHONODOREAE 

1. Caudex hypogeal. 

6. Staminodia fused ........................ 7. PELTANDREAE 

As can be seen from this key, the tribe Aglaonemateae can be defined as having ( 1 ) unilocular ovaries, 

(2) a solitary ovule, (3) free stamens, and (4) an 

epigeal caudex. 

Engler’s tribe Aglaonemateae has only two genera, 

Aglaonema and its only closely related generic relative 

Aglaodorum. Aglaodorum Schott is monotypic and is 

endemic to Malaya, Sumatra, and Borneo. It is limited 

to tidal mudflats, where it is not easily accessible for 

collecting and has apparently been collected only about 

25 times. 

There has been some disagreement as to whether 
SMITHSONIAN CONTRIBUTIONS TO BOTANY 

No other genera of Araceae have so much in common with Aglaonema as does Aglaodorum. When 

only vegetative material is available, however, some 

Dieflenbachia species cannot be distinguished from 

Aglaonema with certainty. Although some affinity may 

be suggested by this fact, Dieffenbachia is native to the 

New World and has the stamens fused into synandria, 

four staminodia subtending each pistillate flower, and 

a 2-3-locular ovary. 

Schismatoglottis occasionally is confused with Aglaonema but is recognized easily by its deciduous upper 

spathe and spadix, sterile apex of the staminate portion of the inflorescence, and multi-ovulate ovary. 

There seem to be three separate trends of evolution within the subfamily Philodendroideae: (1) from 

many locules to incomplete (partial) locules to the 

unilocular condition (respectively represented by 

Philodendron, Homalomena, Schismatoglottis) ; (2) 

from many ovules per locule to few ovules per locule 

to one ovule per locule (represented by Philodendron, 

Zantedeschia, Dieffenbachia) ; (3) from free stamens 

to partially fused stamens to synandria (represented 

by Philodendron, Arnauriella, Peltandra) . These three trends interrelate in such a complex 

fashion that it is not possible to make a meaningful 

two-dimensional “family tree” ; nevertheless, the evidence suggests that Philodendron with its many carpels, 

many ovules, and free stamens, has the most primitive 

flowers in the subfamily. The pistillate flower in Aglaonema (with one ovule in a unilocular ovary) could 

have arisen from the Schismatoglottis type (unilocular, 

multiovulate) by reduction of ovules ; however, it 

could also have arisen from the Homalomena type 

(which has several incomplete locules, many ovules) 

by reduction in ovule and locule number. In a search 

for a possible derivation of Aglaonema, the above facts 

must be considered as well as the fact that, today, all 

three genera have the same distribution. Of the three, 

on the basis of the locular and ovular situation, Hornalomena appears to be closest to the ancestral type while 

Aglaonema and Schismatoglottis appear to be more 

specialized and derivative branches. 

Taxonomic History of Aglaonema 

Because the members of the genus Aglaonema are 

relatively inconspicuous plants of dark places in the 

southeast Asiatic rainforest, they do not appear in much 

pre-Linnaean literature. 

Aglaodorum is a distinct genus or not. A number of 

authors treated Aglaodorum under Aglaonema 

(Hooker, 1883; Ridley, 1925) ; however, specialists in 

the family have maintained Aglaodorum (Schott, 

1860; Engler, 1915 ; van Alderwerelt van Rosenburgh, 

1922). 

Both Schott (1858) and Engler (1915) separated 

Aglaodorum from Aglaonema on the basis of the supposed presence in the former of staminodia among the 

pistils. This appears to be nothing but a misinterpretation of the pressed, fruiting, type-specimen. There 

are staminodia on the lower part of the staminate portion of the inflorescence, which abuts on the single 

whorl of pistillate flowers, but none beneath or between 

the pistillate flowers. As the fruits develop, the pistils 

enlarge greatly and may squeeze themselves out of the 

original single whorl. The lower staminodia may be 

caught in this developmental distortion, as illustrated 

by Schott (1858, pl. 58) in a drawing of the type 

specimen. Engler (1915, p. 35) followed Schott’s misinterpretation, but published an illustration (ibid. 

fig. 16C) of a young inflorescence, showing the pistils 

in a single whorl without staminodia among the pistils. 

Alderwerelt van Rosenburgh (1922, p. 163) correctly 

observed : 

Since . . . I had not yet set eyes on a specimen of the species 

[Aglaodorum grifithii], I took the presence of staminodes 

as a generic character from the said authors [Schott and 

Engler]. Now I have had the opportunity to examine a living 

tpecimen.. . . this specimen agrees exactly with Engler’s fig. 

16 and has no staminodes encircling the ovaries but it has 

the lower stamens sterile, which, however, are in aspect not 

different from the fertile ones when seen from above. 

My two collections of the genus confirm van Alderwerelt van Rosenburgh‘s conclusions. 

Some differences found useful in differentiating 

these genera are tabulated below : 

Aglaonema Aglaodorum 

arrangement of pistillate several spirals one whorl 

length of peduncle to 25 cm 40-50 cm 

habitat forest tidal mudflats 

color of ripe fruits red green 

flowers (usually) 

Aglaodorum probably is closely related to Aglaonema section Chamaecaulon, since the members of 

Chamaecaulon and Aglaodorum are repent and have 

exceedingly short petiolar sheaths. Since Aglaodorum 

is so distinct from Aglaonema in appearance and habitat, it is maintained herein as a distinct genus. 
NUMBER 1 

The first author to publish on a species of Aglaonema 

was Camellus (1704), also known as “Kamel” or 

“Camel,” a Jesuit priest in the Philippines from 1688 

until his death in 1706. Camellus (1704, p. 36) described a species that he called “Dracunculus Luzonis 

primus.” According to Camellus’s unpublished drawing (Figure 23) this is a fruiting Aglaonema. No one 

has yet been able to show with certainty what species is 

represented by Camellus’s description. The problem is 

discussed herein under “Dubious Binomials.’’ 

The only other pre-Linnaean author to discuss (and 

illustrate) Aglaonema was Rumphius (1747) in his 

Herbarium Amboinense. In that work two plants 

were reported, Arum aquaticum Rumphius (1747, vol. 

5, pp. 312-313, pl. 108) and Adpendix erecta Rumphius (1747, vol 5, p. 487, pl. 182: fig. 2). Merrill 

(1917) in his Interpretation of Rumphius’s Herbarium 

Amboinense concluded that both represent the same 

species, Aglaonema marantifolium Blume, as interpreted herein. 

Linnaeus ( 1754, p. 22 ; 1736, p. 137 1 ) misidentified 

Arum aquaticum Rumphius with Arum ouatum L., 

the basionym of Lagenandra ouata (L.) Thwaites. 

This misidentification was initiated by Rumphius 

(1747, vol. 5, p. 313), who indicated that his plant 

matched a plate of Rheede’s Hortus Malabaricus, 

which later became the type of Arum ouatum L. Linnaeus apparently accepted Rumphius’s opinion without question. There is a specimen of Aglaonema simplex in the Linnaean herbarium labelled Arum ovatum 

L. Nothing further is known of this specimen (see discussion of A. simplex). 

Roxburgh (1814, p. 65) published Calla oblongifolia in a list of the plants cultivated at the Calcutta 

Botanic Gardens. In a footnote Roxburgh referred to 

Arum aquaticum Rumphius. Roxburgh‘s Calla 

oblongifolia thus was the first legitimate binomial 

applicable to the genus Aglaonema. 

Jack (1820, vol. 1, no. 1, p. 24) described a new 

species from Penang that he called Calla nitida. This 

was the second legitimate binomial applicable to the 

genus Aglaonema. 

In 1822 Link (p. 394) described Arum integrifolium, 

which he believed to be a new species, in a list of plants 

cultivated at the Berlin Botanic Gardens. This plant 

was the same Malayan species that Jack described two 

years earlier. 

5 

The recognition and publication of Aglaonema as a 

distinct genus by Schott ( 1829), who was the first specialist in the taxonomy of the Araceae, was marred 

by an unfortunate nomenclatural error. The significance of this error has been overlooked by subsequent 

botanists. Schott named the type species Aglaonema 

oblongifoliurn. There was no reference to Calla oblongifolia Roxburgh, nor is the species that was described 

the same as the Roxburghian species; however, “Arum 

integrifolium Hort. Berol.” was cited in synonymy, a 

clear reference to Arum integrifolium Link, which had 

been published seven years earlier. Thus, Aglaonema 

oblongifolium Schott (1829, under Article 63, ICBN, 

1966) is nomenclaturally superfluous and illegitimate. 

Schott later ( 1832, p. 20), perhaps realizing his error, 

renamed his type species Aglaonema integrifolium. 

In 1832 the first complete edition of Roxburgh’s 

Flora Indica was published. In that work Calla picta 

Roxburgh, a Sumatran species of Aglaonema, was 

first published with a description. 

Blume (1837, p. 153), in an important study of 

Asiatic aroids, named three new species of Aglaonema. 

The first, A. simplex, is one of the commonest species 

of the western Malesian area. The second, A. marantifolium, is the commonest species of the eastern Malesian area and included Adpendix erecta Rumphius 

(1747, 5, p. 487, pl. 182: fig. 2). Within the genus 

Aglaonema, this is the epithet available for the Moluccan species. The earlier epithet, oblongifolium, from 

Calla oblongifolia Roxburgh ( 1814), cannot be transferred to Aglaonema since the resulting combination 

becomes a later homonym of Aglaonema oblongifolium 

Schott (1829). The third species, A. maculatum, was 

regarded as dubious by Blume. It is based entirely on 

Dracunculus Luzonis primus Camellus ( 1704), which, 

as mentioned earlier, is still a problem. Blume was the 

first to point out the difference between conspicuous 

and inconspicuous primary venation in Aglaonema, a 

differentiation that has been overemphasized by most 

subsequent authors. 

Kunth (1841) deserves to be recognized as the first 

monographer of the genus Aglaonema since he was the 

first to bring together all of the previous information on 

the genus. He accepted seven species, publishing the 

new combinations A. oblongifolium (Roxburgh) 

Kunth (not Schott, 1829), A. pictum (Roxburgh) 

Kunth, and A. nitidum (Jack) Kunth. 
6 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

By this time, Aglaonema was relatively well accepted 

as a genus and several species were in cultivation as 

ornamental foliage plants. In the following years many 

new species were proposed, frequently from cultivated 

material. A number of authors made major contributions to the knowledge of the genus. 

Schott, in the late 1850s and early 1860s, described 

a number of new species. His most important treatment 

of the genus is in Prodromus Systematis Aroidearum 

( 1860), which recognizes eleven species plus two dubious species, an enlargement of Kunth’s monograph of 

1841. 

Engler (1879), in his first monograph of the family 

Araceae, recapitulated Schott’s (1860) work on 

Aglaonema, recognizing eleven species. 

N. E. Brown, in the late 1880s and early 1890s, described six new species of Aglaonema from cultivated 

plants. It is apparent from N. E. Brown’s autograph 

manuscripts at Kew that he collaborated closely with 

Hooker on Ag1aonem.a in the following work. 

Hooker ( 1893), in his Flora of British India, recognized 13 species, of which 8 were thought to be new. 

This work is important as it covers the western area of 

the distribution of Aglaonema. 

Engler’s (1915) monograph of Aglaonema, which 

is part of his second monograph of the family Araceae, 

is unquestionably the most important taxonomic work 

on the genus. Forty-one species plus two dubious species 

were recognized, of which twelve were considered to be 

new. The criticism one might make of this monograph 

is that no consistent effort was made to clarify relationships. Apparently strong differentiating criteria were 

not found. The need for valid differentiating characters 

and the lack of information sufficient for postulating 

relationships within the genus led to the present study. 

Aldenverelt van Rosenburgh (1922) described ten 

Indonesian species that he considered to be new. None 

of them is accepted in this work since they fall within 

the range of variation of previously established species. 

Ridley (1925, p. 99) discussed the Malayan species 

of Aglaonema. Gagnepain (1942) discussed the IndoChinese species of Aglaonema and accepted 17 species. 

The Index Kewensis lists 76 specific epithets that 

have been published in the genus Aglaonema. The 

most recent monograph (Engler, 1915) accepted 43 

species and 15 have since been proposed, making a 

total of 58 more or less generally accepted species. 

In this work only 2 1 species are accepted. 

Cytology 

A number of major surveys have been made on 

chromosome numbers in Araceae (Ito, 1942; Malvesin-Fabre, 1945 [ex Delay, 19511 ; Mookerjea, 1955; 

Jones, 1957; Pfitzer, 1957). The last mentioned paper 

was the first to be largely founded on meiotic counts. 

There is a great deal of evidence that, as Mookerjea 

(1955) has said, “Specification is affected [sic] 

through karyotypic changes in the soma and through 

their entrance into the growing point.” Tremendous 

variation in chromosome number in somatic tissue 

has been reported several times; for instance, Sharma 

and Datta (1961 ) found eleven different chromosome 

complements in Aglaonema costatum ‘Foxi’ ( =A. 

costatum f. immaculatum) and were obliged to do a 

statistical analysis to decide what was the “normal 

complement.” 

The question of a primitive number for the family 

is far from settled. Gaiser (1927) apparently felt that 

eight was basic. Mookerjea (1955) favored seven. 

Jones (195,7) found indications that six and seven 

were basic. Pfitzer (1957) was impressed by consecutive sequences of numbers within genera and the frequent occurrence of prime numbers ( 11, 13, 17, 19). 

The lowest number reported for the family is 2n= 14 

(Huttleston, 1953, p. 174) in Pistia stratiotes. Gow 

(1907, 1908, 1913) reported n=8 or 2n=16 in 

Aglaonema, Diefenbachia, Symplocarpus and Xanthosoma while Sharma and Mukhopadhyay (1965) 

reported 2n=16, 18 and 26 (n=8, 9 and 13) in 

Typhonium. Jussen (1928), Dudley (1937), and 

Banerji (1947) reported that n=9 in Spathiphyllurn, 

Acorus, and Typhonium, but these counts have not 

all been confirmed by subsequent studies. Vignoli 

(1939) reported n=ll in Ambrosinia. This review 

indicates that low numbers (less than 12) are not 

common in the family; therefore, establishing a low 

basic number for the entire family may be difficult. 

The following list shows the chromosome numbers 

of all genera of the Philodendroideae that have been 

reported in the literature. The known or presumed 

(from somatic material) meiotic numbers are given. 

Only the most frequently reported number is used 

and no attempt is made to include all the various 

complements reported for each genus. The variation 

in number indicates the difficulty of determining a 

basic number for the subfamily. As may be seen, com- 
NUMBER 1 7 

pared with other genera, Aglaonema does not have a 

particularly high number of chromosomes: 

Genera 

Zantedeschia 

Dieffenbachia 

Homalomena 

Philodendron 

Aglaodorum 

Aglaonema 

Anubias 

Schismatoglottis 

Peltandra 

n 

16 

17 

17* 

17 and 18 

18* 

20 

24 

28 * 

56 

References 

Jones, 1957 

Pfitzer, 1957 

Mookerjea, 1955 

Pfitzer, 1957 

Earl, 1956 

Pfitzer, 1957 

Pfitzer, 1957 

Mookerjea, 1955 

Goldberg, 1941 

*Meiotic number was deduced from a mitotic determination. 

By morphological criteria, Aglaodorum is most closely related to Aglaonema. There is some confusion, 

however, over the chromosome number of Aglaodorum. Earl (1956) published a report that Aglaodorum 

(Aglaonema) grifithii had 36 chromosomes. Later, 

she (Jones [nee Earl], 1957) gave more details, stating 

that the chromosome number was 34 but that several 

cells were seen with 36. It would be useful to 

have confirmatory studies to determine the “normal” 

complement. 

Table 1 summarizes the information available concerning chromosome numbers within the genus Aglaonema. The reports are arranged in order of the 

chromosome numbers. 

Ignoring a few deviations, it is reasonably clear that, 

as Pfitzer ( 1957) indicated, “triploid, tetraploid, and 

TABLE 1 .-Chromosome numbers 

n 

hexaploid forms are derived from the basic number of 

20.” Pfitzer’s conclusion was supported by his demonstration of meiosis of a triploid (2n=60) showing 

20 bivalents at the metaphase equatorial plate and 20 

univalents being distributed at random (with 16 to one 

pole) (Pfitzer, 1957, p. 441, fig. 1). Pfitzer also demonstrated that the hexaploid was apomictic and that the 

triploids were sterile, the pollen in both cases aborting 

during meiosis or soon afterward. 

GOW’S (1908) report of n=8 for Aglaonema versicolor (=A. pictum) seems improbable in light of 

subsequent counts in the same species. 

Sharma and Datta (1961) found chromosome complements of 10, 13, 27, 38, and 40 in Aglaonema 

hospitum (=A. brevispathum f. hospitum). Accessory chromosome fragments were found in all complements except 40. Cells with the “normal” complement 

(i.e., most frequent) of 38 had from two to six accessory chromosomal fragments. This suggests that 38 

was probably derived from 40 by fragmentation. 

In Aglaonema costatum, Sharma and Datta (1961) 

found chromosome complements of 14,22, and 40. The 

number most frequently observed was 40, and that was 

termed the “normal” complement. No accessory fragments of chromosomes were observed in cells with the 

“normal” complement. 

In Aglaonema costatum ‘Foxi‘ Sharma and Datta 

( 1961) found eleven different chromosome complements: 16, 21, 25, 29, 32, 34, 36, 40, 44, 48, and 56. 

within the genus Aglaonema 

2n References 

versicolor (=picturn) 

hospitum (= brevispathum) 

hospitum (= breuispathum) 

angustifolium (=simplex) 

pictum 

pictum 

costatum 

costatum 

cortatum ‘Foxi’ 

modestum 

simplex (= modestum?) 

simplex (=modesturn?) 

hatnkei (=densinerLium?) 

roebdinii ( =crispurn) 

roebelinii hyb. (=crispurn) 

treubii (=cornmutaturn) 

commutatum 

commutatum 

8 

38*( 10-40) 

20 

20 

20 

40 

20 

40*( 14-22) 

56 

48*( 16-56) 

30 

60 

60 

60 

’ ca. 30 

40 

119*( 118-120) 

120 

Gow, 1908 

Sharma and Datta, 1961 

Pfitzer, 1957 

Pfitzer, 1957 

Pfitzer, 1957 

Asana and Sutaria, 1939 

Pfitzer, 1957 

Sharma and Datta, 1961 

Sharma and Datta, 1961 

Wang, 1939 

Pfitzer, 1957 

Earl, 1956 

Earl, 1956 

Pfitzer, 1957 

Pfitzer, 1957 

Pfitzer, 1957 

Pfitzer, 1957 

Earl, 1956 

* “Normal” complement, i.e., number most frequently observed; numbers in parentheses are 

extremes of variant complements also observed. 
8 

The number most frequently encountered was 48, and 

this was termed the “normal” complement. Even in 

cells with the “normal” complement there were up to 

eight chromosomal fragments. The detailed karyotypic 

analysis given by Sharma and Datta (1961) does not 

clarify the origin of this number but the number clearly 

is unstable. 

Wang’s (1939) report of 2n=56 in Aglaonema 

modestum is probably derived from a full triploid complement (2n=60). Such a variation is possible according to the variation reported in other species by Sharma 

and Datta (1961). 

Pfitzer (1957) made root tip counts on seven plants 

of the hexaploid A. commutatum and found that one 

plant had 118 chromosomes, four had 119, one had 

120, and one had both 118 and 120. Such variability 

in high polyploids is not unusual. The only peculiarity 

was that the distinctive satellite (SAT) chromosomes 

were only four and sometimes five but never six, as 

would be expected in a hexaploid. Jones (1957, p. 56) 

also mentioned finding four of these chromosomes in 

A. commutatum. 

SuMMARY.-There is not yet enough information to 

be definitive but 20 seems to be established as the basic 

number for Aglaonema; however, only cultivated material has been studied, of which the majority are 

variegated and probably anomalous or derived elements. It would be valuable to have counts from wild 

material of Aglaonema simplex, A. nebulosum, A. 

nitidum, A. hookerianum, and A. marantifolium since 

these species cover the basic range of the genus. It 

would also be useful to have counts from wild, nonvariegated Aglaonema costatum, A. breuispathum, and 

A. pictum to indicate whether or not the variegated 

material-some of which has long been in cultivation 

and propagated vegetatively-has diverged. It may 

be that polyploidy exists only in cultivation. It may be 

that numbers lower than 20 exist in the wild. At any 

rate, the demonstration that several of our cultivated 

plants are sterile polyploids casts doubt on their status 

as “good species.” 

Embryology 

MICROSPOROGENESIS.-~ have found nothing in the 

literature on microsporogenesis in Aglaonema. Threenucleate pollen grains have been reported in Aglaonema pictum by Campbell (1903, p. 677). Th’ is was 

confirmed in Aglaonema pictum and A. costatum by 

Jiissen (1928, p. 239). According to Schnarf (1931, 

SMITHSONIAN CONTRIBUTIONS TO BOTANY 

p. 281 ) , 3-nucleate pollen is common in aroids with 

unisexual flowers while 2-nucleate pollen is common in 

aroids with bisexual flowers. 

Although microsporogenesis has not been demonstrated in Aglaonema, most aroid genera that have been 

studied share the following characteristics : ( 1 ) the tapeta1 cells have amoeboid protoplasts that coalesce into 

the periplasmodium; (2) reduction division of the 

microsporocytes is of the successive type (Schnarf, 

1931, p. 281). 

MEGASPOROCENESIS.-Apparently no one has yet 

observed reduction division of the megasporocyte in 

Aglaonema. In A. commutatum Schott, Campbell 

(1912, p. 102) noted a “group of cells occupying the 

greater part of the nucellus . . . . These cells probably are to be considered as megaspores.” In Aglaonema simplex and A. modestum, Campbell reported that 

the “primary sporogenous cell” (megasporocyte?) 

usually develops directly into the embryo sac (generative apospory?) . 

bryo sac development is accompanied by destruction 

of the nucellus although a micropylar nucellar cap 

almost always remains (Schnarf, 1931, p. 282). 

Embryo sac development in most aroids studied is 

of the Polygonum type (i.e., the normal type) leading 

to an 8-nucleate polarized embryo sac. In Aglaonema 

commutatum, Campbell (1903, p. 683) commented 

on a number of irregularities of embryo sac development: ( 1 ) “a varying number of embryo sacs” [a tendency toward polyembryony]; (2) “in some cases it 

looks as if these [embryo sacs] originated independently 

from hypodermal cells [somatic apospory]”; (3) “extraordinary variation in the number of nuclei in the 

embryo sac . . . from four to twelve”; (4) “multiple 

nuclear fusions.” 

Pfitzer ( 1957, p. 442) presented strong evidence that 

Aglaonema commutatum is apomictic. He found that 

A. commutatum was hexaploid (2n= 118 to 120), had 

all the pollen degenerating in meiosis and yet that fruiting was abundant. 

Embryo sac development in certain other species 

has been reported. The first two nuclear divisions are 

normal in Aglaonema simplex and A. modestum, but 

Campbell (1912, p. 105) could not definitely establish 

the further sequence of events. He said: 

Stages between the four-nucleate condition and that in 

which the egg apparatus was completely developed were not 

found, but is practically certain that only one of the upper 

[micropylar] pair of nuclei divides, giving rise to the synerEMBRYO SAC DEVELOPMENT.-In most aroids, em- 
NUMBER 1 

gidae, while the other remains undivided and becomes the 

nucleus of the egg, there being no polar nucleus present. Thus 

at the time when the egg apparatus seems to be fully developed 

there are only five nuclei in the embryo-sac, instead of the 

eight usually found in angiosperms. 

Other workers (Michell, 1916; Jussen, 1928) found 

a preponderance of 5-nucleate embryo sacs in their 

sections but were able to demonstrate that the 8-nucleate stage was of very short duration and that the 

three antipodal nuclei immediately degenerated and 

easily became confused with the chalaza1 nucellar 

remnants. Both of these workers concluded that Aglaonema did have an 8-nucleate embryo sac. Campbell 

had once (1903, p. 677) described Aglaonema pictum 

as having a typical 8-nucleate embryo sac, but later 

(1912, p. 105) he changed his mind and, speaking of 

what he had formerly called “a typical group of three 

antipodal cells,” he said that, “as the adjacent cells of 

the nucellus are quite like the antipodal cells in appearance, it may be that these ‘antipodal’ cells did not 

really belong in the embryo-sac.” 

The facts that (1) Campbell did not observe the 

peculiar origin of the 5-nucleate embryo sac which he 

described, (2) all authors agree that the antipodal 

cells resemble the nucellar remnants, (3) the 8-nucleate stage is very short in all aroids in which it has 

been definitely established, and (4) the 5-nucleate 

stage is of long duration (the three antipodals having 

degenerated) suggests that the observations of Michell 

and of Jussen were more accurate than those of Campbell and that members of Aglaonema normally have 

the usual 8-nucleate embryo sac. 

Gow (1908, p. 40) reported in Aglaonema pictum 

(A. versicolor) that the final stages of the embryo sac 

had from two to eleven antipodals and that they might 

possibly be even more numerous sometimes. The same 

author (1913, p. 127) reported that, in Aglaonema 

nitidum, ‘‘in the mature embryo sac there are ten cells, 

five of which occupy the position of antipodals, the 

other five performing the usual functions.” Perhaps 

this author has confused the haustorial basal apparatus, 

which is developed by the endosperm nucleus in 

Araceae, with one developed by the antipodals, as in 

other families. 

In most Araceae, after fertilization, the endosperm 

nucleus sets up a basal apparatus, (this will be discussed below) ; however, Gow (1908, 1913) does not 

appear to be speaking of a mature embryo sac but of a 

fertilized one. Two points should be made: (1) Gow 

(1908, pl. 6: fig. 38) shows a rather disorganized 

9 

embryo sac at the time of fertilization, wherein the 

antipodals are two and enucleate; (2) in other aroids 

the antipodals are ephemeral. It is possible that what 

Gow considered to be a “mature embryo sac” was 

actually a fertilized embryo sac with development of 

the endospermous basal apparatus already under way. 

Another explanation for GOW’S basal cells is possible. Buell (1938) reported that, in Acorus calamus, 

after degeneration of the antipodals and fusion of the 

two polar nuclei, the fusion nucleus moved to the base 

and divided, before fertilization. 

Summary: The exact nature and the sequence oi 

events leading to the mature embryo sac are poorly 

understood and what evidence has been reported is 

often conflicting. It appears likely, however, that 

Aglaonema, excepting its apomictic members, has a 

normal type of embryo sac development but that the 

antipodals quickly degenerate, leaving a 5-nuc!eate 

stage which is quite stable. 

FERTILIZATION.-III his studies on Aglaonema commutatum, A. modestum, A. simplex, and A. pictum, 

Campbell (1912, p. 107) did not obtain 

a satisfactory demonstration of the penetration of the pollen 

tube or the fertilization of the egg. Indeed, one is almost 

forced to the conclusion that the development of the embryo 

and endosperm goes on without fertilization. For some time 

after endosperm formation begins the egg apparatus remains 

apparently unchanged, and there is no sign of a degeneration 

of the synergidae such as usually occurs in cases of normai 

fertilization. 

Gow (1908, p. 40, fig. 38) described and illustrated 

in Aglaonema uersicolor ( =A. pictum) an “8-nucleate 

sac and two male cells.” It is difficult to tell from GOW’S 

writing and discussion what he considered to be cells 

and nuclei. For instance, it is not likely that a “male 

cell” would be found inside the embryo sac. Also, cell 

wall formation generally cuts off the nuclei of the 

embryo sac at a fairly early stage. At any rate, Gow 

does seem to be describing fertilization. 

ENDosPERM.-There is a great deal of conflicting 

and irreconcilable information on this subject, not only 

for Aglaonema but also for the rest of the family. Confirmatory studies are badly needed. Maheswari and 

Khanna (1956) conclude that, although there are 

several reports of the free-nuclear endosperm formation type, none of them is authenticated. Most aroids, 

up until the last twenty-five years, have been considered 

as having the helobial type of endosperm formation 

(Jussen, 1928; Schnarf, 1931) ; however, Maheshwari 

and Khanna ( 1956), after reviewing the literature on 
10 

this subject and their own findings, said, “We feel that 

on reinvestigation many reports of Helobial type may 

turn out to be of the Cellular type.” 

Campbell ( 1912, p. 107) reported that in Aglaonema 

pictum the polar nuclei fused, but he did not study 

further development ; however, in Aglaonema simplex 

and probably also in A. modestum, and A. commutatum 

there was no fusion of the polar nuclei, according to 

Campbell (1912, p. 107). These two nuclei remained 

separate and each divided, making four free nuclei, 

after which cell wall formation began. In other cases, 

in A. simplex, discoid groups of flat cells were observed. 

“These are bounded by delicate but evident cell walls 

and in the undivided cavity of the sac may be seen 

several free nuclei between which walls are beginning 

to form (Campbell, 1912, p. 107) .” 

Gow ( 1908, p. 40) reported that in Aglaonema versicolor (A. pictum) endosperm formation is of the cellular type. In A. nitidum, Gow ( 1913, p. 128) reported 

that the endosperm nucleus gives rise to a heavy-walled 

endosperm developing simultaneously with the embryo. 

Campbell (1912, p. 108) reported that, in Aglaonema simplex, the first formation of endosperm is at 

the chalazal end and proceeds upward, filling the sac 

with a few large cells. The smaller endosperm cells 

at the chalazal end look like antipodals. 

In view of the work of Maheshwari and Khanna 

(1956), it is probable that Campbell was correct in 

believing that the first, chalazal, endosperm cells are 

small and look like antipodals. Gow apparently regarded these as antipodals; however, the nuclear type 

of endosperm formation described by Campbell seems 

very peculiar and it may be that Gow’s statement that 

the endosperm formation is cellular is correct. Obviously, confirmatory studies are needed. 

EMBRYo.-campbell (1912, pp. 109-1 12) is the 

only worker who has reported on the development of 

the embryo. The following summary is from his findings on Aglaonema modestum. Since differentiation of 

the organs takes place only after the embryo has 

reached a rather large size, the study of the young embryo does not throw light on the origin of the organs 

of the young plant. 

A large basal cell is sometimes observed and this 

may be considered as a suspensor. The upper part of 

the embryo is composed of an oval mass of similar 

cells. In other cases the embryo is isolated by the 

endosperm and no suspensor is differentiated. Growth 

of the embryo is very rapid and it encroaches on the 

SMITHSONIAN CONTRIBUTIONS TO BOTANY 

endosperm until the latter finally is almost entirely 

destroyed by the embryo, which almost fills the embryo sac cavity. 

The embryo becomes much elongated and the 

micropylar end becomes the apex of the first root while 

the other end develops into a large cotyledon with a 

single vascular bundle. The root remains very short, 

the cotyledon forming much the greater part of the 

embryo. The sheathing base of the cotyledon conceals 

the stem apex. A second leaf is developed before the 

seed ripens. 

As in most Araceae, the basal part of the ovule becomes very massive, so that a section of the seed shows 

that the embryo sac, occupied by the embryo, is only 

about one-half the bulk of the seed; the remainder is 

made up of what Campbell termed “perisperm, comparable to that in Piperaceae or Cannaceae, but not 

developed from the nucellus but from the chalazal 

region of the ovule.” 

Gatin (1921, pp. 154-155) illustrated the condition 

of the embryo in the seed at the time of germination 

in Aglaonema nitidum. Within the embryo is a simple 

vascularization of two irregular circles of strands. In 

the hypocotyl the vascular bundles form an almost 

continuous ring. At the micropylar end the embryo 

develops a number of small scales that are suberized. 

The entire embryo is covered by a secondary layer of 

periderm and under that layer are cells with chlorophyll. Outside the embryo are two teguments. The 

teguments (“perisperm” of Campbell) and the embryo make up the seed. Outside the seed is a fleshy 

pericarp that finally becomes red. 

GERMINATION.-In Aglaonema, according to Gatin’s 

( 1921) description and illustration, the germination 

is hypogeal, i.e., the seed coat containing the cotyledon remains below ground. Engler ( 191 2, figs. 1B-D) 

illustrates epigeal germination in Philodendron with 

the cotyledon aerial and leaflike with the seed coat 

borne at the tip of the aerial cotyledon. According to 

Eames (1961, p. 320), epigeal germination (as in 

Philodendron) is more primitive than hypogeal germination (as in Aglaonema) . 

Engler (1920, p. 37) does n’ot emphasize these two 

types of germination but places great importance on a 

characteristic that may be correlated, i.e., whether 

the mature seed contains endosperm or not. According 

to Engler, in the germination of endospermous seeds a 

foliage leaf immediately follows the cotyledon and the 

cotyledon itself may even be foliate. In the germination 
NUMBER 1 

of endospermless (exalbuminose) seeds, the reduced 

cotyledon is followed by a number of cataphylls before 

a foliage leaf is produced. Aglaonema fits Engler’s second category since, as Campbell ( 1912, p. 110) demonstrated, endosperm is consumed by the developing 

embryo and, as Gatin (1921, pl. 6: figs. 8-10) showed, 

the cotyledon is followed by a number of scale-like 

cataphylls before a foliage leaf is produced. 

Morphology 

Most species of Aglaonema are erect, with relatively 

unbranched stems and generally with distinct internodes. Growth is continuous but the lower leaves fall 

off, leaving the plant with the leaves tufted on the 

upper part of a stem covered below by circular leaf 

scars. The erect habit is rather unusual in the family 

and is also characteristic for most species of the genera 

Dieffenbachia (Philodendroideae) and Montrichardia 

(Lasioideae) . The erect habit, however, is found in 

occasional members of the genera Homalomena, Schismatoglottis, Philodendron (Philodendroideae) , and 

Alocasia (Colocasioideae) . 

Two species, Aglaonema brevispathum and A. 

costatum, are repent, with branched stems and relatively short internodes. The stems of wild plants of 

these species are typically found beneath the humus on 

the forest floor. In cultivation the stems creep on the 

soil surface. The stems are actually epigeal and only 

appear to be hypogeal in the wild because they are 

buried by leaf fall. The creeping or climbing habit 

is typical in most other genera of the Philodendroideae, 

except in Zantedeschia, Peltandra, and Typhonodorum, where the stem is truly hypogeal and functions 

as a storage organ. Thus, these two species of Aglaonema that form the section Chamaecaulon, are intermediate in habit between the typically erect species 

of Aglaonema and the rest of the typically repent or 

or climbing Philodendroideae. 

Several plants of Aglaonema have been observed by 

the author and described in the literature (Engler, 

1915, p. 10) as decumbent. Aglaonema rotundurn has 

been observed to produce many roots along the lower 

part of the erect stem. These roots appear to be contractile (like those described in other aroids [Holm, 

1891 ; Rimbach, 1900; Rosendahl, 1916]), pulling the 

lower stem close to the ground into a decumbent position. Larger species like A. marantifolium rarely stand 

erect over five feet although the stem may be up to ten 

feet long. The stem is not strong enough to support the 

11 

weight of the plant body and the lower stem is forced 

to lie against the ground in a decumbent position. 

Van Tieghem (1867, pp. 122-124, pl. 3: fig. 10) discussed the anatomy of the stem from the epidermis to 

the center. He reported that inside the epidermis and 

one or two tabular, colorless layers (“couches incolores tabulaires” ), there is a continuous parenchyma 

from the periphery to the center of the stem in 

Aglaonema marantifolium; inside a fairly large thickness of stem without any vascular bundles is a ring of 

small bundles with some phloem and one or two vessels (“vaisseaux”) ; inside this ring of vascular bundles 

is a ring of about 20 schizogenous canals, each of 

which is about 0.24 mm in diameter and is full of 

gummy sap (the cells bordering the canals are much 

smaller and have more irregular shapes than those of 

the surrounding parenchyma). Just inside the ring of 

canals and alternating with the canals is another rim 

of vascular bundles. These bundles are simple (like the 

outer ones) but the remaining bundles toward the center of the stem are found to be compound (“faisceaux 

composW) , i.e., with several groups (separated by 

phloem) of two or three vessels each. (According to 

Van Tieghem, it is the separation and turning out of 

these small groups of vessels, accompanied by phloem 

and two laticifers, that give rise to the outer, simple 

vascular bundles which immediately or eventually run 

out into the lateral organs.) 

The stem of Agluonema simplex was found to be 

almost the same except that it lacked the schizogenous 

canals (Van Tieghem, 1867). 

BRANcHmG.-In Aglaonema, as in most Araceae, 

bud primordia are formed in the axils of foliar organs. 

These buds appear to be “reserve” buds that, if separated from the mother-plant with a piece of stem, can 

grow into new plants or, if they remain attached, typically develop only after suppression of the terminal 

shoot (Engler, 1920, p. 12). 

I have observed in a number of plants of Aglaonema 

and other genera of the family that axillary buds do 

develop while attached to the mother-plant if the subtending node is strongly rooted in the ground. The 

creeping stems of the section Chamaecaulon are 

tightly rooted to the ground and typically are well 

branched. 

In general, erect plants of Aglaonema appear to be 

unbranched ; however, all Agluonema, like all other 

Araceae (except for three rather primitive genera, 

Pothos, Pothoidium, and Heteropsis) are supposed to 
12 

have sympodial growth after production of the first 

inflorescence (Engler, 1920, p. 1 1 ) . Engler (1912, pp. 3-5) recognizes two kinds of 

shoots in a sympodium, the lateral continuation shoot 

(“Fortsetzungsprosse” ) and the terminal reproductive 

shoot ( “Vermehrungsprosse” ) . Study of living material of Aglaonerna cornmutaturn (Figure l), A. 

picturn, and A. modesturn indicates that in Aglaonema 

the situation may not be exactly as described by Engler. 

On the mature stem below the inflorescence, the base 

of each leaf clasps and encircles the stem; however, 

when an inflorescence is produced, it is subtended by a 

leaf which seems to clasp only the inflorescence, but 

not the entire stem (Figure 1). This fact suggests that 

simple monopodial growth is not occurring and that 

the. inflorescence is actually terminal. The continuation of the vegetative shoot might then be considered 

the “lateral continuation shoot” ; i.e., the development 

of the axillary bud from the axil of the leaf one node 

lower than, and on the side of the stem opposite to, the 

leaf subtending the inflorescence. This lower leaf, however, was found to have not only the “lateral continuaSMITHSONIAN CONTRIBUTIONS TO BOTANY 

tion shoot” but also an axillary bud (Figure 1). 

Ontogenetic studies are necessary to any attempt to 

establish not only the exact sequence of events but 

also which buds are terminal, which are lateral or 

adventitious, and what happens to each bud. Without 

such study it is not possible to give an adequate 

interpretation. 

CATAPHYLLS AND LEAVES.-According to Engler 

(1920, p. 17) and my own observations, the cataphyll 

of Araceae corresponds to the petiolar sheath (vagina) 

of a foliage leaf. The cataphyll serves the same function 

as the petiolar sheath, protecting the subtended shoot 

during ontogeny. It occasionally ends in a little tip 

that looks like the rudiment of a leaf-blade. Ertl (1932, 

p. 120) supported the concept of the aroid cataphyll 

as homologous with the petiolar sheath when he described the development of the leaf in Aglaonema commutaturn: The leaf-bud clasps the subtended shoots 

and first develops into a thin, cataphyll-like structure; 

later, the petiole is formed by thickening growth along 

the longitudinal axis and the leaf-blade by growth in 

breadth at the upper end; the margins of the thin, 

cataphyll-like structure are left as the petiolar sheath 

on the petiole. It appears that cataphylls originate from 

the same kind of primordia as do the leave? but that 

differential growth does not occur in the primordia of 

cataphylls. Gluck (1919, p. 236) stated that we must 

interpret the cataphylls (“Vorblatter” ) of Araceae, as 

well as the cataphylls (“Niederblatter”) of many 

monocotyledons as bladeless, petiolar sheaths (“Blattscheiden”) . 

In Aglaonema the location of the cataphyll is sometimes of diagnostic importance. In all species of the 

genus cataphylls are found between the inflorescences 

and sometimes at the base of the reproductive shoot J 

however, as was mentioned in the section on stems, 

species of the section Charnaecaulon have one or more 

cataphylls subtending the petioles, as well as those subtending the inflorescences. Thus, in the section 

Chamaecaulon cataphylls are found between the leaves. 

This does not happen in other Aglaonema species. 

The cataphylls in the genus Aglaonema are generally 

cream colored or very light green. They soon dry to a 

brown color and disappear, leaving only a scar. 

The petioles of most aroids clasp the stem and are 

grooved longitudinally on the adaxial side at the base. 

The margins of this groove are generally broad and 

run up on the petiole in what is called a va,‘ oina or 

petiolar sheath. The petiolar sheath provides many 

Peduncle without a 

Cataphyll subten 

FIGURE 1.--Stylized lateral view of a flowering stem of 

Aglaonema cornmutaturn Schott. 
NUMBER I 13 

The relative distinctiveness of the primary veins 

has been considered to be of taxonomic value from the 

time that Blume (1837) described Aglaonema simplex as “protuberati-venosis” and A. marantifolium as 

“obselete parallelo-venosis.” Engler ( 1915, p. 10) used 

distinctiveness of venation in the opening lead of his key 

to the genus, comparing “Foliorum nervi laterales I. 

et 11. subaequales, tenues, densi, subparalleli” with 

“Foliorum nervi laterales I. quam nervi 11. & validiores.” In the latter lead, however, one finds half of 

the species recognized by Engler under a further subdivision, “Nervi laterales I. quam nervi 11. paullum 

validiores,” suggesting that this is not an absolute differentiating character. The possibility has been borne 

out in this study: even in the same species (e.g., A. 

marantifolium or A. nitidum) some plants have been 

found to have venation scarcely differentiated and 

others to have strongly differentiated primary venation. 

The angle of divergence of the primary lateral veins 

has been found to be more or less correlated with the 

width of the leaves. Narrow leaves tend to have smaller 

angles of divergence and broad leaves tend to have 

large angles (cf. A. simplex) ; however, some plants 

(e.g., A. commutatum) show unusually wide angles 

of divergence (almost 90’) of the primary veins for 

the width of the leaves, which are not particularly 

broad. 

Ertl (1932, p. 120-5) described the ontogeny of 

the venation in Aglaonema commutatum in detail. 

Early in ontogeny three vascular bundles are laid 

down. The center #bundle runs to the leaf tip in a 

straight line; the two outer bundles also run all the 

way to the apex, but about half way up the leaf 

they arch out and later turn back to the leaf apex in a 

gentle curve. Soon afterward, the third, fourth, and 

fifth pairs of bundles are successively laid down outside the first pair, arching out progressively lower on 

the leaf blade. All these veins run to the apex and are 

called “primary lateral veins.” 

Secondary lateral veins develop between the primaries and are distinctive since they turn out and 

anastomose with the next outermost primary vein and 

do not go all the way to the apex. 

Tertiary veins develop between the primary and 

secondary veins and are similar to the secondary veins 

except that they turn out and anastomose with the next 

outermost vein much further from the leaf apex and 

margin than do the secondaries. 

variations in length, attachment, and appearance that 

sometimes are useful to the taxonomist. In Aglaonema, 

however, the characteristics of the petiolar sheath may 

vary greatly within a given plant; lower leaves, for 

instance, tend to have longer petiolar sheaths than the 

upper leaves. 

Aglaonema rotundum has been observed by me to 

have the upper end of the petiolar sheath free and 

ligulate, but only for a matter of millimeters. Aglaonema crispum and A. nitidum commonly have 

chartaceous or scarious margins of the sheath that 

appear to be rather saw-toothed. Aglaonema pictuiv 

has a sheath that is more than one-half the length of 

the petiole while the closely related A. nebulosum and 

A. uittatum have sheaths that are much less than onehalf the length of the petiole. In the section Chamaecaulon the sheath is exceedingly short, normally only 

long enough to clasp the stem. 

Ertl (1932, pp. 124-5) described the vascularization in the petiole of Aglaonema commutatum. In a 

cross-section of the petiole, the vascular bundles are 

arranged approximately in three semicircles. The outer 

semicircle is made up of small bundles alternating with 

strands of collenchyma. The second semicircle is made 

up of larger vascular bundles that will be the primary 

longitudinal bundles of the leaf-blade, described below. 

The third, innermost semicircle consists of four to five 

larger bundles that branch above and form veins of 

higher order in the leaf-blade. All these bundles run 

parallel to one another through the petiole and into 

the midrib, with only a few branches and cross connections. In the leaf-blade these veins successively arch 

out, causing the midrib to diminish toward the leaf tip 

until only the medial vascular bundle remains, which 

goes all the way to the leaf tip. 

The leaf-blades of Aglaonema are generally narrowly 

oblong to narrowly elliptic and commonly have an 

acute to acuminate apex and a rounded to obtuse leaf 

base. Occasional specimens have more or less cordate leaf bases. 

The venation of Aglaonema is essentially parallel 

or striate. This type of venation, although common in 

monocotyledons, is only found in a few Araceae (all 

Philodendroideae and many Monsteroideae) . 

The overall venation patterns in species of Aglaonema are similar, but taxonomists like Engler (1915) 

have placed much importance on the relative distinctiveness of the primary lateral veins and on the angle 

of divergence from the midrib. 

326-551 0-69-2 
14 

This process is repeated until lateral veins of a fifth 

or sixth order are formed. 

Finer cross-venation, which is unbranched and 

rather short, develops between the lateral veins. 

The leaf blades of many Aglaonema are variegated 

in patterns that are more or less distinctive for certain 

species or cultivars. Aglaonema crispum has an ashy 

variegation that completely covers about two-thirds 

of the center of the leaf, the outer, marginal areas 

being of a normal dark green color. Aglaonema 

nitidum seedlings frequently show streaks of ashy 

variegation along the primary veins and sometimes this 

pattern is carried on into the adult plants, which are 

known as A. nitidum f. curtisii. Aglaonema commutatum shows similar ashy patterns. Aglaonema 

costatum and A. breuispathum commonly exhibit scattered white or yellowish spots and/or a white midrib. 

Aglaonema uittatum typically has a white midrib. 

Aglaonema rotundum has red variegations following 

the primary veins. Aglaonema pictum has rather ashy 

to silvery irregular blotches spattered over the leaves. 

Bergdolt (1955) studied the ontogeny of variegation 

in Aglaonema roebelinii ( =A. crispum), A. roebelinii 

var. pseudobracteosum ( =A. commutatum cv. 

‘ Pseudobracteatum’), A. costatum, A. marantifolium 

(=A. commutatum var. maculatum) as well as in 

species of other genera of Araceae. In all cases she 

found that in the initial stages all cells of the epidermis, 

mesophyll, and palisade layers contained colorless 

plastids of the same size. During ontogeny the chloroplasts of certain groups of cells of one or more layers 

failed partially or entirely to enlarge and become 

green, creating the variegated patterns. Some of these 

chloroplasts degenerated, making the process irreversible. In other cases the inhibition was found to be 

reversible. The damage to the plastids was found to 

be correlated with cell size; the cells losing their ohloroplasts were distinctly smaller than cells with normal 

plastids. Experiments with Dieflenbachia showed that 

development of variegation was retarded by lowering 

light exposure. Less than normal daylight resulted in 

less variegation. A chemical, disruptive factor (“Storfaktor” ) that was distributed through the vascular 

tissues was made responsible for the damage to the 

plastids. Unfortunately, this factor was not isolated. 

Webber (1960) found that the lateral walls of the 

cells of the upper leaf epidermis were undulate in 

Aglaonema costaturn, A. pictum, and A. commutatum 

var. maculatum. This undulation has also been obSMITHSONIAN CONTRIBUTIONS TO BOTANY 

served in single species of Nephthytis (Webber, 1960), 

Amorphophallus, Homalomena, and Alocasia 

(Dalitzsch, 1886) . 

The epidermal cuticle showed a “ribbing” of more 

or less parallel striations in the three species of Aglaonema studied. This was also observed in Porphyrospatha, Pothos, Homalomena, and Spathiphyllum. 

Stomata in Aglaonema pictum were found only on 

the lower leaf surface. They had only one pair of accessory cells. Stomata were found on both leaf surfaces 

in Aglaonema costatum with varying numbers of 

accessory cells. 

Webber (1960) concluded from his study of the 

epidermal structure of Araceae that “it is doubtful if 

any taxonomic significance can be drawn from these 

findings.” 

INFLORESCENCE.-The morphology of the peduncle, 

spathe, and spadix could be discussed separately but so 

little is known anatomically and morphologically in 

Aglaonema that it is convenient to consider them together. According to Van Tieghem ( 1867, p. 180), the 

structure of the inflorescence axis is similar in all aroids 

except Acorus. Van Tieghem (1867, p. 176; 181) 

found evidence that the peduncle of Acorus calamus 

L. includes two separate vascular systems: one, an 

outer, subepidemal circle of small vascular bundles 

that run into the spathe and the other, a V-shaped 

series of nine large bundles that run into the spadix. 

Such evidence suggests the possibility that in Aglaonema and other Araceae the entire inflorescence may be 

composed of two parts: a shoot with an elongate naked 

base (peduncle) terminating in a spike (spadix) that 

is fused with a foliar organ (spathe) that actually subtends the shoot. Gliick (1919, p. 238) supported this 

interpretation when he concluded that the spathe of 

Araceae must be a leaf lamina without a petiole; however, Van Tieghem (1867, p. 124) reported that the 

vascularization of the peduncle of Aglaonema simplex 

is identical with that the petiole. 

In the same work Van Tieghem described the general structure of the aroid spadix, excluding Acorus. 

Under the epidermis there is cortical parenchyma, then 

a circle of vascular bundles. Inside are scattered a great 

number of similar bundles. Each of these bundles is 

made up of an arc of sclerenchyma with thin walls 

bordered by laticifers-if the plant has them, and 

Aglaonema does-a bundle of phloem, and a large 

group of vessels. These bundles send branches out to 

the flowers. 
NUMBER 1 

Staminate Flower: In Aglaonema it is almost impossible to determine how many stamens there are in 

a staminate flower. To phrase it another way, it is 

almost impossible to define a staminate flower in Aglaonema. There are no subtending perianth parts, there is 

no consistent pattern of groups of stamens, and there 

is no evidence of fusion of stamens. The upper spadix 

is simply a mass of thick stamens, each with a single 

vascular strand and four microsporangia that are 

grouped in two pairs on opposite sides of the stamen. 

With nothing more than external appearances to judge 

by, one could just as well conclude that each stamen 

is a staminate flower. 

Indeed, one worker has said as much. Van Tieghem 

( 1907, p. 315) observed that the endothecium around 

each microsporangium in Aglaonema was complete 

and independent of those of other microsporangia, that 

the xylem of the solitary vascular trace divided at a 

certain level, and that near the apex of the stamen the 

trace itself divided into the two branches which diverged toward the sporangia pairs. From this evidence, 

Van Tieghem concluded that the quadrisporangiate 

body everyone had been calling a stamen in Aglaonema 

was the product of fusion of two bisporangiate anthers 

along their ventral faces. In other words, each quadrisporangiate structure was considered as a staminate 

flower, a synandrium of two bisporangiate stamens. 

This interpretation (Van Tieghem, 1907) has not 

been accepted herein for several reasons. Van Tieghem’s observation that each endothecium was complete and independent has not been supported by my 

observations, nor, for that matter, by Van Tieghem’s 

own illustrations (1867, pl. 8: figs. 11-13) ; even if 

his observations were correct, it would not necessarily 

indicate the conclusion reached by Van Tieghem. 

Van Tieghem’s observation that the xylem branched 

toward the sporangia pairs at the upper end of the 

vascular strand may have a more conventional explanation. For instance, the bifurcation could indicate 

the dichotomous nature of the origin of the stamen. 

Wilson (1942) reached this conclusion from the same 

type of evidence in Tilia and Enallagma. To my 

knowledge, bisporangiate stamens have not otherwise 

been reported in Araceae. In fact, genera related to 

Aglaonema and in which there is no question of the 

definition of the stamen or the staminate flower, have 

exactly comparable quadrisporangiate stamens : e.g., 

Dieflenbachia, where there is fusion of four to five 

quadrisporangiate stamens into synandria, and the 

15 

genera Homalomena and Philodendron, where the 

grouping and orientation of from two to six free 

quadrisporangiate stamens define the staminate flower. 

In addition, the origin of the vascular traces, discussed 

in the following paragraphs, indicates that these quadrisporangiate bodies also can occur in groups comparable to those in the related genera mentioned 

above. Thus, we may describe the stamen in Aglaonema as a quadrisporangiate structure with a single 

trace that bifurcates near the apex. 

The question still remains: How many stamens to 

a staminate flower? Perhaps because external morphology gave no clue, the author of the genus, Schott 

(1829, 1832, 1856, 1859, 1860), refrained from stating 

the number of stamens in a staminate flower of Aglaonema. The first person to mention the number of 

stamens was Engler ( 1879, p. 315), who stated, “Flores 

masculi 2-andri.” This statement also appeared in his 

most recent monograph (Engler, 1915). Since I could 

find no external evidence to support this report of two 

stamens, I attempted to trace the origin of vascular 

traces of stamens by serial cross-sections at ten microns. 

A portion of an inflorescence of Aglaonema nitidum 

(Nicolson 1022) and a portion of an inflorescence of 

A. nebulosum f. nanum (Nicolson 1123) with seven 

stamens were sectioned. 

In Aglaonema nitidum I found that the solitary vascular traces of three stamens arose from a single trace 

that gave off two lateral traces at 170 microns below 

the surface of the spadix. The traces of the other 

nine stamens moved out of the sections before any 

branching or fusion of traces was observed. The common origin of the three traces suggests that the three 

stamens compose a highly reduced staminate flower. 

The sections in the material of Aglaonema nebulosum f. nanum, however, went much deeper (660 

microns below the surface of the spadix), and all 

seven staminal traces were found to be more or less 

related by various branchings and fusions. Such evidence suggested that the deeper one went, the more 

difficult it became to interpret the number of stamens 

from the origin of the staminal traces. 

The results indicated that the number of stamens 

to a pistillate flower might range from one to seven. 

For practical and gross morphological purposes, one 

can say that each stamen appears to represent a 

staminate flower. 

Pistillate Flower: The pistillate flower in Aglaonema 

is unilocular and uniovulate. Only two workers (Van 
16 

Tieghem, 1867; Engler 1915, 1920) have reported its 

structure. 

Relatively little is known of the floral anatomy and 

vascularization or aroid flowers although the flowers 

are of basic importance in the taxonomy of the family. 

Van Tieghem (1867) reported on the vascularization 

of the pistillate flowers in Zantedeschia, Alocasia, and 

Aglaonema. Birdsey ( 1955) discussed the vascularization of pistillate flowers in Syngonium. Eyde et al. 

(1967) surveyed floral anatomy in the family with 

studies on 23 species in 18 genera in six subfamilies. 

Floral structure was found to be extremely diverse. 

Van Tieghem (1867, p. 188) found that the pistil 

in Aglaonema rnarantifolium was unilocular and uniovulate. The ovule was found to be obliquely attached 

at the base and therefore derived from a parietal 

placenta. The wall of the pistil was found to have six 

bundles that were interpreted as three dorsal traces and 

three fused ventral pairs from three carpels. Van Tieghem regarded Zantedeschia as having a basic type of 

trimerous pistil (precisely like the Liliaceae) with parietal placentae and 3-trace carpels. Aglaonema’s organization was derived from this basic type by reduction. 

Engler (1915, p. 10) stated: “Flores feminei I-, 

rarissime 2-gyni.” In the same description he stated: 

“Ovarium 1-, raro 2-loculare.” The ovule was described and illustrated as being anatropous and 

obliquely attached to an indistinct basal placenta or “in 

ovariis 2-locularibus ad basin dissepimenti.” Engler did 

not say in what species he observed the rare 2-locular 

condition, nor have I observed it. 

Engler later ( 1920, p. 35) cited Aglaonema as a case 

wherein a uniovulate ovary was certainly developed 

from more than one carpel. He did not say how many 

carpels were involved, nor did he state what evidence 

led him to this conclusion, but he did indicate that 

he considered that the pistillate flower of Aglaonema 

was derived by reduction from more than one carpel. 

Serial sections of six pistillate flowers (four in crosssection and two in longitudinal section) of Aglaonema 

nitidum (Nicolson 1022) and five pistillate flowers (two 

in cross-section and three in longitudinal section) of 

Aglaonema nebulosum f. nanum (Nicolson 1123) were 

made at ten microns. The vascularization was found to 

vary from flower to flower in an exceedingly confusing 

fashion. The base of each flower had a ring of from 

four to nine traces, and the stylar regions had a ring of 

from four to seven traces. Since each flower differed 

from all others, it was not possible to say if any were 

SMITHSONIAN CONTRIBUTIONS TO BOTANY 

representative of the spadix, the species, or the genus. 

It was not possible to differentiate the carpellary 

traces from the lateral traces in a consistent fashion applicable to each flower. It was not even possible to 

determine, with any degree of certainty, the number of 

carpels composing a pistillate flower. Perhaps this was 

due to reduction, to the peculiar nature of monocotyledonous venation, or to both. Depending upon how 

much weight one might give a particular piece of evidence, it was possible to interpret a given pistillate flower as being composed of one, two, three, or more carpels. 

The inconclusiveness of vascular evidence demands 

that one consider the carpellary facts in related genera. 

Of 17 genera placed in the Philodendroideae by Engler 

(1920), only 6 have unilocular pistils. Except for 

Aglaonema, all of these genera are minor, having only 

three or fewer species. The other 11 genera, including Philodendron, the largest and possibly the most 

primitive genus in the subfamily, have two or more 

locules (carpels). This suggests that Aglaonema may 

have arisen from syncarpous stock. 

Comparison of sections shows a certain bilateral 

symmetry in the arrangement of vascular traces in the 

lower stylar region. Typically, more than half of the 

traces are in an arc of about 120” and the remaining 

traces are widely scattered in the other 240”. This 

positioning of bundles to one side suggests abortion 

of one or more carpels or the presence of sterile carpels. 

If one assumed that abortion occasionally does not 

occur, this would explain Engler’s (1915) report of 

bilocular pistils in Aglaonema. 

Plant Diseases 

According to the Review of Applied Mycology, 

members of Aglaonema have been reported with a leaf 

spot caused by anthracnose fungi and Pythium root rot. 

Baker (1914, p. 2157) reported Gleosporium gra@ 

Sydow on a Philippine plant from the wild that was 

determined as Aglaonema densinervium. Graham and 

Strobe (1958) reported a related fungus, Colletotrichium, on leaves of several genera of oranamental 

foliage plants cultivated in Washington State, including Aglaonerna cornmutatum. 

Pythium splendens root rot, first noticed in Orange 

County, Florida, nurseries in 1943 on Aglaonema 

modestum, has been the subject of several studies. Tisdale and Ruehle ( 1949) discussed the symptoms, which 

develop over a 3-week period. Plants were first observed to show stunting in their aerial parts and drooping of the leaves. Finally the stems rot off at ground 
NUMBER 1 

level and the plants fall over. The roots were found 

to have water-soaked lesions that rot and remain in 

the ground, even if the plants are pulled up. Other 

aroids (Dieffenbachia, Nephthytis, Philodendron, 

Rhaphidophora) were affected by the root rot, but not 

as seriously as Aglaonema. 

A paper discussing control of Pythium root rot in 

Chinese evergreen (A. modestum, although the author 

17 

calls it A. simplex) was published by Miller (1959). 

He took sterile peat, mixed it with infested debris and 

let it sit for four weeks. Various fumigation methods 

were applied (Vapam soil drench, mylone powder 

broadcast, methyl bromide injection) and three weeks 

elapsed before the beds were planted. The highest percentage of loss was 1.27 percent (Vapam) except for 

the untreated control (87.4 percent). 

Aglaonema Schott 

Aglaonema Schott, 1829 (8 Sept.), p. 892. 

Stem smooth, usually green, often rooting at nodes, 

erect and unbranched or creeping and often branched. 

Internodes, usually longest near the stem-base. 

Petioles usually equaling or slightly shorter than the 

leaf-blade. Petiolar sheaths membranous to scarious, 

rarely slightly prolonged, equaling the stem circumference in a few species to extending the length of 

the petiole. Leaf-blades ovate-elliptic to lanceolatenarrowly elliptic, rarely broadly ovate or sublinear ; 

base often unequal, obtuse to cordate or attenuate; 

apex often apiculate, usually acuminate to acute, 

sometimes gradually or abruptly acuminate ; variegation none or of various patterns; venation undifferentiated to strongly or weakly differentiated into 5-8 

primary lateral veins diverging from the midrib at 

40'-60' ; texture coriaceous to membranaceous. Cataphylls usually subtending inflorescences, occasionally 

subtending petioles. Peduncles erect, solitary to less 

than 10 together, from half as long as to longer than 

the subtending petiole, deflexing in fruit. Spathe 

ovate, erect, not differentiated into a tube and blade, 

often apiculate, green to yellowish white, slightly to 

strongly decurrent, usually finally marcescent. Stipe 

almost none to half the spadix length. Spadix thincylindric to clavate, erect, from half as long as to 

longer than the spathe; pistillate portion below, pistils 

1-locular, 1-ovulate, style short and thick, stigma broad 

and disk-like; staminate portion white to cream 

colored, stamens solitary but tightly compacted, opening in pairs at tops of the four sporangia. Fruits ellipsoidal, the outer layer fleshy green but turning yellow 

(rarely white) and finally red. Seed solitary, erect, ellipsoidal, almost as large as the fruit. Embryo macropodal. Endospenn none. 

TYPE OF GENus.-Aglaonema oblongifolium Schott, 

superfluous name (Arum integrifolium Link, Aglaonema integrifolium (Link) Schott) [ = Aglaonema 

nitidum (Jack) Kunth]. 

DISTRIBUTION.-Tropical Southeast Asia from 

northeastern India through New Guinea. 

Key to the Species of Aglaonema 

1. Stem repent, frequently branching; petiole sheath less than 20% the petiole length, less than 

1 cm long; cataphylls subtending the petioles . . . . . . . . . . . . Section Chamaecaulon 

2. Cataphyll subtending petiole usually drying-persistent ; petiole usually equaling or exceeding 

blade; leaf-base usually obtuse to subrounded . . . . . . . . . . . . 1. A. brevispathum 

2. Cataphyll subtending petiole usually immediately deliquescent; petiole usually less than 

blade length; leaf-base subcordate to rounded . . . . . . . . . . . . . 2. A. costatum 

1. Stem erect, rarely decumbent, rarely branching; petiole sheath rarely 20% the petiole length 

usually more than 1 cm long; cataphylls usually only subtending peduncles, rarely among the 

petioles . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Section Aglaonema 

3. Spadix sessile, the stipe and major part of pistillate portion of spadix adnate to spathe. 

4. Pistillate flowers separated from functional staminate flowers by several series of staminodia, 

these slightly larger than and of a different color from the functional staminate flowers (even 

in dried material) ; peduncle 3.5-7.0 ( 10) cm long . . . . . . . . . . . 3. A. ovatum 

4. Pistillate flowers contiguous with functional staminate flowers, or staminodia, if present, few 

(i.e., only a single series) and indistinguishable from functional staminate flowers except for 

their lack of pores; peduncle 5-21 cm long. 

5. Apex of spadix short 1 cm or more of spathe apex; leaf-acumen elongate, over 2.5 cm 

from point of 1 cm blade width; peduncle 5-10 cm long . . . . . . 4. A. modestum 

5. Apex of spadix equaling (within 0.5 cm) spathe apex; leaf-acumen shorter, less than 2.5 

cm from point of 1 cm blade width; peduncle (6) 10-21 cm long . . 5. A hookerianum 
18 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

3. Spadix stipitate, the stipe free from spathe for 0.2 cm or more (at anthesis or after). 

6. Spadix clavate; spathe globose. 

7. Spadix exserted from spathe at anthesis; upper leaf-surface dull; leaves not variegated or 

else covered with irregular silver or gray blotches. 

8. Petiole sheath more than '/z petiole length. Plants of mountain slopes, commonly over 

1OOOm. Sumatra. ...................... 6. A. pictum 

8. Petiole sheath less than 36 petiole length. Plants of lowland swamp forest. Malaya and 

Borneo ........................... 7. A, nebulosum 

7. Spadix included in spathe, even at anthesis; upper leaf surface shiny, leaves not variegated 

or else only with reddish or whitish midrib and/or primary lateral veins. 

9. Leaf-blade lanceolate, 1.7-3.0 cm wide .............. 8. A. vittatum 

9. Leaf-blade ovate, 5-10 cm wide ................ 9. A. rotundum 

10. Apex of spadix equaling, exceeding, or short no more than 1 cm of spathe apex; pistillate 

10. A. nitidum 

6. Spadix cylindric; spathe elongate. 

flowers usually more than 20 (except in A. cochinchinense and A. nitidum var. helferi). 

11. Margin of petiole sheath scarious; leaf-base acute to attenuate . . 

11. Margin of petiole sheath usually membranous; leaf-base obtuse to rounded. 

12. Venation undifferentiated ............... 1 1. A. cochinchinense 

12. Venation differentiated into primary and secondary veins. 

13. Spathe 1.8-2.5 (4) cm long ................. 12. A. tenuiper 

13. Spathe (2.3) 3-5 (6.5) cm long ............... 13. A. simplex 

length. .......................... 14. A. cordifolium 

10. Apex of spadix short 1 cm or more of spathe apex; pistillate flowers usually less than 20. 

14. Leaf-blade broadly ovate with cordate base; petiole sheath less than )$ the petiole 

14. Leaf-blade narrowly elliptic-oblong with acute to rounded base; petiole sheath 

15. Stipe (usually over 1 cm long) 0.3 cm longer than spathe decurrency (this less than 

15. Stipe (usually less than 1 cm long) shorter or to 0.2 cm longei than spathe deusually more than $5 the petiole length. 

1 cm; see also A. commutatum). .............. 15. A. marantifolium 

currency (this usually more than 1 cm long). 

16. Leaves variegated. 

17. Variegation (usually in form of bars, rarely in spots) oriented along primary 

lateral veins; margins of petiole sheaths usually membranous 

........................ 16. A. commutatum 

17. Variegation forming solid band down center of each side of leaf parallel to 

....... 17. A. crispum 

18. Leaf-blade over 4 times longer than wide; petiole length 56 the leaf-blade or 

18. Leaf-blade less than 3.5 times longer than wide; petiole length X the leaf19. Petiole sheath 17-30 cm long, 2/5 to equaling the petiole; leaf-blade 12-19 cm 

wide; peduncle 20-32 cm long ........... 19. A. densineruium 

19. Petiole sheath 1-17 cm long, )/43//4 the petiole; leaf-blade 2.5-13 cm wide; 

peduncle 2-1 7 cm long. 

20. Petiole (8.5) 14-25 cm long ........... 20. A. philippinense 

20. Petiole 3.5-7.0 cm long ............... 21. A. pumilum 

....... 

leaf margin; margins of petiole sheaths scarious 

16. Leaves not variegated. 

less ........................ 18. A. stenophyllum 

blade or more. 

Aglaonema section Chamaecaulon Nicolson, 

new section 

Caudex repens, internodiis vix 0.5 cm longis; cataphylla petiolum plerumque amplexa, nonnumquam, 

deliquescens; petioli vagina usque ad 1 cm longa, vix 

v5 petioli longitudinis. 

Roots frequently approaching the stem thickness. 

Stem repent and often branching, usually less than 1 

cm thick. Internodes usually less than 0.5 cm long. 

Petiolar sheaths inconspicuous, to 1 cm long, 0.03-0.15 

times as long as the petiole. Leaf-blades lanceolate to 

ovate, rarely narrowly elliptic ; variegation none or with 

white spots affecting both sides of leaf and/or a central 

white stripe on or over the midrib; venation usually 

differentiated. A cataphyll subtending and clasping 

each petiole, this soon deliquescent in A. costaturn. 

Peduncle solitary. Spathe ( 1.5) 2.5-4.0 (5) cm long. 

Spadix ellipsoidal-cylindric, (1.2) 1.8-3.5 cm long, 

commonly exceeding spathe by 0.3-1.0 (1.5) cm; 
NUMBER 1 

pistillate portion 0.2-0.5 cm long, pistils 6-13. Fruits 

red, 1.2-1.7 cm long, 0.5-1.0 cm thick. 

TYPE OF sEcTIoN.-Aglaonema costatum N. E. 

Brown. 

DISTRIBUTION.-TWO species of continental Southeast Asia. 

HABITAT.-Humid shady places below 500 m. 

The section Chamaecaulon is characterized by a 

creeping and branching habit, exceedingly short petiolar sheaths, and the occurrence of cataphylls among the 

leaves. In these characters it is similar to two genera 

that are considered to be closely related, Aglaodorum 

(west Malesia) and Anubias (Africa) ; however, unlike 

Aglaonema, both these genera are tidal mudflat or 

swamp dwellers. 

Within the genus Aglaonema, several of the smaller 

species, such as A. pumilum and A. rotundum, seem 

to be intermediate between the two sections. These 

rare and poorly known species are decumbent, occasionally have short petiolar sheaths, and may have 

cataphylls among the leaves. 

1. Aglaonema brevispathum 

Aglaonema brevispathum (Engler) Engler, 1915, p. 32, fig. 

Stem repent and often branching, rarely with more 

than three leaves, 0.5-1.0 cm thick. Internodes ca. 

0.3 cm long, very rarely 1.5 cm. Petioles (7) 13-25 

(33) cm long, (0.5) 0.8-1.2 (1.6) times as long as the 

blade length. Sheaths 0.5-1.0 cm long, 0.03-0.15 times 

as long as the petiole. Leaf-blades lanceolate, rarely 

narrowly elliptic, (10) 14-20 (24) cm long, (3.9) 4.4- 

15. 

19 

8.0 cm wide, lengthjwidth ratio 1 : 2.54.2; base obtuse, 

occasionally rounded or broadly acute, often unequal ; 

apex usually acuminate, sometimes acute to subacuminate or gradually acuminate; variegation none, in the 

form of a central white stripe, or in the form of 

scattered spots on both sides of the leaf; venation 

weakly or strongly differentiated into 4-7 primary 

lateral veins diverging from the midrib at (30) 40”- 

50” ; texture coriaceous. Cataphylls like brown paper, 

commonly clasping the petiole for most of their length. 

Peduncle solitary, (5.5) 7-12 (15.5) cm long, usually 

‘/3-3/4 the petiole length. Spathe usually apiculate, 

1.5-3.5 cm long, decurrent for 0.5-1.3 cm. Stipe 0.2- 

1.0 cm long. Spadix ellipsoidal, 1.2-2.5 cm long, equaling to exceeding spathe by 0.3-0.6 cm; pistillate portion 

0.2-0.5 cm long, pistils less than 10; staminate portion 

1-2 cm long, 0.3-0.6 cm thick. Fruits red, 1.2-1.7 cm 

long, 0.5-1.0 cm thick. 

DIsTRIsuTIoN.-EaStern continental Southeast Asia 

(Figure 3). 

HAsITAT.-Near streams or on heavily wooded 

slopes below 500 m. 

FLOWERING TIME.-March-July. 

Aglaonema breuispathum is obviously most closely 

related to A. costatum. The differences used in the key 

seem to be more subjective than objective (Poilane 

10746 from South Vietnam would key to A. costatum 

but its narrow leaves (1 : 4), few primary veins, and 

obtuse leaf base are more similar to A. breuispathum). 

Further studies and evidence would be necessary before 

these species could be united. 

Key to the Forms of Aglaonema brevispathum 

Leaf-blade without scattered spots ..... 

Leaf-blade with scattered spots. ...... 

la. Aglaonema brevispathum f. brevispathum 

Aglaonema brevispathum (Engler) Engler, 1915, p. 29. 

Homalomena brevispatha Engler, 1902, p. 172. 

Aglaonema ridleyanum Engler, 1915, p. 28, fig. 12. 

A. hospitum var. obtusatum Engler, 1915, p. 29. 

A. costatum var. viride sensu Gagnepain, 1942, p. 1124 [not 

Leaf not blotched; venation usually strongly differentiated into 3-6 primary lateral veins diverging from 

the midrib at (35”) 45”-50”. Spathe apiculate, 3-3.5 

(1.8, fide Engler, 1915) cm long. Stipe 0.2-0.8 cm 

long. Fruit red, 1.2-1-7 cm long, 0.5-1 .O cm thick. 

TYPE coLLEcTroN.-Thailand, Koh Chang, Skoven 

nar Klong Son, 1 March 1900, Schmidt 675a (holotype : B ; isotype : C) . 

Engler, 19 151. 

........... 1 a. forma breuispathum 

.............. 1 b. forma hospitum 

DISTRIBUTION .-Eastern Southeast Asia. 

HABITAT.-Near streams or on heavily shaded 

FLOWERING TIME.-March-July. 

Aglaonema ridleyanum must be typified on the 

drawing published by Engler (1915, fig. 12) since no 

specimen or collector was cited and no authentic 

specimens have been found. Engler cited “Siam 

(Bangkok.-Bot. Gart. Singapore) .” The illustration fits 

A. breuispathum but the description has several mis1onga”-the “cm.” is presumably an error for “dm.” 

although 20-30 cm would be an excessively long leaf; 

slopes below 500 m. 

leading statements: ( 1 ) “Lamina .... 2-3 cm. 

(2) “nervis ... angulo circ. 80°”-which is exces- 
Figures 2-4.-Distribution of nine species of Aglaonema 
Figures 5-8.-Distribution of twelve species of Aglaonema. 
22 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

FIGURE 9.-Isotype of Aglaonema brevispathum (Engler) 

Engler : Schmidt 675a (C) . 

sively broad, not only for this species but for the type 

drawing; (3) “Pedunculi plures . . .”-which is very 

disturbing because members of the section Chamaecaulon have solitary inflorescences. 

Gagnepain (1942, p. 1129), who had no more information than that published by Engler, regarded A. 

ridleyanum as having an erect stem; however, Englcr 

placed A. ridleyanum next to A. hospitum, which is 

treated herein as part of A. brevispathum. With the 

exceptions noted above, the descriptions and illustration of A. ridleyanum fit well into A. brevispathum. 

Since the description of A. ridleyanum did not mention 

variegation, I am placing it with this forma. 

The typification and interpretation of A. hospitum 

var. obtusatum are also problematic. Engler cited his 

material as “Cochinchina (L. Pierre.-Herb. Mus. Paris, 

Berlin).” A diligent search by me in the Berlin herbarium failed to reveal any specimens, now presumed 

lost. In the Paris herbarium there are only three Pierre 

sheets annotated by Engler as A. hospitum although 

none of them are annotated as A. hospitum var. obtusatum. Since two of the three sheets clearly have erect 

stems and are not referable to section Chamaecaulon, 

they were excluded from further consideration; they 

have been identified, in fact, as Aglaonema tenuipes 

Engler. According to my notes, the third sheet consists 

of separate specimens, one of which has a central white 

stripe on the leaf-blade and no spots, while the other 

specimens appear to have the leaves totally unmarked. 

In the absence of other evidence, we must rely on 

Engler’s description : “Foliorum lamina basi obtusata 

atque immaculata.” Since all leaves of these Paris 

specimens are obtuse, “immaculata” becomes definitive. I therefore recognize the unmarked material as 

typifying Aglaonema hospitum var. obtusatum. 

Another sheet (Pierre s.n.), not annotated by Engler, 

is a mixture of three forms of A. brevispathum, no 

markings on one specimen, blotches on a second specimen, and a central white stripe on a third specimen. 

The mixture of forms on these sheets indicates that 

the taxa are of minor significance. If the white striped 

form becomes better known, it probably will be named 

as a forma. 

All specimens annotated and cited by Gagnepain 

(1942) as A. costatum var. viride have been placed 

here. 

lb. Aglaonema brevispathum f. hospitum 

Aglaonema brevispathum f. hospitum (Williams) Nicolson, 

Schismatoglottis siamensis Bull, 1885, p. 15 [not Aglaonema 

Aglaonema hospitum Williams, 1904, p. 226. 

Leaf variegation in form of small spots that affect 

both sides of the leaf; venation weakly to strongly differentiated into 4-7 primary lateral veins diverging 

from the midrib at 40”-50”. Spathe 2.5-3.5 cm long. 

Stipe 0.5-1 cm long. Fruits unknown. 

TYPE COLLECTION.-HOng Kong Botanic Garden 

No. 59/88, received from Bangkok several years ago. 

30 May 1888 [without collector] (holotype: K). 

DIsTRIsuTIoN.~Thailand (Koh Chang) and “Cochinchine.” 

HABITAT.-LOwland rainforest. 

UsEs.-widely cultivated as an ornamental for its 

1968, p. 124. 

siamense Engler, 19021. 

variegated leaves. 
NUMBER 1 

The type of Schismatoglottis siamensis Bull is: England, Hort. Bull, Siam, 10 Nov. 1885 [N. E. Brown?] 

(holotype: K) . This epithet is the oldest applicable 

not only to this forma but to the entire species. It is 

unavailable at the specific level as its transfer to Aglaonema would create a later homonym of A. siamense 

Engler ( 1902) ( =A. tenuipes) . 

In collecting this forma in the wild on Koh Chang, 

I noted that plants with unvariegated leaves occurred 

in the same population (Nicolson 1623), This indicates 

that the presence of variegation is only a minor 

character. 

2. Aglaonema costatum 

Aglaonema costatum N. E. Brown, 1892, p. 426. 

Roots up to 0.5 cm thick. Stem repent and branching, 0.6-1.3 cm thick. Internodes 0.1-0.5 cm long. 

Petioles 5-12 (19) cm long, (0.3) 0.5-1 times as long 

as the leaf blade. Sheaths 1 cm long, 0.05-0.15 (0.3) 

times as long as the petiole length. Leaf-blade usually 

ovate, rarely lanceolate, 9.5-20 (26) cm long, (4.7) 

6-10 cm wide, length/width ratio 1 : 1.5-2.5 (3.6) ; 

base often unequal, subcordate to rounded ; apex often 

apiculate, acute to subacuminate, occasionally abruptly 

acuminate; usually variegated with scattered spots on 

both sides of the leaf and/or with a white midrib, rarely 

not variegated; venation differentiated into 7-15 primary lateral veins diverging from midrib at 50’-70’. 

Cataphylls 5-8 cm long. Peduncle solitary, (2.5) 5-15 

(23) cm long, 3/4 to longer than subtending petiole. 

Spathe often apiculate, (2.2) 2.5-4.0 (5) cm long, 

decurrent for 0.5-1.5 (3) cm. Stipe 0.2-0.6 cm long. 

Spadix ellipsoidal-cylindrical, exceeding spathe by 

0.3-1.0 (1.5) cm, (1.8) 2-3.5 cm; pistilate portion 

0.2-0.5 cm long, pistils 6-13; staminate portion 1.5-3.3 

cm long, 0.5-0.8 cm thick. Fruits unknown. 

DIsTRrsuTIoN.-Continental Southeast Asia from 

Langkawi Island to Vietnam (Figure 3) . 

HABITAT.-Humid, shady places on limestone or 

near streams. 

FLOWERING TrME.-March-June. 

This species is closely related to Aglaonema brevispathum and occasional specimens prove difficult to 

determine. These specimens involve A. brevispathum 

f. brevispathum, which may be difficult to separate 

from A. costatum f. immaculatum and f. concolor. 

Field studies are needed to determine how similar or 

different A. costatum and A. brevispathum really are. 

Aglaonema costatum has been popular in cultivation for its variegated forms that have been introduced 

from the wild. Birdsey (1951, p. 14) dismissed the 

forms because he had observed that the amount and 

kind of variegation varied greatly on leaves of the 

same plant. It is true that the patterns are strongest on 

youngest leaves and that they fade with age, but I have 

not observed the great variability on the same plant 

emphasized by Dr. Birdsey. Perhaps such forms do 

not warrant taxonomic recognition but, in this case, all 

the variants have been named and I believe it would 

be well to clarify them and their nomenclature. 

Key to the Forms of Aglaonema costatum 

1. Leaf-blade with a white midrib. 

2. Leaf-blade with irregularly scattered spots 

2. Leaf-blade without spots .................. 2b. forma immaculatum 

3. Leaf-blade heavily and irregularly blotched ........... 2c. forma virescens 

............ 2a. forma costaturn 

1. Leaf-blade with a green midrib. 

3. Leaf-blade not variegated ....... .. 

2a. Aglaonema costatum f. costatum 

Aglaonema costatum N. E. Brown, 1892, p. 426. 

A. costatum var. maculatum Engler, 1915, p. 17. 

A. costatum var. foxii Engler, 1915, p. 17. 

Petioles 5.0-7.6 (18) cm long. Leaf-blades 9.5-18 

cm long, 5.3-10 cm wide, lengtn/width ratio 1 : 1.5- 

1.9; variegation in the form of a central white stripe 

and scattered spots on both sides of the leaves; venation differentiated into 7-15 primary lateral veins diverging from the midrib at 60”-70’. Peduncle (2.5) 

4.5-9.0 cm long. 

TYPE coLLEcTroN.-England, Hort. Veitch, intro- 

............ 2d. forma concolor 

duced from Langkawi Island by Curtis, 26 March 

1892 [N. E. Brown?] (holotype: K) . In the original 

publication Brown stated : “A native of Perak.” Curtis, 

the original collector, noted on another specimen collected by him on Langkawi Island: “The habitat 

Perak in G[ardeners’] C[hronicle] is an error.” Brown, 

author of the error, corrected his note on the holotype 

so that it now reads: “Introduced from Langkawi Island by Mr. Curtis.” 

DIsTwsuTIoN.-Endemic on Langkawi Island. 

HABITAT.-“Damp shady places at foot of limestone 

hills” (Curtis 2813) . 
24 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

FLOWERING TIME.-March, coinciding with the onset of the rainy season. 

Aglaonema costatum var. maculatum is typified by: 

Malaya, Langkawi Island, Coah, March 1892, Curtis 

2813 (K, SING). In his description Engler did not 

mention that the midrib is white as it is on the typespecimens. It does not differ from the typical element 

of the species. 

Aglaonema costatum var. foxii is a rather dubious 

element. The type is apparently no longer extant, the 

citation of the type-locality includes errors, and the 

short description does not exactly fit any plants known 

to me. Engler’s description-“Leaf dark green, marked 

with a pale green blotch which irregularly flows out 

from the midrib on each side”-best fits Furtado s.n., 

22 September 1928 (SING). Photographs published 

by Graf (1957, 1959, 1963) as Aglaonema costatum 

foxii also show this, which is rather different from the 

simple white midrib found in A. costatum f. immaculatum; however, all of these also have a few scattered 

spots, which Engler did not mention. Perhaps these 

spots had faded on a dried specimen. Interpreted in 

this fashion, since A. costatum var. foxii is no more than 

typical A. costatum with broader variegation on the 

midrilb and fewer spots than usual, it should be regarded as a cultivar. 

2b. Aglaonema costatum f. immaculatum (Ridley) 

Nicolson, new status 

Aglaonema costaturn var. irnrnaculatum Ridley, 1907, p. 20. 

A. costaturn var. lineatum Engler, 1915, p. 17. 

Petioles 5-18.5 cm long, grooved in upper quarter. 

Leaf-blades 12-23 cm long, 4.7-8.0 cm wide, length/ 

width ratio 1: (1.6)2.1-3.1(3.6) ; variegation in form 

of a white midrib; venation differentiated into 7-12 

primary lateral veins diverging from the midrib at 

60”-70”. Peduncle (6.5) 8-13 cm long. 

TYPE coLLEcTIoN.-Singapore Botanic Gardens, 

1898, Ridley s.n. (holotype: K). 

DIsTRIBuTIoN.-Continenta~ Southeast Asia. 

HABITAT.-Forest, near streams. 

FLOWERING TIME.-April-June. 

The type of Aglaonema costaturn var. lineatum presumably was deposited at Berlin but, if so, it is no longer 

extant. No material was found elsewhere, including 

Bogor, that could qualify as a type. The description 

“Folia supra atroviridia, costa albida” is sufficient to 

warrant placing the taxon here. 

The wild material shows considerable variability. 

Kerr 21263 has leaves ranging from 1.6-3.1 times longer than broad. Kerr also noted “leaves sometimes with 

white stripe along midrib.” This suggests that the taxon 

hardly can be regarded as more than a form occurring 

in the same population with other forms. 

Alston 792 (cultivated in Ceylon) has long petioles, 

a cataphyll subtending the petiole, and narrow leaves 

like those of A. brevispathum, but the base is subcordate and the veins are many and wide-spreading as 

in A. costatum. 

2c. Aglaonema costatum f. virescens ( Engler) 

Nicolson, new status 

Aglaonerna costaturn var. uirescens Ridley ex Engler, 1915, p. 

Petioles 9-19 cm long, grooved for entire length in 

fresh material. Leaf-blades 15-26 cm long, 8.0-10 cm 

wise, length/width ratio 1 : 1.8-2.4; variegation in the 

form of irregularly scattered blotches of white sometimes on both sides of the leaf; venation differentiated 

into 10-14 primary lateral veins diverging from the 

midrib at 50”-70”. Peduncle 11-15 cm long. 

TYPE COLLECTION.-“Kultiviert im Bot. Garten 

Singapore (A. Engler, Reise nach Java und Brit. Indien n. 3811)” (holotype: B, presumed destroyed). 

England, Hort. Kew, 324-05, Singapore, 1914 [without collector] (neotype: K) . In designating a neotype 

I have selected a specimen fitting Engler’s description; 

the original plant came to Kew from Singapore in 

1905, the year of Engler’s visit to Singapore. 

17. 

DISTRIBUTION.-Western Laos. 

Aglaonema costatum var. virens Ridley (in Anderson, 1912, p. 111) is a nomen nudum, presumably 

being referred to by Engler when he adopted the 

epithet “virescens.” 

2b. Aglaonema costatum f. concolor Nicolson, 

new form 

A forma typica folius concoloribus sine notis differt. 

Pedunculus 18-23 cm longus. Pistilis paucis, 7-8. 

TYPE COLLECTION.-Malaya, Kedah, P. Langkawi, 

Selat Panchor, 21 November 1934, Henderson s.n. 

(holotype: SING). 

DISTRIBUTION.-weStern Thailand and southern 

Laos. 

HABITAT.-& limestone, 280 m. 

FLOWERING TIME.-May. 

Aglaonema costatum var. inornatum Ridley (in 

Anderson, 1912, p. Ill), a nomen nudum, is placed 

here with some doubt. Authentic material could not be 
NUMBER 1 

found and I place the epithet here solely on the basis 

of its descriptive name. 

Forma concolor approaches A. brevbpathum in its 

general size although its proportions are those of typical 

A. costatum. The presence of few (7-8) pistils is more 

usual for A. brevispathurn. 

Kostermans, Kwae Noi Exped. 998 from Thailand 

has long petioles and there is a cataphyll subtending the 

petiole, as in A. brevispathum; however, the leaf base 

is subcordate and the veins are many, as in A. costatum. 

Vidal 1852 from Laos has rather narrow leaves (2.5 

times longer than broad) but otherwise seems to fall 

here. 

25 

Aglaonema section Aglaonema 

Roots rarely approaching the stem in thickness. 

Stem erect to partially decumbent, 10-200 cm tall, 

0.2-6.0 cm thick. Internodes 0.4-3.0 (5) cm long. 

Petiolar sheaths conspicuous, (0.8) 4-17 (25) cm long, 

(0.1) 0.3-0.8 (1) times as long as the petiole. Leafblades ovate-elliptic to lanceolate-narrowly elliptic, 

occasionally broadly ovate to sublinear; variegation 

rarely affecting the lower leaf surface, never with a 

central white stripe ; venation undifferentiated to 

weakly or strongly differentiated. Cataphylls usually 

found only among the inflorescences (except possibly 

in A. rotundum and A. pumilurn). Peduncles solitary 

to 7 together. Spathe (1) 3-7 (12) cm long. Spadix 

thin-cylindric to clavate, (1.2) 2.5-5.0 (7) cm long, 

from % the spathe length to exceeding the spathe by 

1 cm; pistillate portion 0.2-1.0 (2) cm long, pistils 

(3) 10-25 (38) ; staminate portion rarely (A. tenuipes) with a gap between staminate and pistillate 

portions. Fruits ellipsoidal, green but turning yellow 

and finally red, up to 3 cm long and up to 1.5 cm 

thick. 

3. Aglaonema ovatum 

Aglaonema ovaturn Engler, 1898, p. 21. 

Stem erect, 0.5-1.5 cm thick. Internodes 0.7-3.0 

cm long. Petioles (5) 8-14 (18) cm long, (0.3) 0.5- 

0.8 (1.1) times as long as the leaf-blades. Sheath 

membranous, (1.5) 3-8 cm long, (0.3) 0.4-0.6 (0.7) 

times as long as the petiole. Leaf-blades ovate to 

lanceolate, rarely elliptic, ( 10) 13-21 (26) cm long, 

(3) 5-10.5 cm wide, length/width ratio 1: (1.7) 

2-3 (5.5) ; base rounded to obtuse, rarely subcordate; 

apex often apiculate, acuminate (acumen 1 .O-2.8 cm 

from point of 1 cm blade width to apex) ; rarely variegated with scattered blotches; venation weakly to 

strongly differentiated into 5-10 primary lateral veins 

diverging from the midrib at 35'-60'. Peduncles 1-2 

together, 3.5-7.0 (10) cm long, usually about f/2 as 

long as the petiole. Spathe 3.5-4.7 cm long, decurrent 

for 0.4-1.5 cm. Stipe none. Spadix cylindric, 3.0-4.3 

cm long, usually exceeding spathe but sometimes 0.5 

cni short of spathe apex; pistillate portion 0.4-1.6 cm 

long, pistils ca. 10-20; staminate portion 2.8-3.5 cm 

long, 0.6-1.0 cm thick, with lower 0.5 cm sterile and 

distinctly differentiated in color and size from the fertile area. Fruits red, 1.1-2.0 cm long, 0.7-1.0 cm thick. 

DrsTRrsuTIoN.-Northem continental Southeast 

Asia (Figure 4) . 

HABITAT.-In evergreen forest along streams or in 

shaded and humid places. 

FLOWERING TIME.-April-May (July), probably influenced by the onset of the summer monsoonal rains 

( March-April) . This species is most closely related to Aglaonema 

hookerianum and A. modestum. It is distinguished by 

having the spadix equaling or exceeding the spathe 

and by rather strongly differentiated staminodia between the pistillate and staminate flowers. Fruiting 

specimens can be distinguished from A. hookerianum 

by their shorter peduncles. Fruiting and sterile specimens cannot be satisfactorily distinguished from A. 

modesturn. The leaves of A. modestum are very longacuminate (acumen over 2.5 cm long from point of 1 

cm blade width to apex), a condition not attained in 

A. ovaturn; however, flowering material of A. modesturn from Laos lacks the extremely long acumen. It is 

hoped more and better flowering material may provide the means of recognizing better vegetative 

characters. 

It should be noted that Aglaonema tenuipes occasionally has subsessile spadices and differentiated 

staminodia, suggesting possible confusion with A. 

ovaturn. I am not certain that all fruiting or sterile 

specimens can be satisfactorily separated from A. 

tenuipes. 

Key to the Forms of Aglaonema ovatum 

Upper leaf surface not variegated . . . . . . . . . . . . . . . . . . . . 3a. forma ovaturn 

Upper leaf surface variegated with ashy blotches . . . . . . . . . . . . 3b. forma rnaculaturn 
26 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

3a. Aglaonema ovatum f. ovatum 

Aglaonema ovatum Engler, 1898, p. 21. 

Scindapsus tonkinensis Krause, 1932, p. 31 1. 

Aglaonema decurrens Buchet ex Gagnepain, 1942, p. 1130 

Leaf-variegation none ; venation usually strongly 

differentiated into 5-10 primary lateral veins diverging 

from the midrib at 35”-60”. Peduncles 1 or 2, 3.5-7.0 

(9.0) cm long, usually less than half as long as the 

petiole. Spathe 3.5-4.7 cm long, decurrent for 0.4-1.5 

cm. Spadix 3.0-4.3 cm long, usually equaling to 

exceeding spathe. 

TYPE coLLEcTIoN.-Cochinchina, in ditione fluvii 

Attopeu (Laos) Harmand 1435 (holotype: P; isotypes: P) . Harmand 1495 nad Herb. Pierre 3347 are 

the same collection and both collectors’ names and 

numbers are found on the holotype. 

DIsTRIBuTIoN.-Northern continental Southeast 

Asia. 

FLOWERING TIME.-March-July. 

The type of Scindapsus tonkinensis is Viet-Nam, 

Hanoi, April 1924, Pe‘telot 1371 (holotype: B; isotypes: BO, P, UC). Krause erroneously placed this 

specimen in Scindapsus, which has bisexual flowers. 

Study of the flowers shows that they are the usual unisexual flowers of Aglaonema. Krause’s description of 

the stem as “scandens” is not supported by the specimen. E. D. Merrill, apparently unaware that he had 

isotypes, correctly determined the unidentified isotypes 

at BO and UC as Aglaonema ovatum. Gagnepain 

( 1942) overlooked Scindapsus tonkinensis in his treatment of the aroids of Indochina. 

The basis of Aglaonema decurrens Buchet ex 

Gagnepain (1942, p. 1130) is the drawing (fig. 107: 

5-8, on p. 1128). The original drawing, in the Paris 

herbarium, was made from material cultivated in the 

Paris Botanic Garden. This material was received as 

Cadiire 27 from Cua-tung, near Hue. This binomial 

was invalidly published since there was no Latin diagnosis. Although Gagnepain regarded A. decurrens as 

new, he placed it among the “esptces insuffisamment 

connues,” along with A. ovatum. The decurrent and 

prolonged spadix is common in A. ovatum. I see no 

reason to consider A. decurrens distinct from A. 

ovatum f. ovatum. 

Dr. Shiu Ying Hu has annotated Liang 69877 from 

Kwangsi as Aglaonema liangii, new species. I do not 

regard the specimen as sufficiently distinct from A. 

ovatum f. ovatum to warrant recognition. 

In leaf shape, Kerr 20899,21776, and Nicolson 1681 

[without Latin diagnosis]. 

represent the narrowest extreme. Other specimens show 

all stages of intergradation to the typical ovate leaf 

shape and, thus, do not warrant recognition. 

3b. Aglaonema ovatum f. maculatum Nicolson, 

new form 

Aglaonema ovatum var. maculatum Gagnepain 1942, p. 1130 

A forma ovato foliorum laminis irregulariter cinereomaculatis differt. 

Leaf-variegation of scattered ashy blotches about 1 

cm square, these frequently irregularly confluent ; venation weakly differentiated into 6-8 primary lateral 

veins diverging from the midrib at 50”. Peduncles 1 or 

2, 8-10 cm long, over half the petiole. Spathe decurrent for 1 cm. 

TYPE coLLEcTroN.-Thailand, Petchabun, Ta Luang, 200 m, evergreen forest, 5 March 1931, Kerr 

20370 (holotype : K; isotypes : K) . This form is known only from a single fruiting collection that shows no evidence of a stipitate inflorescence. Its locality and vegetative characters indicate 

that it belongs with A. ovatum rather than with A. 

modestum or A. hookerianum. It was called A. ovatum 

var. maculatum Gagnepain (1942, p. 1130), but this 

name was invalidly published without a Latin 

description. 

4. Aglaonema modestum 

Aglaonema modestum Schott ex Engler, 1879, p. 442. 

A. acutispathun N. E. Brown, 1885, p. 39. 

A. costaturn var. viride Engler, 1915, p. 17. 

A. laoticum Gagnepain, 1941, p. 117. 

A. marantifolium sensu Troll, 1928, p. 123, figs. 10-13 [not 

A. simplex sensu Bailey, 1930, p. 31 [not Blume, 18371. 

A. modestum cv. ‘Variegatum’ Graf, 1963, pp. 118, 1527. 

Stem dark green, erect, 20-50 cm tall, 0.4-2.0 cm 

thick, internodes 0.4-3.0 cm long. Petioles (6) 10-20 

(22) cm long, (0.4) 0.5-0.9 ( 1.1 ) times as long as the 

leaf-blade, upper portion flattened. Sheaths broad and 

membranous, (2.5) 4-8 (11) cm long, (0.2) 0.4-0.8 

times as long as the petiole. Blades ovate to sublanceolate, rarely elliptic, ( 12) 14-25 cm long, (5) 8-1 1 cm 

wide, length/width ratio 1 : 1.9-2.5( 2.8) ; base often 

unequal, obtuse to rounded ; apex gradually longacuminate (typically 2-3.5 cm from point of 1 cm blade 

width to apex) ; variegation none; venation weakly to 

strongly differentiated into 4-6 primary lateral veins 

diverging from the midrib at 30”-50”. Peduncles 1-3 

together, 5-12.5 cm long, usually more than half as 

long as the petiole. Spathe apiculate, (3) 6-9 cm long, 

[without Latin diagnosis]. 

Blume, 18371. 
NUMBER 1 27 

FIGURE 10.-Reproduction of the type (no longer extant) of 

Aglaonema modestum Engler. 

( 1.5) 2.4-2.6 cm wide, turning yellow with age, decurrent for 0.5-2.0 cm. Stipe none. Spadix thin-cylindric, 

( 1.5) 3-4.3 cm long, (0.6) 1.2-3.1 cm short of spathe 

apex; pistillate portion 0.5-1.0 cm long, adnate to 

spathe, pistils 9-13; staminate portion 2.3-3.5 cm long, 

0.3-0.6 cm thick, not shedding pollen. Fruits orange, 

2-3 cm long, 1.0-1.3 cm thick. 

TYPE CoL.LECTIoN.-Philippines, Manila, Gaudichaud s.n. (holotype: P, fide Engler, 1915, but not 

found). Schott, Aroideae Nr. 56 (W), a drawing of 

the holotype, is suggested here as a neotype, if the 

holotype has been lost. The neotype is the basis for the 

drawing distributed as Engler, Araceae 74 (seen in B, 

G, K, SING, Z) . 

D1sTRIBUTION.-southern China, northern Laos, 

and northern Thailand (Figure 4). 

HABITAT.-Humid ravines in tropical and semideciduous forest. 

FLOWERING TIME.-March-April. 

This species is commonly called “Chinese evergreen” 

in the United States and is popular as a house plant. It 

is called “Wan-lien-tching” in Chinese (Morse 374), 

literally “10,000 years green,” i.e., evergreen, and is 

considered to bring good fortune to the owner. 

Hitherto, this species was thought to be native to the 

Philippines since the type supposedly was collected 

there. Merr2ll (1923, p. 184) suggested that the type 

actually was collected in Macao. Merrill (1932, p. 94) 

later reconsidered and thought it highly probable 

that Gaudichaud 

actually did secure his material from Manila but that the 

specimens were taken from plants imported from southeastern 

China by the Chinese and grown for ornamental purposes in 

Manila; it seems probable that the species has been cultivated 

for a long time by the Chinese, and that it was more or less 

disseminated by them in the Orient, quite as in more recent 

times it has been disseminated by them in the United States. 

This hypothesis is supported by my findings. 

Aglaonema modestum is most closely related to A. 

hookerianum and A. ovatum. It differs from both in 

having the spadix distinctly shorter than the spathe 

(by more than 1 cm) . Vegetative or fruiting material 

is difficult to determine with any degree of certainty. 

Aglaonema modesturn typically has a long acumen on 

the leaf apex; however, this is not found in the material 

called A. laoticum although the spadix is 1.3 cm short 

of the spathe apex and the species clearly is referable 

to A. modestum. 

The type of Aglaonema acutispathum N. E. Brown 

is: England, Hort. Kew., from Hong Kong by H. V. 

Knaggs, 1 June 1885 [Brown] (holotype: K) . Brown 

(1903, p. 185) himself reduced his taxon to synonymy 

saying, “In my description I stated that it was distinct 

from A. modestum, but Engler’s reproduction of 

Schott’s figure of the species (Engler, Araceae No. 74), 

which I had not then seen, demonstrates that the two 

are identical.” 

‘The type of Aglaonema costatum var. viride Engler 

is: Hort. Bot. Calcuttensis, Perak [origin], April 1896, 

Kunstler s.n. (holotype: CAL). The origin “Perak” 
28 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

into 7-13 primary lateral veins which diverge from the 

midrib at (30’) 40”-50”. Peduncles 1-3 together, (6) 

10-21 cm long. Spathe 3.7-6.0 cm long, decurrent for 

(0.6) 1-1.5 (2) cm. Stipe none. Spadix thin-cylindric, 

2.5-4.0 cm long, equaling to 0.8 cm short of spathe 

apex; pistillate portion 0.3-0.6 cm long, attached to 

spathe, pistils ca. 10-15; staminate portion 2.0-3.7 cm 

long, 0.3-0.6 cm thick. Fruits red, large when ripe, 

(1.7) 2-3 cm long, (0.7) 0.9-1.4 cm thick. 

TYPE CoLLECn0N.-India, Churra [Cherrapunji], 

2000 ft, 15 June [18]50, Hooker and Thomson s.n. 

(holotype : K) . 

DIsTRrsuTIoN.-Northeastern India, East Pakistan, 

and coastal western Burma (Figure 3). 

HABITAT.-BelOW 1000 m, in deep shade of forest 

receiving more than 80 inches of annual rainfall. 

FLOWERING TIME.-June-July, probably influenced 

by the onset of the summer monsoon. 

Aglaonema hookerianum Schott, as indicated by the 

key to species, is very similar to A. modestum and A. 

ovatum. The primary differences between these species 

seem to be in the spadices. Most of the specimens 

available are in fruit; because of the paucity of flowering material, I am not certain just how distinctive these 

taxa are. Secondary considerations like plant size, leaf 

shape, and distribution were used for identification. 

Also, lack of collections from northern Burma and 

southwestern China made it impossible to know 

whether or not intergradations occur in the areas where 

they would be most expected. The distinctive characteristics of A. hookerianum are : ( 1 ) sessile spadix, [ 2) 

spadix equaling the spathe (3) large fruits, (4) a long 

peduncle (compared with A. ovatum and A. modestum) , and (5) restriction to western Burma, southern East Pakistan, and northeastern India. 

At one time it seemed to me justifiable to segregate 

Aglaonema ovatum, A. modestum, and A. hookerianum 

as a separate section of the genus because of their 

sessile spadices and northern distribution. The absence of any other distinguishing characters and the 

occasional occurrence of subsessible spadices in A. 

simplex, A. tenuipes, and A. cochinchinense made this 

segregation untenable. 

The type of Aglaonema clarkei Hooker f. is: East 

Pakistan, Kaselong [Kaselang River], 10 Jan. 1869, 

Clarke 8266 (holotype: K). Aglaonema clarkei is 

similar to A. hookerianum with its large seeds, sessile 

spadix, and long peduncle. It differs primarily by its 

exceedingly short petiolar sheath and scandent habit. 

noted on the specimen seem unlikely unless it was 

from cultivation. The specimen is annotated by Engler 

only as “Aglaonema costatum N. E. Br.” without mention of a varietal name; however, this is the only specimen I have seen that fits Engler’s citation “Malakka, 

Perak (Kunstler. - Kult. im Bot. Gart. Calcutta) .” 

Only one leaf and one inflorescence (one destroyed) 

are present, but the material seems to be A. modestum 

or, possibly A. hookerianum. Furtado annotated the 

specimen as “certainly not A. costatum.” 

The types of Aglaonema laoticum Gagnepain are: 

Lzos, Luang Prabang, Ken Luong [no date], Spire 726 

(lectotype: P) ; [Laos] Paklai, 1866-1868, Thorel 5.71. 

(syntype: P) ; Thailand, Chieng Mai, Doi Soo-tep, 

3000 ft, 2 May 1910, Kerr 1160 (syntype: K). This 

lectotype was selected because it is the most complete 

(with flowers and fruits) and because Gagnepain made 

sketches of the inflorescence on it. The spadix is sessile, 

reaching 1.3 cm short of the spathe apex, and without 

conspicuous staminodia, indicating that this is A. 

modestum although the acumen of the leaves is short 

for A. modestum. The large seed (3 x 1 cm) is usual 

for A. hookerianum, but the plants are much smaller 

than those of that species. 

The binomial Aglaonema simplex is often misapplied 

to A. modestum. This misidentification has been 

pointed out by Merrill (1932, p. 93). True A. simplex 

has a stipitate spadix and only rarely has the long 

acumen commonly found in A. modestum. 

Aglaonema modestum cv. ‘Variegatum’ was reported to come from Pennock’s Puerto Rico Nursery. 

The photograph published by Graf (1963) is not 

clear but the plant looks as though the variegation 

might be caused by a disease. 

5. Aglaonema hookerianum 

Aglaonema hookerianum Schott, 1859, p. 30. 

A. clarkei Hooker f., 1893, p. 529. 

Stem erect, 40-50 cm or more, 1.5-2.0 cm thick. 

Internodes 1.5-3.0 cm long. Petioles 14-24 cm long, 

0.7-0.9 (1.2) times as long as the leaf-blade. Sheaths 

membranous, (4) 8-15 cm long, (0.2) 0.5-0.6 (0.8) 

times as long as the petiole. Leaf-blade ovate to elliptic 

or lanceolate to narrowly elliptic, 20-27 cm long (5.3) 

7-12 cm wide, length/width ratio 1 : 2.0-2.9( 3.8) ; 

base often unequal, rounded, obtuse or broadly acute, 

rarely acute; apex often apiculate, acuminate to gradually or suddenly long acuminate (acumen to 2.1 cm 

long from point of 1 cm blade width to apex) ; variegation none; venation weakly to strongly differentiated 
NUMBER 1 

The report of scandent habit is surely an error for the 

reason that, although large plants of Aglaonema tend 

to recline against their neighbors, they are not climbers 

in the usual sense of the word. The short petiolar 

sheath is represented only by the one leaf on the type. 

Superficially the taxon resembles A. nitidum in its 

acute leaf-base and lack of differentiated venation, 

characters occasionally found in A. hookerianum 

(Prazer s.n. and Leslie 306). It seems best to regard 

A. clarkei as a probable synonym of A. hookerianum. 

The type-locality, on the Kaselang River, may now be 

drowned by the Karnaphuli Reservoir. 

29 

( 1844, vol. 3, pl. 804). The typification of this epithet 

was discussed by Sealy ( 1957). 

DISTRIBUTION.-SUmatra and Nias Island (Figure 

HABITAT.-usually reported between 1000 and 2000 

m on the slopes of major volcanos. Occasionally collected as low as 35 m (Jacobson 127 from Padang) . 

FLOWERING TIME.-April or August. Most fruiting 

material was collected in March. Seasonality is weak 

and probably variable according to the exposure of the 

mountain side. 

The type of Aglaonema gracile is: Sumatra? 

[Korthals] (holotype: L) . The origin of this specimen 

has long been in doubt because the original label was 

lost. In the original publication Schott stated: “Borneo, 

Sumatra vel Iava (Sched. deperd.) .” The type is 

presently annotated as being from “Java.” Schott had 

a drawing (Aroideae Nr. 45) labelled “Borneo?” that 

was prepared from the type. I have annotated the type 

as probably being from Sumatra, the only island of the 

three cited in the original publication in which A. 

pictum is native. Engler (1879, p. 437) reduced A. 

gracile to synonymy under A. pictum. Engler later 

( 1915, p. 33) placed A. gracile as an unpublished name 

under A. sim9lex. This disposition seems mistaken since 

the leaves are variegated and the spathe decurrency is 

very short. I am returning A. gracile to synonymy under A. pictum. 

I designate the plate on page 6 in Bull (1894) as the 

neotype of Aglaonema versicolor. This plate was published subsequently to the original publication but 

clearly represents the plant Bull earlier had in mind. 

Bull’s plant differs from A. pictum only in its variegation: “. . . irregularly blotched with patches of dark 

velvety green interspersed among patches of lighter 

green, and some of milky white.” Thus, the leaves have 

three colors and it may be difficuIt to tell which is the 

basic leaf color. Plants of A. pictum with 3-colored 

leaves with white patches probably are best handled as 

a cultivar. 

The type of Aglaonema pictum var. tricolor is: 

England, Hort. Veitch, September 1886, without collector [N. E. Brown?] (lectotype: K). No specimen 

was cited in the original description, but the name was 

attributed by Engler to N. E. Brown in manuscript. 

This is the only specimen anotated by N. E. Brown as 

A. pictum var. tricolor. Engler (1915) described A. 

pictum var. tricolor as having the blade marked with 

yellowish and yellowish-green spots. This may be truer 

of dried specimens than fresh specimens. A slightly 

7) * 

6. Aglaonema pictum 

Aglaonema pictum (Roxburgh) Kunth, 1841, p. 55. 

Calla picta Roxburgh, 1832, p. 516 [1814, p. 65, nomen 

Aglaonema gracile Schott, 1864, p. 279. 

A. versicolor Hort. Bull [sic], 1893 (Apr. 8), p. 414. 

A. pictum var. tricolor N. E. Brown ex Engler, 1915, p. 33. 

Stems erect, 30-50 cm tall, 0.3-2.0 cm thick. Internodes 0.5-5.0 cm long, longest near the base of the 

plant. Petiole (2.1) 3-7 (14) cm long, 0.3-0.4 (0.6) 

times as long as the leaf-blade. Sheaths broad, hyaline 

to scarious, often slightly prolonged, 1.5-5.0 (13) cm 

long, (0.25) 0.5-0.8 (1.0) times as long as the petiole. 

Leaf-blades dull above, frequently with a crispulate 

margin, narrowly elliptic to elliptic or ovate, rarely 

lanceolate, (5.7) 10-16 (20) cm long, (2.7) 3.5-6.0 

(9.5) cm wide, length/width ratio of 1 : (1.6) 

2.3-3.0 (4.0) ; base often unequal, obtuse to rounded, 

rarely truncate ; apex often apiculate, acuminate; 

variegated or not with irregular light green to silvery 

irregular maculations on a dark green background; 

venation strongly differentiated into 4-8 (10) primary 

lateral veins diverging from the midrib at (30’) 

40’35”. Peduncles 14 together, 3-6 (15) cm long. 

Spathe light green to yellowish, globose, abruptly 

apiculate, 1.5-3.0( 5) cm long, 7 cm wide, decurrent for 

0.2 cm (rarely 0.5-1.0 cm). Stipe 0.3-1.2 cm long, 

elongating in anthesis and fruit. Spadix clavate, exserted from spathe for 0.5 cm in anthesis, 1.4-3.0 cm 

long; pistillate portion 0.2-0.5 cm long, pistils 12-26 

( 15 and 26 on Nicolson 676), stigmas broad as ovary; 

staminate portion 1-2 (2.5) cm long, lower half often 

sterile, 0.3-0.5 cm thick at base, 0.7-0.8 cm thick at 

middle or slightly above middle. Fruits becoming red, 

1-13 cm long, 0.5-1.0 cm thick. 

TYPIFICATroN.-The type is Roxbuqrh’s drawing 

No. 1662 (K) . This drawing was published by Wight 

nudum]. 

326-551 049-3 
30 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

7. Aglaonema nebulosum 

Aglaonema nebulosum N. E. Brown, 1887, p. 67, pl. 24. 

Stem erect, 10-60 cm tall, 0.5-1.0 cm thick. Internodes 0.5-2.0 cm long. Petioles 2-5 cm long, (0.1) 0.2- 

0.4 times as long as the leaf-blade. Sheaths membranous, short, 0.5-1.5 (2) cm long, 0.15-0.3 (0.6) times 

as long as the petiole. Leaf-blades dull above, oblanceolate to narrowly elliptic-oblong, rarely obovate to 

elliptic, (6.5) 9-18 (20) cm long, (2.5) 3-5.5 (7) cm 

wide, lengthlwidth ratio 1 : 2.5-3.5; base obtuse to 

acute; apex often apiculate, acuminate to abruptly 

acuminate; variegation rare, then in the form of ashy 

or silvery irregular blotches on the upper leaf surface; 

venation usually strongly differentiated into 5-10 or 

more pairs of primary lateral veins diverging from the 

midrib at 4O04O0. Peduncles solitary, rarely 2 together, 2-4.5 cm long, equaling the petioles. Spathe 

apiculate, light green to whitish, globose, 1.7-2.7 cm 

long, to 4 cm wide, decurrent for 0.2-0.5 cm. Stipe 

0.3-1.0 cm long. Spadix clavate, exserted from spathe 

for 0.5 cm at anthesis, 1.2-2.0 cm long; pistillate portion 0.3-0.6 cm long, ca. 0.6 cm thick, pistils 10-20; 

staminate portion 0.9-1.5 cm long, 0.3-0.4 cm thick at 

base, O.GO.8 cm wide at or above middle, white. Fruits 

dark red, 0.7-2.0 cm long, 0.4-0.9 cm thick. 

DIsTRIsuTIoN.-Malaya, Borneo, and islands off the 

east coast of Sumatra (Figure 7). 

HABITAT.-Peat and freshwater forests. 

FLOWERING TIM E.-Nonseasonal. 

Aglaonema nebulosum N. E. Brown has been taxonomically confused since its inception. It originally 

was described from vegetative material cultivated by 

Linden and was reported to have originated in Java. 

All available evidence is against this source, for three 

other cultivated specimens sent earlier to Brown by 

Linden and Bull were noted as having originated from 

Borneo, and the species is not known to reach Java. 

It is not clear to me why Brown chose to describe only 

vegetative material and chose to overlook the flowering material of A. nebulosum, which he had already 

prepared. It is this flowering material that makes it 

possible to identify A. nebulosum with later collections 

from the wild and with epithets subsequently applied 

to the species. 

Hooker (1893) did not include A. nebulosum, but 

had three nonvariegated collections of material assignable to this species, each of which he recognized 

as the type of a species: A. minus, A. nanum, and A. 

different definition was given by van Aldenverelt van 

Rosenburgh (1922, p. 327) who reported the leafblade as “rather marbled with dark green, bright green 

and pale-grey-green so that the bright green portions 

are dominant.” 

Aglaonema pictum differs from A. nebulosum; the 

most closely related species, by its longer petiolar 

sheath. In A. nebulosum the sheath rarely is over twofifths as long as the petiole and in A. pictum the sheath 

is rarely less than half as long as the petiole. The species 

also occupy different ranges and habitats. Aglaonema 

nebulosum is found in lowland swamp forests of Malaya 

and Borneo, and A. pictum normally is found at over 

1000 meters on slopes of Sumatran volcanos. 

A number of the specimens appear not to have any 

leaf variegation: Meijer 6431, Rahmat Si Boeea 10262, 

Steenis 6407. I cannot justify taxonomic recognition 

of this nonvariegated material since variegation may 

fade in dried material and I have found no other characters correlating with the lack of variegation. 

Two specimens had definitely lanceolate leaves with 

the length almost four times the width; Steenis 8887 

from Atjeh and Dames 39 from Sibolga. These extreme 

specimens are joined with the more usual specimens 

by intermediates such as Bartlett 8169 and Rahmat Si 

Boeea 6030, in which the lengthjwidth ratios vary 

from 3.0-4.0 on different leaves of the same specimen. 

This indicates that the extremes do not represent 

distinct taxa. 

The spathe of A. pictum normally seems to be quite 

globose at anthesis in fresh material, but this is practically impossible to determine in dried and pressed 

material. The globosity seems to be correlated with a 

short spathe decurrency (0.2 cm) . In nonvariegated 

fruiting material a short spathe scar is very useful in 

distinguishing A. pictum from A. simplex since A. 

simplex typically has the spathe scar 0.5 cm long or 

more in Sumatran material. Unfortunately, some exceptions are known. There are several specimens with 

flowering and fruiting material in which the spadix 

is clearly clavate but the spathe scar is 1 cm long, e.g., 

Rahmat Si Boeea 10262 and Bunnemeijer 4636. 

Rahmat Si Boeea 10262 is extremely anomalous, 

having peduncles 10-15 cm long, a spathe 5 cm long, a 

spathe decurrency of 1.0 cm, leaves 9.5 cm wide, petiole 

14 cm long, and the sheath 13 cm long. All these measurements are extremes for the species. Since the material, however, has a clavate spadix and its ratios fall 

with A. pictum, I have decided to place it here. 
NUMBER 1 31 

scortechinii. Ridley (1907) reduced A. nanum and A. 

scortechinii to varieties of A. minus. Engler (1915), 

who did not cite Ridley’s work, recognized all the four 

species A. nebulosum, A. minus, A. nanum, and A. 

scortechinii, noting that A. nebulosum and A. scortechinii were “Species imperfecte cognita.” Ridley 

(1925) added a new element of confusion when he 

reduced A. minus to synonymy with A. pictum and 

recognized A. nanum and A. scortechinii as varieties 

of A. pictum. Aglaonema pictum is, to be sure, closely 

related to A. nebulosum but seems to be distinct. In 

A. pictum the petiolar sheath is consistently well over 

half the petiole length while in A. nebulosum the sheath 

is rarely over one-third as long as the petiole. Apparently Ridley confused the species because variegation 

patterns in the two species are almost identical. 

Aglaonema vittatum may be confused with A. nebulosum since both have short sheaths and overlapping 

distributions, but the spadix is not exserted in A. vittatum as it is in A, nebulosum. 

Key to the Forms of Aglaonema nebulosum 

Leaves variegated .......................... 7a. forma nebulosum 

Leaves not variegated. ......................... 7b. forma nanum 

7a. Aglaonema nebulosum f. nebulosum 

Aglaonema nebulosum N. E. Brown, 1887, p. 67, pl. 24. 

A. rotundum sensu N. E. Brown, 1893b, p. 379, fig. 56 [not 

A. pictum sensu Ridley, 1925, p. 102 [not Kunthl. 

Leaf variegation in irregularly ashy to silvery blotches 

over the upper leaf surface. 

TYPE coLLEcTIoN.-[Be~gium, Ghent, Hort. Linden] Java, from Mr. Linden, July 1887 [N. E. Brown?] 

(holatype : K) . 

DISTRIBUnON.--SingapOre and southern Malaysia 

(Johore) . It is curious that Brown made no reference to the 

flowering material sent to him before 1887, not only by 

Linden but by Bull. This flowering material, with its 

N. E. Brown, 1893al. 

variegated leaves, short petioles, short sheaths, exerted 

and clavate spadix, and globose spathe, clearly relates 

A. nebulosum to A. nanum, which appears to be only a 

form with nonvariegated leaves. The type seems to have 

somewhat longer leaves than the wild material. The 

type and some of the wild collections do not show their 

variegation after drying. 

7b. Aglaonema nebulosum f. nanum (Hooker f.) 

Nicolson, new status 

Aglaonema nanum Hooker f., 1893, p. 530. 

A. minus Hooker f., 1893, p. 530. 

A. scortechinii Hooker f., 1893, p. 530. 

A. minus var. scortechinii (Hooker f.) Ridley 1907, p. 22. 

A. minus var. nanum (Hooker f.) Ridley, 1907, p. 22. 

A. obovatum van Alderwerelt van Rosenburgh, 1922, p. 322. 

A. pictum var. scortechinii (Hooker f. )Ridley, 1925, p. 102. 

A. pictum var. nanum (Hooker f.) Ridley, 1925, p. 102. 

Leaf variegation none. 

TYPE COLLECTION.-Malaya [Selangor], Ulu Bubong, July 1886, King’s Collector 10421 (holotype: K; 

isotypes: BM, CAL, SING). In this case, Dr. King’s 

collector was Kunstler. 

DIsTRIsuTroN.-Malaysia, Borneo, and islands off 

the eastern coast of Sumatra. Highest elevation reported is 2000 ft (Derry 613) on Mt. Ophir, Malacca. 

FLOWERING TIME.-Appears to be nonseasonal. 

The type of Aglaonema minus is: Singapore, October 1822, Wallich, E. India Co. 89608 (holotype: K) . 

The type is in abortive fruit and the smallish leaves 

have three to four pairs of weakly differentiated veins. 

I regard this specimen as a poorly developed individual 

of A. nebulosum, which generally has six or more 

rather strongly differentiated pairs of veins. 

The type of Aglaonema scortechinii is: Perak [no 

date], Scortechini 255 (holotype : K, isotype : CAL) . 

FIGURE 11 .-Holotype of Aglaonema nebulosum N. E. Brown : 

N. E. Brown? (K). 
32 

The Kew specimen is regarded as the holotype since 

it was annotated by N. E. Brown, who assisted Hooker 

with Araceae for the Flora of British India. The leaves 

of this specimen are ovate (2.0-2.4 times longer than 

broad) rather than oblanceolate ; however, some of 

these lower leaves are broader than the upper, more 

vigorous leaves, making it difficult to justify maintenance of A. scortechinii solely on the basis of leaf shape. 

The type of Aglaonema obovatum is: Indonesia, 

Lingga Archipel, Pulo Lingga, Kampong Panggak, 30 

m, Sago bosch, 27 July 1916, Biinnemeijer 7039 (holotype: BO). Alderwerelt van Rosenburgh did not 

comment about the relationships of his species nor 

indicate what was distinctive about it. I cannot maintain it since the type is adequate and falls completely 

within my concept of A. nebulosum f. nanum. 

8. Aglaonema vittatum 

Aglaonema vittatum Ridley ex Engler, 1915, p. 33. 

Stem 10-15 cm tall, 0.2-0.7 cm thick in dried material. 

Internodes 0.4-1.4 cm long. Petioles 2-4.2 cm long, 

0.3-0.6 times as long as the leaf-blade. Sheaths 0.8-1.2 

cm long, 0.2a.4 times as long as the petiole. Leafblade lanceolate, 6.5-10.5 cm long, 1.7-3.0 cm wide, 

length/width ratio 1 : (2.6) 3-4.4; base often unequal, 

obtuse ; apex gradually attenuate ; variegation none 

or in form of a narrow central white stripe along 

the midrib; venation differentiated into 2-3 primary 

lateral veins diverging from the midrib at 30’. Peduncle solitary, 2.5-3.0 cm long (4-5 cm, fide Engler, 

1915). Spathe 2-3 cm long, decurrent for 0.3-0.5 cm. 

Stipe 0.1-0.4 cm long. Spadix clavate, about half the 

spathe length (fide drawing at Kew) ; pistillate portion 

with ca. 10 pistils. Fruits red. 

TYPE coLLECnoNs.-Indonesia, Lingga, Batu Gajah, 1-2000 ft, 19 July 1893, Hullett s.n. (lectotype: 

SING) ; Indonesia, Sungei Kelantan, Siak, November 

1897, Ridley s.n. (syntype: SING). 

DrsTRIsuTIoN.-Restricted to eastern Sumatra and 

the Lingga Archipelago (Figure 6). 

HABITAT.-h primary rain forest from a few meters 

above sea level (Buwalda 6629) to 300400 m 

(Hullett s.n.) . 

FLOWERING TIME.-July-August on the few specimens available. Seasonality is not expected since all 

collections are from lowlands and within one degree 

of the equator. 

Some of the collections show no evidence of the 

white midrib. Teysmann s.n. (BO 100514) is made up 

SMITHSONIAN CONTRIBUTIONS TO BOTANY 

of four plants; two are variegated, one is not, and I am 

uncertain about the fourth. Biinnemeijer 7161 is 

variegated and Biinnemeijer 7173 is not although both 

were collected the same day, at the same elevation, and 

in the same locality. 

Aglaonema uittatum is most similar to A. rotundum, 

judging by the unpublished drawings of the inflorescence at Kew. It resembles A. nebulosum in its short 

petiole and sheath but differs by its much narrower 

leaf. 

9. Aglaonema rotundum 

Aglaonema rotundum N. E. Brown, 1893a, p. 86. 

Stem decumbent, erect for the terminal 5-6 cm, 0.3- 

0.9 cm thick. Internodes 1 cm long. Petioles 2.54.5 cm 

long, 0.3-0.4 times as long as the leaf-blade. Sheaths 

membranous, broad, frequently prolonged for 0.5 cm, 

1-3.5 cm long, 0.4-0.8 times as long as the petiole. 

Leaf-blade broadly ovate, 6-13.5 cm long, 5.8-10.6 cm 

wide, length/width ratio 1 : 1.0-1.3; base obtuse to 

truncate; apex slightly apiculate, obtuse; variegation 

in form of whitish or pink midrib and veins on a dark, 

glossy green upper surface, lower surface dark red; 

venation differentiated into 4-6 primary lateral veins 

diverging from midrib at 50’-60’ ; texture rather coriaceous. Cataphylls to 3.5 cm long, like brown-paper, 

mixed with petioles. Peduncle solitary, 6.5-7.0 cm 

long, exceeding the petioles. Spathe large 6-7 cm long, 

bent forward, decurrent for 0.6-1.5 cm. Stipe present? 

Spadix clavate, 3 cm long, bent forward; pistillate portion 0.6 cm long, pistils in 2-3 series (ca. 15?) ; staminate portion 2.5-3.5 cm long, 1.3 cm thick (fide 

Brown). Fruits 0.8 cm long, 0.6 cm thick. 

TYPE cOLLEcTIoN.-England, Hort. Veitch, 4 July 

1893 [N. E. Brown?] (holotype : K) , 

DISTRIBUTION. (Figure 6) .-The only collection 

(Bartlett 6739) definitely taken from wild-growing 

material is from near Medan in North Sumatra. 

Huttleston 1907 reports his plant was collected by W. 

H. Hodge in Thailand, but it may have been cultivated there. Graf (1963, p. 1527) states that he 

found the plant in northern Thailand, but again it may 

have been from cultivation. The earliest collection was 

made from Hort. Veitch on 27 July 1889 and was 

reported to have been “imported from Calcutta without information.” On the holotype N.E. Brown wrote, 

“Native country unknown, the plants were received by 

Mr. Veitch from Calcutta and Penang.” 

HABITAT.-Near a waterfall, 360 m (Bartlett 6739). 
NUMBER 1 33 

10. Aglaonema nitidum 

Aglaonema nitidum (Jack) Kunth, 1841, p. 56. 

Calla nitida Jack, 1820, p. 24. 

Stem erect or with lower part reclining on ground in 

larger plants, to 1 m or taller, 0.5-5 cm thick. Internodes 0.5-2.0 cm long. Petioles (8) 11-26 (29) cm 

long, 0.4-0.9 times as long as the leaf-blade, often in 

two spirals. Sheaths with scarious margins, (5) 9-22 

(28) cm long, (0.4) 0.7-1.0 times as long as the 

petioles. Leaf-blades narrowly elliptic to narrowly 

oblong or oblanceolate, ( 11 ) 20-45 (50) cm long, (4) 

7-16 (20) cm wide, length/width ratio 1 : (2.1) 2.7- 

3.6(4.3) ; base acute to attenuate, rarely broadly acute; 

apex often apiculate, acuminate to broadly acute or 

shortly acuminate; variegation usually not present; 

when present, either in bars following the venation or 

in rather irregularly scattered blotches; venation usually undifferentiated in dry material but sometimes 

weakly differentiated into 5-9 or more primary veins 

diverging from the midrib at 35 "-55" ; texture coriaceous. Peduncles rarely solitary, 2-5 together, (5) 

10-17 (21) cm long, commonly equaling or surpassing the subtending petiole. Spathe light green but frequently turning white with age, eventually withering, 

3-7 (8.5) cm long, decurrent for (0.4) 1-1.6 (2.0) 

cm. Stipe 0.2-0.9 cm long. Spadix cylindric, equaling 

or slightly exceeding spathe, (1.3) 4-7 cm long; 

pistillate portion (0.2) 0.5-1.0 cm long (to 2 cm in 

fruit), pistils 16-37, scattered at anthesis, stigmas 

yellowish, to 0.3 cm broad, the style weakly constricted; 

staminate portion ( 1.1) 2.5-5.0) (6.0) cm long to 

1.5 cm thick, commonly rotting or shriveling soon 

after anthesis. Fruits white. 

DIsTRrBUTI0N.-southern Burma, Malaysia, Sumatra, and Borneo (Figure 2) . HABITAT.-Lowland primary dipterocarp forest in 

damp and well-shaded place. Found in freshwater 

peat-swamp forests and on limestone but in pockets 

and damp ravines. 

FLOWERING TIME.--NOnSeaSOnal. The species is 

largely restricted to the equatorial region, where rainfall is daily (convectional) rather than seasonal and 

the plants live in dark places where the humidity is 

always high. 

There are two important taxonomic problems in 

Aglaonema nitidum. The first problem involves the 

northwestern extremity of the distribution of the 

species. In that area the species appears to undergo 

FIGURE 12.-Holotype of Aglaonema rotundum N. E. Brown: 

N. E. Brown? (K). 

USES.-A very desirable foliage plant. In Thailand 

I was shown a wellgrown specimen in a home and was 

told that the plant brings good luck to the owner. In 

the market of Bangkok I saw a depauperate specimen 

for sale for what seemed to be an extremely high price. 

A living specimen was given to me by Prof. Kasin 

Suvatabandhu of Chulalongkorn University, but I regret to note that it has disappeared fron my collections. 

The inflorescence disintegrates by the second day 

(Huttleston 1907) after opening and it is not possible 

to determine from material available to me whether or 

not there is a stipe. On Huttleston 1907 there appears 

to be a stipe, but this may be the result of distortion 

in pressing. In his original description Brown noted 

the presence of a stipe, but he may have been referring to the naked portion of the spadix rather than to 

the stipe as the term is used here (measured next to 

the spathe). I assume, with doubts, that it is stipitate. 

The wild specimen is much smaller than the cultivated material and has shorter petiolar sheaths. This 

is not considered to be of significance since both characters may be influenced by the care given to cultivated 

material. 

The spadix is much shorter than the spathe. In this 

respect A. rotundum is similar to A. uittatum. 

The presence of cataphylls among the petioles and 

the decumbent habit is very suggestive of section 

Chamaecaulon. 
34 

several tendencies : smaller size, shorter petiolar 

sheath, loss of the scarious petiolar margin, membranous rather than a coriaceous texture, more ovate 

leaves with rounded leaf-bases, and fewer pistillate 

flowers. The plants of central Thailand, Cambodia, 

and South Vietnam are so strongly differentiated in 

these characters that they could be maintained as a 

distinct species, A. cochinchinense. The plants of 

southern Burma, however, show only enough differentiation to justify recognition as a variety, A. nitidum 

var. helferi. Further collections from southern Burma 

and western Thailand would be very desirable to 

clarify the complex. 

The second problem involves the northeastern extremity of the species. In the Philippines there seems 

to be some overlapping with the A. philippinenseSMITHSONIAN CONTRIBUTIONS TO BOTANY 

densiiiervium-stenophyllum-crispum complex. Some of 

the distinctive characters of A. nitidum (as opposed 

to other common species, such as A. simplex and A. 

marantifolium) are found in material from the Philippines, i.e., acute leaf base, long petiolar sheaths, and 

scarious sheath margins; however, these characters are 

either not in combination or, when they are, they are 

further combined with characters not at all typical of 

A. nitidum. This had led me to consider that the 

Philippine complex may have a hybrid origin in which 

A. nitidum, along with A. simplex and A. marantifolium, may have had a part. True A. nitidum, as 

known in Borneo, Malaya, and Sumatra, occurs in 

the Philippines only on Balabac Island (which is, 

incidentally, also the farthest penetration of the mouse 

deer [chevrotain] into the Philippines) . 

Key to the Varieties and Forms of Aglaonema nitidum 

1. Spathe (3.5) 4.5-8.5 cm long ...................... var. nitidurn 

2. Leaf-blade not variegated ....... .. 

2. Leaf-blade variegated ......... 

1. Spathe 3.0 cm long ........... 

10a. Aglaonema nitidum var. nitidum f. nitidum 

Calla nitida Jack, 1820, vol. 1, no. 1, p. 24. 

Arum integrifolium Link, 1822, p. 394. 

Aglaonema oblongifolium Schott, 1829, p. 829. 

A. integrifoliurn (Link) Schott, 1832, p. 20. 

A. nitidurn (Jack) Kunth, 1841, p. 56. 

A. princeps Kunth, 1841, p. 55. 

A. oblanceolaturn van Alderwerelt van Rosenburgh, 1922, 

A. oblanceolatum f. maximum van Alderwerelt van RosenPetioles (8.5) 11-26 (29) cm long, 0.4-0.9 times as 

long as the leaf-blade. Sheaths (7) 10-20 (28) cm 

long, 0.6-1 .O times as long as the petioles. Leaf-blades 

(19) 25-45 (50) cm long, (5) 7-16 (20) cm wide, 

length/width ratio 1 : (2.1) 2.7-3.6(4.3) ; variegation 

sometimes seen on juvenile leaves. Peduncles (8.5) 

10-17 (21 ) cm long. Spathe 3.5-8.5 cm long, decurrent 

for (0.8) 1-1.6 (2) cm. Spadix (2.5) 4.5-6.0 (6.9) 

cm long; pistillate portion 0.3-1.0 cm long, pistils (10) 

16-37 ; staminate portion 2-6 cm long. 

TYPE coLLEcTIoN.-Malaya, Penang, August 1822, 

Wallich, E. India Co. 89654 (neotype: K). 

DISTRIBUTION.-PeninSUlar Thailand, M a 1 a y a , 

Sumatra, and Borneo. 

HABITAT.-LowlandS, in damp and shady places. 

FLOWERING TIME.-Nonseasonal. 

A neotype must be selected for Calla nitida Jack. All 

of Jack’s types have been elusive since the bulk of his 

p. 322. 

burgh, 1922, p. 323. 

............ 10a. forma nitidurn 

............ lob. forma curtisii 

1Oc. var. helferi ............. 

collections burned with the ship Fame in 1824. Cowan 

( 1954) reported on newly recognized Jack specimens 

at Edinburgh, but none of these were Araceae. The 

classical commentary on Jack is Burkill’s (1916) publication of the letters written by Jack to Wallich from 

1819 to 1821. Jack‘s letter of 7 May 1819 from Penang 

includes the following: “Capt. Fergusson of the Boyne, 

has agreed to take charge of a box of growing plants, 

and a parcel of specimens, which hope will arrive safe.” 

Although nothing is said about aroids in the parcel of 

dried specimens, Jack continues, “I hope the box of 

plants will please you, it contains a good many which 

I have never seen in flower .... There are besides, 

two species of Arum, one of Calla ... .” Jack ( 1820) 

described three species of Araceae from Penang: Calla 

[Homalomena] humilis, Calla [Homalomena] angustifolk, and Calla [Aglaonema] nitida. I believe that all 

three of these species (two called Arum at that time) 

were included in that shipment of living plants. Burkill 

(1916) tends to support this view, saying that the “one 

of Calla [is] Aglaionema [sic] oblongifolium Schott” on 

page 174 although on page 268 he says it is Aglaonema 

marantifolium Blume. This second determination 

seems to be part of the nomenclatural confusion involving the type-species. It is not likely that Burkill intended 

to refer to a Moluccan species. 

It seems probable that living material of Calla nitida 
NUMBER 1 35 

FIGURE 13.-Painting of Aglaonema nitidum (Jack) Kunth, 

neotype of Aglaonema oblongifolium Schott and of Aglaonema integrifolium (Link) Schott: Schott, Aroideae Nr. 59. 

Jack was sent by Jack from Penang to Wallich in Calcutta, but there is no record there of a dried specimen 

from Jack. Study of the material of the East India 

Company, which was distributed by Wallich, shows 

that there are two collections from Penang that, apart 

from mixed material, belong to what is now known as 

Aglaonema nitidum (Jack) Kunth. Neither was collected by Jack. Porter, E. India Co. 4443 was labelled 

as Pothos marginatus Wallich (nomen nudum) . According to the numerical list of the East India Cornpany Museum, issued by Wallich (1828-1849), No. 

4443 was collected by “G. P.” [George Porter], who 

travelled with Wallich to Penang in 1822 and stayed 

to start the Penang Botanic Garden; however, material at Brussels of Porter, E. India Co. 4443 is a mixture of three species, Aglaonema nitidum (Jack) 

Kunth, Anadendrum marginatum Schott, and an 

aquatic plant that is not an aroid. According to Engler 

(1905, p. 47), “Wallich, 4443 - Herb. Kew u. a. [und 

anderny’ is Anadendrum marginatum Schott. Porter’s 

collection, distributed by Wallich as E. India Co. 4443, 

need not be further considered for neotypification of 

Calla nitida Jack since it is a mixed collection. 

The only other specimen of Aglaonema nitidum 

(Jack) Kunth in the East India Company’s herbarium 

from Penang is No. 89658. According to Wallich 

(1828-1849)) No. 8965A was collected in Penang in 

August 1822. The collector’s name is not given and 

therefore it is presumed that it is Wallich‘s own collection. It is known that Wallich was in Penang in 

August 1822 (Steenis-Kruseman, 1950, p. 557). 

This neotypification is the same as that inferentially 

used by previous workers. Schott (1860, p. 302) cited 

only one specimen under Aglaonema nitidum: “Pulo 

Penang. Wallich-V.V. et s. sp.” Engler (1879, p. 

438) cited “India orientalis; Pulo Penang (Jack, Wallich n. 8965) . . . .” 

A neotype must be designated for Arum integrifolium Link. In describing this species from cultivation in the Berlin Botanic Garden, Link (1822) did not 

cite any type. There is no herbarium specimen that 

can be connected with the original description ; however, there is every indication that Link‘s Arum 

integrifolium was Aglaonema nitidum: the description 

spoke of the leaves being attenuate at the base. I have 

seen two specimens and one watercolor painting made 

from plants cultivated in the Berlin Botanic Gardens, 

all of which have the epithet “integrifolium” appearing 

on them, and all of which are identifiable with 

Aglaonema nitidurn. Previous monographers of the 

family, i.e., Schott (1860, p. 303) and Engler (1879, 

p. 438), have treated Arum integrifolium as a synonym 

of Aglaonema nitidum. It seems best to select the 

watercolor Schott, Aroideae Nr. 59 (W) as neotype 

since the description fits it well and the specimen is 

annotated “Ar. integrifolium Hort. Berol.” Selection 

of this neotype also settles a nomenclatural problem involving the type-species of the genus Aglaonema, as 

explained below. 

Aglaonema oblongifolium Schott is the type-species 

of the genus Aglaonema. Beginning with Hooker’s 

( 1893, p. 529) unwarranted inclusion of Calla oblongifolia Roxburgh as a synonym of Aglaonema oblongi- 
36 

folium Schott, most botanists have, in effect, typified 

the Schottian binomial on the type of the Roxburghian 

binomial. This was easy to assume since the epithets 

were identical. Study of the original descriptions and 

reliquiae, however, clearly shows that each epithet has 

its own type, that each pertains to a different species, 

and that it was only by coincidence both authors chose 

the same descriptive epithet. The typification of Calla 

oblongifolia Roxburgh is discussed under Aglaonema 

marantifolium Blume and the typification of Aglaonema oblongifolium Schott follows. 

In describing Aglaonema oblongifolium from material cultivated at Schoenbrunn Garden in Vienna, 

Schott did not cite any specimens; however, he did cite 

“Arum integrifolium Hort. Berol.,” which may be an 

indirect reference to the earlier binomial Arum 

integrifolium Link (1822) or it may only be a reference 

to the label name and the source of the plant at 

Schoenbrunn. 

Supposing Aglaonema oblongifolium Schott is not a 

superfluous name for Arum integrifolium Link, we 

must select a neotype. The entire Schottian herbarium, 

with the exception of one bundle, was destroyed after 

the World War 11, but a collection of 3282 drawings 

and paintings of Araceae was saved. There are only 

four drawings pertinent to Aglaonema nitidum, and 

of these none is labelled Aglaonema oblongifolium 

Schott although one is labelled “Ar. integrifolium Hort. 

Berol.” That one is “Schott, Aroideae Nr. 59,” which 

I designated above as the neotype for Arum integrifolium and now designate as the neotype of Aglaonema 

oblongifolium Schott also. The designation of the 

same neotype for these two binomials so links them 

that Aglaonema oblongifolium Schott must be regarded as a superfluous name. Actually, Schott seemed 

to regard his own epithet as superfluous since he almost immediately (1832, p. 20) published the correct 

combination, Aglaonema integrifolium (Link) Schott, 

and never again referred to his original publication. 

Aglaonema princeps Kunth was described from material cultivated in Berlin labelled “Caladium princeps 

Hort. Berol.” Schott (1860, p. 303) treated it as a 

synonym of Aglaonema nitidum; however, Engler 

(1879, p. 439) transferred it later to synonymy under 

Aglaonema simplex. The only material extant today 

is a drawing of the inflorescence with dissections, 

Schott, Aroideae Nr. 70 (W) . The pistillate flowers 

are scattered like those of A. nitidum rather than A. 

simplex. The description of a plant with an acute leaf 

SMITHSONIAN CONTRIBUTIONS TO BOTANY 

base and a petiole that is sheathed almost to the apex 

clearly pertains to A. nitidum rather than A. simplex. 

I see no reason to accept Engler’s treatment and am 

following Schott in placing A. princeps under A. 

nitidum. 

Aglaonema oblanceolatum was based on Lorring 

5352 (BO, L) and Lorzing 5887 (BO) , both from 

Sibolangit, Sumatra. I designate Lorring 5352 [Herbarium Bogoriense 1003551 (BO) as lectotype, with 

Lorzing 5352 [Herbarium Bogoriense 1003561 (BO, L) 

as isolectotypes, leaving Lorzing 5887 (BO) as a syntype. I can see no reason for this material to be regarded 

as distinct from Aglaonema nitidum. The author evidently had no idea of the identity of Aglaonema nitidum although he stated: “Probably very near A. oblongifolium Kunth [which was true, sensu Engler, not 

Kunth].” 

Aglaonema oblanceolatum f. maximum was described from cultivated material at Bogor. Search of the 

Gardens Herbarium revealed the holotype, van Alderwerelt van Rosenburgh 372 (BO), which had been 

collected from “XI. B. IX. 113.” Study of the Garden 

Register showed that material designated as “XI. B. 

IX. 113” had originally been collected by Docters 

van Leeuwen from Sibolangit, Sumatra, in 1919, and 

that in October 1923, it had been transferred to 

“XI. B. IX. 166.” Any subsequent collections from 

“XI. B. IX. 166” such as Nicolson 994 may be regarded 

as clonotypes. 

The inflorescence of A. nitidum var. nitidum can be 

identified by its many scattered pistils on a spadix equaling the spathe, which is long-decurrent. The leaves are 

also distinctive by their long, scarious petiolar sheaths, 

and blades more or less acute at the base with undifferentiated lateral venation. This is the only species 

that often has a white spathe in fruit and white berries. 

lob. Aglaonema nitidum var. nitidum f. curtisii 

Aglaonema nitidum var. nitidum f. curtisii (N. E. Brown) 

A. oblongifolium var. curtisii N. E. Brown, 1897, p. 70. 

Leaf-variegation of ashy or silvery stripes along the 

courses of ten or more lateral veins or irregularly 

scattered. 

TYPE coLLEcTIoN.-England, Hort. Veitch, from 

Penang by Mr. Curtis, 22 November 1896 [N. E. 

Brown?] (holotype: K) . 

Nicolson, 1968, p. 126. 

DISTRIBUTION .-Malaya and Lingga Archipelago. 

USES.-Popular in conservatories for its variegated 

leaves. 
NUMBER 1 37 

This form differs from the typical form only in having variegated leaves. Occasional collections do not 

show the usual striped variegation known in cultivation but have rather irregularly blotched leaves. 

In the original publication, N. E. Brown spoke of a 

plant at Kew that “had one leaf near the basal part of 

the stem variegated in exactly the same way as Messrs. 

Veitch & Son’s plant, and all the other leaves on the 

plant were of the usual deep glossy green of the typical 

form.’’ In the wild I have also observed variegation 

restricted to juvenile and seedling plants (Nicolson 

2246). Perhaps the variegated form only represents 

perpetuation of this occasional characteristic of 

juvenile plants. 

1Oc. AgZaonerna nitidurn var. heZferi (Hooker f.) 

Nicolson, new status 

Aglaonema helferi Hooker f., 1893, p. 529. 

Petioles 11.5-12.5 [15, fide Hooker] cm long. Sheaths 

5-9 crn long, 0.4-0.7 times as long as the petioles. 

Leaf-blades [15, fide Hooker] 21-25 cm long, 5.6-7.5 

cm wide, length/width ratio of 1 : 3.3-4.0. Peduncles 

5-8 cm long. Spathe 3 cm long, decurent for 0.4-0.5 

cm. Spadix 1.3-1.7 cm long; pistillate portion 0.2 cm 

long, pistils ca. 4-16; staminate portion 1.1-1.5 cm 

long. 

TYPE COLLECTION.-Burma, Tenasserim [no date], 

Helfer, E. India Co. 5994 (holotype: K). 

DrsTRIsuTIoN.-Southern Burma. 

This variety approaches A. cochinchinense by its exceedingly short spathe; however, it is more similar to 

A. nitidum by its scarious petiolar sheath margins and 

acute to attenuate leaf-bases. Short spathes are known 

in A. nithidum (3.5 cm long in Korthals s.n. from 

Borneo). The number of specimens available is insufficient to make firm decisions, but what is available 

does not justify maintenance of A. helferi as a species. 

Further collections and study in southern Burma may 

shed light on its status. 

1 1. Aglaonema cochinchinense 

Aglaonema cochinchinense Engler, 1915, p. 16. 

A. cambodianum Gagnepain, 1941, p. 116. 

Stem erect, 1-1.5 cm thick. Internodes 0.4-1.0 cm 

long. Petioles (8) 10-18 (22) cm long, (0.4) 0.6-0.8 

(0.9) times as long as the leaf-blade. Sheaths with 

membranous margins, (3) 5-10 (13) cm long, 0.3-0.7 

times as long as the petiole. Leaf-blades ovate or 

lanceolate to elliptic to narrowly elliptic, (1 1) 15-25 

(28) cm long, (4) 5.5-10 (12.5) cm wide, length/ 

width ratio 1: 2.1-3.2; base obtuse to rounded, frequently unequal; apex obtuse to acute, sometimes 

apiculate; variegation none; venation undifferentiated; 

texture membranous. Peduncles 1-2 together’ (6) 

9-15 cm long, frequently exceeding the subtending 

petiole. Spathe green, 3-5 cm long, often apiculate, 

decurrent for (0.5) 1-1.5 (2) cm. Stipe 0.3-1.0 cm 

long (0.1 in Kerr 20525). Spadix cylindrical, 1.9-3.2 

cm long, often exceeding the spathe at anthesis; pistillate portion 0.4-0.7 cm long, the pistils ca. 10, 

scattered; staminodia occasional; staminate portion 

white, tapering. 1.5-3.0 cm long, 0.4 cm thick. Fruits 

red, to 2 cm long and 1 cm thick. 

TYPE coLLEcTIoN.-Cochinchina, Pierre s.n. (holotype: B) . DIsTRlsuTIoN.-southern Thailand, Cambodia, 

and South Vietnam (Figure 4) . HABITAT.-Lowland rainforest, on limestone or calcareous soil. 

FLOWERING TIME.-March-July. 

Aglaonema cochinchinense appears to be most 

closely related to A. nitidum, both of which have undifferentiated venation and spadices that equal or 

exceed the spathe at anthesis. The principal differences from A. nitidum are: smaller size, shorter petiolar 

sheath, membranous petiolar sheath margins, lack of 

the thin, translucent margin of the leaf-blade, membranous leaf texture, and few (ca. 10) pistils. Aglaonema nitidum var. helferi, however, seems to be intermediate between A. nitidum and A. cochinchinense. 

It is possible that A. cochinchinense should be given 

subspecific rank under A. nitidurn, but it seems well 

enough differentiated, even in upper peninsular Thailand, where the two species overlap, to justify its maintenance as a distinct species. 

The type of Aglaonema cambodianum is: Cambodge, Pursat, 14 March 1914, Chevalier 3201 I (holotype: P) . As illustrated by Gagnepain ( 1942, fig. 107), 

the spadix exceeds the spathe by almost half its length. 

Other specimens do not show this. In differentiating A. 

cambodianum from A. cochinchinense in his key to 

species, Gagnepain (1942, p. 1120) indicated that A. 

cambodianum has a rounded to obtuse leaf-base while 

A. cochinchinense has an attenuated leaf-base. This 

separation is not supported by material of the typecollection of A. cochinchinense, which I found to have 

rounded to obtuse leaf-bases with occasional, smaller 

leaves having an acute leaf-base. At any rate, three of 

the four specimens cited by Gagnepain as A. cochinchinense have rounded leaf-bases. 
38 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

explicable in terms of their difference in developmental 

stages, A. siamense being in ripe fruit and A. tenuipes 

being in bud. Vegetatively they differ principally in: 

(1) leaf shape (A. siamense having leaves with a 

lengthjwidth ratio of 1 : 2.5-2.9, and a truncate base; 

A. tenuipes having leaves with a length/width ratio 

1 : 3.0-3.4, and an obtuse leaf base) ; and (2) petiolar 

sheath length (A. siamense having the sheath 8 to 

10 cm long [about 0.7 the petiole length] and A. 

tenuipes having the sheath 3 to 4 cm long [about 

0.4 the petiole length]). Neither difference is significant. Leaf shape appears to be variable in the 

pertinent plants of Aglaonerna from Koh Chang. 

Flowering material (Kerr 6799) from Klong Mayon 

on Koh Chang (about one mile from Klong MunsC) 

includes duplicates that vary in their length/width ratio 

from 1 : 2.4-4.0. Length of petiolar sheath also seems 

to be rather variable. Nicolson 1616 has (from typelocality) petiolar sheaths 6-7 cm long, almost halfway 

between those described for A. siamense and 

A. tenudpes. 

Gagnepain (1942) described the spathe of A. 

siamense as 3.5-4.5 cm long. Study of specimens cited 

shows that this excessively long spathe is based on Kerr 

20899 from Laos, which I identify as A. ovatum. 

I see no reason to maintain A. siamense and A. 

tenuipes as sympatric species. Both epithets have equal 

priority. No one previously has united them. I am 

adopting the name A. tenuipes since it is based on 

flowering material more easily identifiable with other 

specimens than the fruiting specimens typifying A. 

siamense. 

The types of A. pierreanurn are: Cochinchina, 500 

m, June 1877, Pierre s.n. (lectotype: P) ; Bienhoa, 

March 1877, Harmand, Herb. Pierre 1936 (syntype: 

B). The many Pierre specimens of A. tenuipes are 

mostly incompletely labelled and many have been 

determined by Engler as A. pierreanum. In choosing a 

lectotype, one can consider only material actually cited 

in the description, restricting the choice to the two 

cited above. I have chosen the first specimen as lectotype since it has flowering material that is necessary to 

recognize A. tenuipes definitely. One reason for not 

selecting the second specimen is that the description 

depicted the leaves as being 6-8 cm wide whereas the 

leaves on the second specimen are only 4.5 cm wide. 

Engler (1915) regarded A. pierreanum as distinct from 

A. tenuipes. He placed A. pierreanum in his key as 

having the primary veins more or less incised above 

and highly prominent below, while he placed A. 

12. Aglaonema tenuipes 

Aglaonema tenuifies Engler, 1902, p. 275. 

A. siamense Engler, 1902, p. 275. 

A. pierreanum Engler, 1915, p. 24, fig. 8. 

A. subfalcatum Engler, 1915, p. 23. 

Stem erect, 40-150 cm tall, 0.6-2.5 cm thick. Internodes 0.5-2.0 cm long. Petioles (5) 7-1 7 (22) cm long, 

0.5-0.8 times as long as the leaf-blade. Sheaths membraneous, (2.5) 5-8 (15) cm long, (0.4) 0.5-0.7 (0.8) 

times as long as the petiole. Leaf-blades ovate or elliptic 

to lanceolate, frequently unequal or falcate, (6) 15- 

25 (35) cm long, (2.4) 5-13 (21) cm wide, length/ 

width ratio 1 : (1.7) 2.3-3.2(6.7) ; base truncate to subacute, commonly rounded, sometimes unequal ; apex 

usually gradually acuminate, sometimes acute or 

acuminate ; variegation none ; venation strongly (sometimes weakly) differentiated into 4-8 primary lateral 

veins diverging from the midrib at (30") 40"-50" 

(60" ) ; texture membranaceous to subcoriaceous. Peduncles 1-5 together, 2.5-8.0 (13) cm long, deflexing 

in fruit. Spathe green, 1.8-2.5 (4 in Kerr 7010) cm 

long, to 3 cm wide, often apiculate, decurrent for 

0.3-1.0 cm. Stipe 0.2-1.0 cm long. Spadix short-cylindric to subclavate, often slightly exceeding the spathe, 

1.4-2.5 cm long; pistillate portion 0.2-0.7 cm long, 

pistils 16-19 or more (only 3 in Kerr 19940) ; staminate portion white, sometimes separated from pistillate 

portion by small gap (as in Kerr 3005), 0.7-2.9 cm 

long, 0.3-0.8 cm thick. Fruits turning red, 1.5 cm long 

and 0.8 cm thick. 

TYPE COLLECTION .-Thailand, Koh Chang, Klong 

Munst, 8 February 1900, Schmidt 451a (holotype: C) . 

DISTRIBUTION.-Southern continental Southeast 

Asia from South Vietnam, Laos, and Cambodia to 

peninsular Thailand (Figure 6) . 

HABITAT.-In evergreen forest up to 1000 m in damp 

areas. 

FLOWERING TIME.-February-April. 

The type of Aglaonema siamense is: Thailand, Koh 

Chang, Skoven n. f. Lem Dan, 6 January 1900, 

Schmidt 90 (holotype: C; isotype: B) . Engler (1902, 

p. 275) erred in describing the leaves as 2.5 cm wide, 

implying a length/width ratio of 1 : 8-10. In the type 

collection the leaves are 8.5 cm wide with a length/ 

width ratio of 1 : 2.7. 

The types of Aglaonema siamense and A. tenuipes 

were taken from the same general locality (within three 

miles of each other). In the original publication both 

were cited as being from "jungle near Lem Dan." The 

differences in their reproductive structures are largely 
NUMBER 1 39 

tenuipes among the species with primary veins slightly 

stronger than the secondary veins. Reliance on venation alone and without further supporting characters 

is not justified since individual specimens show much 

variation. Aglaonema tenuipes generally has rather 

strongly differentiated venation but occasionally this is 

not the case. I believe this to be more environmentally 

than genetically determined. 

The type of A. subfalcatum is: France, Paris, Jardin 

des Plantes, without collector s.n. (holotype: B) . This 

species, represented only by the type collection taken 

from cultivated material probably originating from 

South Vietnam (fide Engler, 1915), was placed next to 

A. pierreanum Engler. The difference between these 

two species, published simultaneously, was that A. 

pierreanum was supposed to have a cordate leaf base 

and A. subfalcatum an acute or rounded one. This distinction does not exist on the types. In the description 

of A. pierreanum, Engler was much more accurate 

when he described the leaf base as “emarginata vel 

subcordata.” In the published illustration of A. pierreanum the leaf bases are only rounded, definitely not 

cordate. 

Gagnepain (1942) separated A. pierreanum from A. 

subfalcatum, A. siamense, and A. tenuipes on the erroneous assumption that the spadix did not surpass the 

spathe in A. siamense, A. subfalcatum, and A. tenuipes. 

This distinction is not borne out by the types of 

A. subfalcatum and A. siamense, in which the spadix 

clearly surpasses the spathe although it appears to be 

true in the type of A. tenuipes. The fact, however, that 

the spathe is closed and that the peduncle is so short 

indicates that the type of A. tenuipes is in bud and that 

the stipe and spadix may not have fully elongated. 

Gagnepain (1942) regarded A. tenuipes as most 

closely related to A. subfalcatum since both had more 

or less acute leaf bases, the main difference being that 

A. subfalcatum had an oblique leaf base and A. tenuipes 

did not. It seems to me that the vegetative dissimilarities emphasized by Engler and Gagnepain are outweighed by the reproductive similarities. Therefore, 

I reduce all three binomials to A. tenuipa. 

The following variations are of note since they 

represent extremes of morphological variation within 

A. tenuipes. I have not accorded them formal taxonomic recognition since intermediates were frequent 

and widely scattered. Nicolson 1625 and 1638 have 

some of the broadest leaves observed in the species, 

15-20 cm wide with length/width ratios less than 1 : 2. 

Evrard 478 has the narrowest leaves observed in the 

species, to 2.6 cm wide with length/width ratio up to 

1 : 6.7. A number of plants seem to be rather smaller 

than usual for the species: Kerr 10765, Put 1444, Kiah, 

SFN 35244, Put 1615, Rock 499, Pierre s.n. (24 February 1877), and Put s.n. (20 June 1926.) One specimen (Kerr 19440) seems quite anomalous in that it has 

only three pistils in the one. inflorescence I studied. 

Aglaonema tenuipes obviously is closely related to 

A. simplex but may intergrade with A. ovatum. At 

times the stipe seems very short (approaching that of 

A. ovatum). There is a possibility that A. tenuipes is 

only a geographical subspecies of A. simplex. These 

species are separable only when the spathe is present. 

Fruiting material has been identified from geographical evidence; for instance, all flowering material from 

Koh Chang has the short spathe and blunt spadix of 

A. tenuipes; the type of A. siamense is from Koh Chang 

but is in fruit, and so I have made the assumption that 

A. siamense has the A. tenuipes type of inflorescence. 

Because of the doubt surrounding each fruiting specimen, I cannot be certain what actually are the ranges 

of A. simplex and A. temipes. I therefore hesitate to 

make further taxonomic reductions. 

13. Aglaonema simplex 

Aglaonema simplex Blume, 1837, p. 152, pls. 36D, 65. 

Caladium simplex Reinwardt ex Blume, 1823, p. 103 [nomen 

Aglaonema schottianum Miquel, 1856 (Aug. 15), p. 565; 

A. malaccense Schott, 1859, p. 30. 

A. longicuspidatum Schott, 1860, p. 304. 

A. propinquum Schott, 1864, p. 280. 

A. birmanicum Hooker f., 1893, p. 529. 

A. nicobaricum Hooker f., 1893, p. 530. 

A. angustifolium N. E. Brown, 1895, p. 18. 

A. angustifolium var. undulatum Ridley, 1907, p. 21.-FurA. borneense Engler, 1915, p. 22. 

A. schottianum var. genuinum Engler, 1915, p. 20. 

A. schottianum var. genuinum f. angustifolium (N. E. Brown) 

A. schottianum var. brownii Engler, 1915, p. 21. 

A. schottianum var. malaccense (Schott) Engler, 1915, p. 21. 

A. schottianum var. winkleri Engler, 1915, p. 21, fig. 7. 

A. breuivaginatum van Alderwerelt van Rosenburgh, 1922, 

A. elongatum van Alderwerelt van Rosenburgh, 1922, p. 326. 

A. emarginatum van Alderwerelt van Rosenburgh, 1922, p. 

A. grande van Alderwerelt van Rosenburgh, 1922, p. 325. 

A. latiusvan Alderwerelt van Rosenburgh, 1922, p. 325. 

A. nieuwenhuisii Engler ex van Alderwerelt van Rosenburgh, 

nudum]. 

1856 (Dec. 25), p. 216. 

tado, 1937, p. 243. 

Engler, 1915, p. 20. 

p. 324. 

324. 

1922, p. 320. 
40 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

A. simplex f. inaequale van Alderwerelt van Rosenburgh, 

A. simplex f. typicum van Aldenverelt van Rosenburgh, 1922, 

A. subarborescens van Aldenverelt van Rosenburgh, 1922, p. 

In the following description some exceptional measurements have been included in brackets which were 

given by van Aldenverelt van Rosenburgh in his description of A. subarborescens. 

Stem erect, 15-120 [300] cm tall, 0.4-1.7 (2.5) cm 

thick, thickest near the base. Internodes usually less 

than stem diameter, 0.3-2.5 cm long, longest near base 

of stem. Petioles (4.3) 5-12 (21.5) [27.5] cm long, 

(0.2) 0.3-0.5 (0.9) times as long as the leaf-blade. 

Sheaths usually with a membranous but sometimes 

slightly scarious margin, ( 1) 3-10 (22) cm long, (0.25) 

0.5-0.8 (1) times as long as the petiole. Leaf-blades 

narrowly oblong, narrowly elliptic to lancmlate, occasionally linear, elliptic or ovate, (10) 13-25 (35) 

cm long, (1.9) 4-10 (25) cm wide, length/width 

ratio 1 : ( 1.5) 2.8-5.0( 10) ; base often unequal or 

oblique, obtuse, .rounded or subtruncate, rarely acute or 

subcordate; apex often apiculate, acuminate, sometimes abruptly or gradually acuminate; variegation 

none; venation strongly differentiated into (3) 5-10 

( 14) primary lateral veins diverging from the midrib at 

(15”) 30”-50” (60°), rarely weakly differentiated; 

texture coriaceous. Peduncles ( 1) 2-4 (6), ( 1.5) 4-9 

(12) [17.5] cm long. Spathe often apiculate, (2.3) 3-5 

(6.5) cm long. decurrent (0.3) 0.4-1.0 (1.5) cm long. 

Stipe (0.2) 0.4-0.8 (1.2) cm long. Spadix equaling or 

slightly exceeding spathe, (1.7) 2-4 (4.3) cm long; 

pistillate portion 0.3-1.0 cm long, pistils (12) 18-30 

(38) ; staminate portion (1.5) 2.0-3.1 (3.8) cm long, 

0.5 cm thick. Fruits turning red, 1-1.7 cm long, 0.5- 

0.8 cm wide. 

TYPE COLLECTIONS.-)aVa, voet Pulusarie, [Kuhl 

and uan Hasseltl 12 (lectotype: L) ; Java, Gedogang, 

nahe van Kapang dungar, Kuhl and uan Hasselt s.n. 

(syntype: L) . Out of all the specimens in various herbaria, principally Leiden, that might be considered as 

type material, these are the only specimens with original labels that definitely associate them with Blume’s 

citations and discussions. I have selected the specimen 

from Mt. Pulusarie as lectotype since it shows a longstipitate spadix. Shortness of the stipe was used by 

Miquel to segregate A. schottianum from A. simplex. 

DIsTRIBuTI0N.-Southern Burma to the western 

Moluccas (Figure 2) . 

1922, p. 323. 

p. 323. 

326. 

FIGURE 14.-Lectotype of Aglaonema simplex Blume : KuhE 

and van Hasselt 12 (L) . 

HABITAT.-h primary or secondary forests in ravines 

or damp places. It rarely occurs over 1500 m but has 

been reported at “7000 feet” (over 2000 m) on Mt. 

Kinabalu (Clemens 40985). 

FLOWERING TIME.-Essentially nonseasonal. 

LOCAL NAMEs.-Only one name has been recorded 

more than twice, “Lempoei” on three different plants 

cultivated at Bogor and its probable variant, “Lempoeng.” Although the language involved is Sundanese, 

“lempung” or “lempong” means “light or soft wood” in 

Indonesian and Malayan and may be more of an adjective than a name. 

The existence of a specimen of Aglaonema simplex 

in the Linnaean herbarium, identified by Linnaeus 
NUMBER 1 

(presumably) as Arum ovatum, opened the question 

of whether it typified the Linnaean binomial. Linnaeus 

(1753, p. 967) defined Arum ovatum, “Arum acaule, 

foliis ovato-oblongis,” and cited “Karinpola. Rheed. 

mal. 11. p. 45. t. 23. Habitat in India.” Rheede’s (1692) 

plate of “Karinpola” is clearly Lagenandra ovata 

(L.) Thwaites. The specimen of Aglaonema in the 

Linnaean herbarium is devoid of any information except for being noted as “ovatum.” In view of this it 

seems clear that Linnaeus placed primary, if not total, 

emphasis on the information in Rheede. This contention is supported by the fact that the specimen of 

Aglaonema, although broken off near the leaves, is 

clearly caulescent and thus does not fit the critical 

phrase name of Linnaeus. Therefore, the Linnaean 

specimen of Aglaonema, referred to Arum ovatum by 

Linnaeus, should not be considered to typify the Linnaean binomial. 

Aglaonema simplex Blume is the most variable species in the genus. On the fringes of its distribution it 

tends to intergrade, to some extent, with other species. 

These complexities have led taxonomists to segregate 

many taxa within the complex and thereby obscure 

the relatively distinctive characteristics of Aglaonema 

simplex, sensu lato: strong venation, short petiole 

(rarely over 12 cm, i.e., one-half as long as the blade), 

short peduncles (rarely over 9 cm), usually more 

than 20 pistils, stipitate spadix, and spadix equaling 

the spathe. 

Within the species it is tempting to segregate the 

specimens into three groups: (1) narrow-leaved, (2) 

broad-leaved, and (3) intermediate. The narrowleaved plants might be defined as having the leaf-blade 

7-11 times longer than broad and 1.3-4 cm wide. 

Roughly speaking, the following taxa would fit this 

category: A. angustifolium, A. angustifolium var. undulatum, A. schottianum var. brownii, and their 

isonyms. The broad-leaved plants might be described 

as having the blade 1.4-2.5 times longer than broad 

and 8-25 cm wide and would certainly include A. 

nicobaricum, A. borneense, A. emarginatum, A. 

grande, A. latius, and probably also A. nieuwenhuisii 

and A. subarborescens. The intermediate plants have 

the leaf-blade 2.6-6.9 times longer than broad and (4) 

5-9 (11) cm wide and include A. simplex, A. schottianum, A. propinquum, A. malaccense, and probably 

A. schottianum var. winkleri, A. elongatum, and A. 

brevivaginatum (the latter three might be placed 

among the narrow-leaved plants if the definitions were 

41 

changed slightly). Simultaneous comparison of the 

types of these taxa with the specimens cited herein indicates that these taxa are parts of a spectrum, not 

district entities. I have tried grouping the specimens 

along the lines suggested above and also by geographical areas, and have found no evidence of a correlation 

of geographical areas with the morphological groups. 

Therefore, I have not recognized any of these taxa, 

even at the form level. 

Among the previously proposed taxa, two, though 

intergrading into the more usual forms through specimens from the same or other areas, do represent extremes. One is A. nieuwenhuisii from central Kalimantan, which represents A. simplex in its smallest 

aspect (leaf-blades only 12 cm long). The other is A. 

subarborescens from central Sumatra, which represents A. simplex in its largest aspect (leaves up to 35 

cm long and 24 cm across). Other collections from 

the same areas indicate that these specimens represent 

only extreme individuals. 

There are two areas where A. simplex appears to 

hybridize or intergrade with other species : Burma and 

Thailand, where it reaches the northwestern limit of its 

range, and the Philippines and Celebes, where it 

reaches the eastern limit of its range. 

In Burma and western Thailand the occurrence of 

subsessile spadices (for example, on the single stem of 

Kostermans 359 there is both flowering material with a 

clear stipe and fruiting material that shows no evidence of a stipe although the stipe is normally longest 

in fruiting material) and the vegetative variability of 

A. simplex make it difficult to be certain about determinations. It seems quite possible that the northernmost species with sessile spadices, A. hookerianum, A. 

ovatum, and A. modestum, as a group, are most closely 

related to A. simplex and/or A. tenuipes. 

In the Philippines and Celebes it is quite possible 

that A. simplex, along with A. nitidum and A. marantifolium, contributed or contributes to the variability found there. I have no other explanation for 

why the distinctive characteristic-combinations defining these three species, while remaining valid elsewhere, seem to break down in the Philippines and 

Celebes. Unlike the other two species, A. simplex does 

occur in the Philippines (A. nitidum does reach 

Balabac Island, but it is only 60 miles from the Bornean 

coast). 

The type of A. schottianum Miquel is: Java, 

Blambangan, Horsfield s.n. (holotype: K; isotype: 
SMITHSONIAN CONTRIBUTIONS TO BOTANY 

BM). The holotype, annotated by Miquel, does not 

bear the original field labels but only information 

written in what I presume to be Miquel’s hand. The 

isotype bears only field labels in Horsfield’s hand. 

There is another Horsfield specimen at Kew that may 

be type material; however, Horsfield labelled it as 

“Arum” while he labelled the isotype as ‘CPothos.” 

In the original description of Aglaonema schottianum, Miquel stated that it was close to A. simplex. 

Miquel’s (1856) descriptions of A. simplex and A. 

schottianum are not mutually exclusive. The only real 

distinction that Miquel seemed to make was that the 

stipe was short “circiter 1% lin.” in A. schottianum 

but was “longuisculus” in A. simplex. Miquel had a 

keen eye to segregate a new species on the basis of a 

stipe 0.32 cm long instead of 0.4 cm or more as described and illustrated by Blume for A. simplex. 

A review of the history of A. schottianum is necessary since its diagnostic characters changed as different 

authors sought other characters with which to separate this species from A. simplex. Schott (1860) says 

that he had not only seen dried material but also 

living material although he did not have colored plates 

prepared of the living material as he usually did. 

Schott first added a description of the spathe (note: 

Miquel’s types are in fruit and lack spathe material), 

apparently basing it on Spanoghe material, which, although it is not cited by Schott, is in Schott’s herbarium as a tracing annotated as A. schottianum. 

Schott noted that the spadix was about 1.5 inches 

( “subsesquipollicaris” ) long and slightly shorter than 

the spathe. At the same time, he noted that the spadix 

of A. simplex was finally almost longer than the spathe 

(“tandem fere spatha longior”) . At this point it must 

be noted that the spathelspadix ratio in A. simplex is 

virtually 1 : 1. In bud the spadix is shorter than the 

spathe, or in flower the spadix may equal or occasionally exceed the spathe but never by a significant margin. Indeed, I suspect that distortions introduced by 

pressing may affect whether or not the spadix is 

longer than or slightly shorter than the spathe when 

the original material actually indicated that the 

spadix equalled the spathe. 

Engler ( 1879), apparently wishing to follow Schott 

in maintaining A. schottianum, introduced some 

errors. The most serious error was reversing the one 

character that Miquel had used in segregating A. 

schottianum from A. simplex. According to Engler, A. 

simplex has a short stipe (“breviter stipitati”) while A. 

schottianum has a long stipe (“longe stipitati”) . Another error introduced by Engler was that he described 

the spadix as 1.5 cm long. Apparently he meant 4 cm 

but forgot to change inches to centimeters in translating from Schott’s “subsesquipllicaris.” Curiously, Engler (1879) included two of the same specimens under 

both A. simplex and A. schottianum, including a syntype of A. simplex. The syntype of A. simplex is cited 

under A. simplex as “Kapandangan” and under A. 

schottian’um as “Kapangdungor.” The other specimen 

is Zollinger 695B, which was correctly cited under A. 

schottianum but incompletely cited under A. simplex 

as “Zollinger 695.” Up to that time, no one had found 

anything significantly different between A. schottianum and A. simplex. 

In a later publication, Engler (1915) maintained 

his earlier findings but introduced a series of new 

collections. New Malayan specimens were cited under 

A. schottianum although the type of this species was 

from the farthest eastern tip of Java and new Javanese 

specimens were cited under A. simplex. Ridley ( 1925), 

following these identifications, did not question 

whether or not A. simplex occurred in Malaya but 

simply accepted the name A. schottianum. Gagnepain 

(1942) included both species in Indochina, placing 

them together as not having the spadix surpassing the 

spathe and differing only in their leaf bases. Bakhuizen 

van den Brink (1957) was the first to identify A. 

schottianum with A. simplex in his treatment of Araceae of Java. I entirely concur with this disposition. 

The type of A. malaccense is: [Malaya, Malacca] 

Pulo Bissar [Pulau Besar], Grifith, E. India Co. 5985 

(lectotype: K) . Although Schott originally (1859) 

cited only “Malacca, Griffith,” without citing a number or otherwise clearly designating a holotype, he did 

a year later (1860, p. 302) add the notation ‘‘u. s. in 

Herb. Hook.” There are two Griffith specimens from 

Malacca at Kew, one of which fits Schott’s description 

better than the other. This I have selected as lectotype 

because Schott described the leaf-base as obtuse to very 

obtuse; the leaf-bases on Grifith, E. India Co. 5984 

(K) are subcordate and highly unequal. Also, Schott 

(1860) described the leaf-blade as 2.5-3 inches wide; 

Grifith, E. India Co. 5984 has leaves fully 4 inches 

wide. 

A specimen at Kew (McClelland s.n., Burma, Phangrun Valley, Pegu ad Rangoon, 10 February) has been 

annotated by N. E. Brown as follows: “This is probably 

the type [of A. malaccense] rather than Griffith‘s speci- 
NUMBER 1 

mens, as the spathe and berries are undescribed.” Since 

Schott, in his original ( 1859) and a subsequent (1860) 

description, cited only Griffith’s material from Malacca, however, I have typified his A. malaccense on the 

Griffith material. The Schott descriptions support this 

typification since Schott (1859) described the leaf 

apices as acuminate to cuspidate, whereas they are 

merely tapered in McClelland’s material, and in 1860 

he described the petiole as 3.5-4 inches long, whereas 

one of the most conspicuous petioles on the McCelland 

specimen is fully 5.5 inches long. 

The type of Aglaonema longicuspidatum is: Indonesia, Borneo, Bangarmassing, 1857-8, Motley 876 (hob 

type: K) . Study of Schott’s descriptions indicates that 

the most distinctive features of the type are the longcuspidate apex and the undulate margin of some of 

the leaves. Actually, the leaf apices are gradually longacuminate, something which is frequently found, especially in the narrower-leaved A. simplex. The undulate 

margin appears to be nothing but an artifact of pressing, particularly since only two of the four leaves on 

the type show this characteristic. Engler (1879, p. 440) 

reduced A. longicuspidatum to synonymy of A. simplex. 

The type of Aglaonema propinquum is: Indonesia, 

Borneo, Korthals s.n. (lectotype : L-898, 87-85). 

There are three duplicates in the Leiden herbarium, 

all annotated by Schott as A. propinquum. The 

lectotype was designed on the basis that it is 

represented by a tracing in the Schottian herbarium 

(Schott, Aroideae 79). Schott (1864) noted that this 

material was close to A. malaccense and A. longicuspidatum, both of which are synonyms of A. simplex. 

Engler ( 1879, p. 440) made A. propinquum a synonym 

of A. schottianum, which in turn I consider synonymous with A. simplex. 

Aglaonema fallax Schott ex Engler (1879, p. 439) 

was published only as a synonym of A. simplex. Color 

drawings in Schott’s herbarium (Schott, Aroideae 43 

and 44), presumably prepared from cultivated material, indicate that its placement here is entirely 

correct. 

The type of Aglaonema birmanicum is: Birma [sic], 

toward Nempean, low woods, Grifith, E. India Co. 

5981 (holotype: K). This locality is unknown to me 

although Hooker (1893) cites it as from “Upper 

Burma.” The type and associated collections from 

Burma and western Thailand frequently have very 

short to obscure stipq. This occasionally happens 

throughout the range of A. simplex (Scheepmaker s.n. 

43 

from Java has a stipe 0.2-0.5 cm long; Rahmat si 

Toroes 4609 from Sumatra has stipes 0.3-0.6 cm long; 

several specimens from Borneo and Celebes have stipes 

0.3 cm long). The fact, however, that all the Burmese 

specimens have short stipes (except Helfer 5992 from 

Tenasserim and McClelland s.n. from Pegu, both of 

which have stipes 0.4 cm long) suggests that A. 

birmanicum is not a distinct species but some sort of 

integradation between A. simplex and a species 

with sessile spadices, possibly A. ovatum. The short 

peduncles preclude its being identified with A. 

hookerianum. 

The type of Aglaonema nicobaricum is: India, 

Nicobars, 23 November 1884, King’s Collector 537 

(lectotype : K; isolectotype: BM, GAL) . King’s Collector 537 is selected as lectotype since the description 

of the spathe was obviously made from it and another 

collection for the Nicobars, Kurz s.n. (February 1875), 

lacks the spathe. In fruiting material the spathe is decurrent for 0.5 cm and the stipe is 0.7 cm long. Other 

specimens with leaves 1.5-2.5 times longer than broad 

are scattered throughout the range of A. simplex: 

Burma-Meebold 15467; Thailand-Kostermans 359; 

Sumatra-Henderson SFN 20122, Biinnemeijer 4311, 

Meijer 5645; Borneo-Brooke 10700, Nicolson 1306; 

Celebes-Elbert 3169. I have been unable to find any 

other distinguishing morphological characters that 

correlate with the broad leaves, and it is clearly impossible to draw an arbitrary line based solely on leaf 

width because the specimens, in toto, simply form a 

spectrum of leaf widths from 1.5-11 times longer 

than broad. Therefore, I am not even accepting any 

subspecific taxa based solely on leaf width. If later 

workers wish to do so, A. nicobaricum is the oldest specific epithet applicable to the broad-leaved specimens of 

the Aglaonema simplex complex. 

The type of Aglaonema angustifolium N. E. Brcwn 

is: Great Britain, Kew Gardens, “Aroid from ?amore. 

Received from Mr. Curtis in 1893,” 23 November 

1894 [N. E. Brown?] (lectotype: K). The lectotype 

has some of the narrowest leaves in Aglaonema simplex, ten times longer than broad ; however, the distinctiveness of this character begins to fade when one 

finds that the leaves of the syntype are only seven times 

longer than broad. Later collections from the typelocality (Burkill 186 and 187, from South Pangkore 

Forest Reserve) show even more variation in adjacent 

plants. Burkill 187 can be indentified with the lectotype as the leaves are over eight times longer than 
44 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

at Bogor, Koorders 161 79, was not mentioned and (by 

my own measurements) has petioles only 9-13 cm long, 

it is apparent that he was not discussing or describing 

the taxon, but only the specimens that he was excluding. Association with Koorders 161 79 holds this 

name to the species in which it used by Engler and 

discussed by van Aldenverelt van Rosenburgh. 

Although Ridley accepted the taxon Aglaonema 

angustifolium N. E. Brown in his works on the flora of 

the Malay peninsula (1907, 1925), he felt it necessary 

to distinguish the typical element described by Brown 

as A. angustifolium var. undulatum. His reasoning was 

that “the type-plant described by Brown was the very 

local form var. undulatum. The leaves of young plants 

of typical Tie., common] A. angustifolium are almost 

black.” This action resulted in his application of the 

name A. angustifolium to a taxon considered different 

from that represented by the type on which it was based 

and in the renaming of the typical element. This is, of 

course, inadmissible. Aglaonema angustifolium var. 

undulatum Ridley is a synonym both nomenclaturally 

and taxonomically of A. angustifolium N. E. Brown. 

The undulate margins occur in A. simplex in other 

localities. 

The holotype of Aglaonema borneense is: Hort. 

Buitenzorg, January-February 1906, Engler, Reise 

nach Java und Brit. Zndien 4089 (B). According to 

Engler (1915), this material came from Borneo. The 

specimen has two detached leaves, one 7 cm wide and 

one 12 cm wide. The smaller leaf appears to be very 

thin and probably is immature. Since the larger leaf 

agrees very closely in all respects with Nicolson 1306 

and Brooke 10700 from Sarawak, there is no reason to 

doubt the Bornean origin. The only deterrent in recognizing this as a broad-leaved specimen of A. simplex is 

that the spadix appears to be fully 1.5 cm short of the 

spathe apex. The spadix, however, has been sliced 

longitudinally and, since the cut was diagonal, it appears that a substantial portion was taken from the 

apex of the spadix. Engler indicated that his taxon was 

most closely related to A. siamense [A. tenuipes herein], 

but this is not supported by the spathe length in the two 

taxa (less than 2.5 cm in A. tenuipes, 5 cm in A. borneense). The species appears to be no more than another broad-leaved A. simplex, similar to that which 

was earlier described as A. nicobaricum. 

Aglaonema schottianum var. genuinum f. angustifolium (N. E. Brown) Engler is a change in rank of 

A. angustifolium N. E. Brown. Engler cites only the 

broad. Burkill 186 has leaves only 4.3-4.6 longer than 

broad and is noted: “Seems to be an age form of 

specimen HMB 187. Both plants were growing adjacent.” It is probable that Burkill, aware that he was 

in a variable population, collected extremes to demonstrate the range of variability. I strongly suspect he 

could also have collected intermediates. Intermediates 

certainly are known as Engler (1915, p. 20) pointed 

out when he reduced Aglaonema angustifolium to the 

rank of a forma. Curtis 3820 (SING) from Wellesley 

has n2rrow leaves at the base up to 9.5 times longer 

than broad and upper leaves only 4.1 times longer 

than broad, clear evidence that the width of the leaves 

can be variable on the same plant. Equally narrowleaved specimens have been collected in Borneo: (1) 

Amjah 90, (2) material in cultivation from Borneo 

named A. elongatum Aldenverelt, (3) material called 

A. schottianum var. winkleri Engler. There is no reason to believe that intergradations do not also occur 

here. Despite the striking aspect of individual narrowleaved specimens, we seem to be dealing with something that is transient or a spectrum, not with a distinct 

taxon. Therefore, I am following Bakhuizen van den 

Brink (1957, p. 35) in completely synonymizing A. 

angustifolium with A. simplex. 

Aglaonema simplex f. macrophyllum (Engler, 1898, 

p. 21 ) , although a nomen nudum, can be associated 

with a preserved specimen (Koorders 16179 [BO] 

from Tatokok, Celebes) and thus assigned to synonymy 

with A. simplex. When Engler first used the name 

(without description), he cited five specimens, Koorders 16126, 16129, 16155, and 16179 from Minhassa, 

Celebes, and Warburg I4984 from Jolo, Philippines. 

The last-mentioned specimen, presumably formerly at 

Berlin, is no longer extant. Alderwerelt van Rosenburgh ( 1922, p. 323) undertook a discussion of some of 

the specimens of A. simplex f. macrophyllum at Bogor 

and stated that the petioles were 17.5-25 cm long. This 

at first appears to be a validating description for 

Engler’s name, but van Alderwerelt van Rosenburgh 

went on to indicate that the material from which these 

measurements were taken actually belonged to another 

species, A. oblongifolium [A. philippinense herein]. He 

cited only Koorders 16126, 16129, and 16155 and 

stated that at Bogor there was no material of Koorders 

16129. The other two specimens agreed “in [his] 

opinion more with A. oblongifolium than with A. 

simplex.” Since the one other specimen cited by Engler 

that was available to van Alderwerelt van Rosenburgh 
NUMBER I 

syntype, Scortechini 5.71. There is no significant morphological difference between the types of A. 

schottianum var. genuinum f. angustifolium and A. 

schottianum var. brownii Engler (their types are even 

almost topotypes) . Furtado ( 1937, p. 243) discussed 

this involved nomenclatural problem. 

The type of Aglaonema schottianum var. brownii is: 

Perak, Dindings, Lumut, April 1898, Ridley 9508 (lectotype: SING; isolectotypes: CAL, K) ; Perak, Dindings, Lumut, February 1899, Ridley 10144 (syntypes: 

CAL, SING). Engler actually cited Prain as a parenthetical author (Engler’s types are determined [in 

Ridley’s hand] as tlglaonema brownii Prain) . Prain 

was interested in Araceae and published several new 

species of Typhonium but never published anything on 

Aglaonema. Examination of the lectotype and syntype 

collections of A. schottianum var. brownii reveal them 

to be scarcely distinguishable from the type of N. E. 

Brown’s A. angustifolium. The leaves are light green, 

extremely narrow, and the margins are crispulate. 

Aglaonema schottianum var. malaccense is a change 

in rank of A. malaccense Schott. Engler does not cite 

Schott’s types but his description of the leaves as being 

only three times longer than broad fits the types. 

The type of Aglaonema schottianum var. winkleri is: 

Sudost-Borneo, zwischen M. Uja and Kundimbaru, 

July 1908, Winkler 2727 (TBG or HBG) . Unfortunately, I have not seen the type, but there is no reason 

to think that it does not exist. I have interpreted it on 

the basisof the illustration published by Engler ( 1915). 

As described and illustrated by Engler, this specimen 

with narrowly oblong leaves looks very much like A. 

propinquum Schott, which Engler places as a synonym 

of A. schottianum. The leaves, however, are slightly 

narrower, 4.5-5 cm wide instead of 5-7 cm wide. I see 

no reason to maintain this variety, especially when it 

seems possible that narrow leaves can be transient (see 

discussion under A. angustifolium) . 

The type of Aglaonema brevivaginatum is: Lingga 

Archipel, P. Lingga, Kp. Panggak, 60 m, 27 July 1919, 

Bunnemeijer 7044 (holotype: BO-100314; isotypes: 

BO-100315, L-922,297-1858). This material appears 

to be no more than a rather small Aglaonema simplex. 

As the specific epithet implies, the patiolar sheath is 

very short, “5-1 (raro 1v2-2) cm longa.” On one 

isotype (L) I found a sheath 2.5 cm long, almost half 

the petiole length. Although short sheaths are not common in A. simplex, the sheath length is exceedingly 

variable, even on the same specimen, and it is im326-551 -94 

45 

possible to segregate taxa on this characteristic alone. 

The following specimens also have sheaths less than 

2 cm long: Jelinek 32 from Nicobar and Nicolson 11 15 

from Singapore. 

The holotype of Aglaonema elongatum is: Cult. in 

Hort. Bog. as IX. 61 (Borneo) [March 1921 - fide description], van Alderwerelt van Rosenburgh 355 (BO) . 

In the original description it was reported that the 

plant had been collected on Mt. Tenampak in eastern 

Borneo by Amjah, a collector on Capt. van Genderen 

Stort’s North Borneo Boundary Commission. The living plant is no longer at Bogor. The description reported the leaves five to seven times longer than broad and 

that the plant was nearest to Aglaonema schottianum, 

but unfortunately, nothing was mentioned about what 

made it different from A. schottianum. The leaves seem 

to be dark green, narrow, smooth margined, and 

gradually long-acuminate or tapered like the material 

that Ridley (1907, 1925) regarded as the most 

“typical” [i.e., common] form of Aglaonema angustifolium. As pointed out in the discussion of that 

synonym, however, the narrowness of leaves does not 

furnish enough evidence to warrant recognition of 

taxa. 

The holotype of Aglaonema emarginatum is: Eiland 

Sebesi, 100 m, 29 April 1921, Docters van Leeuwen 

5412 (BO). The author gives no clue as to what 

species this specimen might be related nor how it is 

differentiated from other species. The leaves are 

“16-23 x 8-11 cm.,” and thus clearly identical in size 

with those of Aglaonema nicobaricum. 

The holotype of Aglaonema grande is: Cult. in Hort. 

Bot. Bog. XI. B. IX. 62, [March 1921 - fide description] 

van Alderwerelt van Rosenburgh 361 (BO) . According 

to the description, the living plant was collected in 

Borneo as Nieuwenhuis 863. I have not seen a specimen with this number collected by Nieuwenhuis or 

by any of the collectors who accompanied him: 

Amdjah, Hallier, Jaheri, Molengraff, or Sekarang. The 

author gives no clue, except the specific epithet, as to 

what distinguishes this taxon or its relatives. The type 

appears to be nothing but another broad-leaved 

Aglaonema simplex, this time with leaves “25- 

35 x 11.5-15 cm.” It is larger, to be sure, than the 

large-leaved plants described as Aglaonema nicobaricum and A. emarginatum but that is hardly sufficient 

reason to maintain it, even at a subspecific level. 

The type of Aglaonema latius is: Eil. Simaloer. 

29 March 1919, Achmad 1009 (lectotype: BO-100331; 
46 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

isolectotypes: BO-100332, L-992, 297-539) ; Eil. 

Simaloer, 12 June 1918, Achmad 487 (syntypes: BO, 

L, U). I have designated Achmad 1009 as lectotype 

since it has the widest leaves. Judging by the epithet, 

the author was emphasizing leaf width; in his description he states that the leaves are 10.5 cm wide. The 

leaves on Achmad 487 are less than 10 cm wide but 

they are occasionally 10.5 cm wide on Achmad 1009. 

The author gives no clue as to related species. It seems 

clear, however, that the specimens represent another 

broad-leaved Aglaonema simplex, this time a bit 

smaller than the material described as Aglaonema 

nicobaricum, A. emarginatum, and A. grande but 

hardly warranting recognition as a distinct taxon. Even 

the author suggested that leaf width may not be very 

significant when he described the leaves as sometimes 

three times longer than broad. 

The holotype of Aglaonema nieuwenhuisii is: 

Borneo, Soengei Bloe-oe [tributatry of Mahakan River;], 

1896-7, Jaheri, Exp. Nieuwenhuis 673 (BO) . Engler 

(1915) cited this collection as Aglaonema minus 

Hooker f., having longer petioles, the petiolar sheath 

more than 0.5 the petiole length, and a longer leaf 

blade. This is true. The only question is whether it 

should be recognized as a taxon within A. simplex. The 

main characteristic is that the leaves are very small, 

“9-12 x 4%-6 cm.” Recently the Bailey Hortorium received photographs and a cutting of a similar plant 

collected in southeastern North Borneo by L. Maurice 

Mason. Considering the variability known to exist in 

Aglaonema simplex, I have decided not to recognize 

what amounts to a group defined merely as “‘plants 

small.” 

The holotype of Aglaonema simplex f. inaequalz is: 

Lingga Archipel, P. Singkep, Weg Manggoe, 50 m, 

Bosch, 2 August 1919, Biinnemeijer 7189 (BO) . In the 

original publication this collection was erroneously 

cited as Biinnemeijer 7180. I have no record of Biinnemeijer 7180 but Biinnemeijer 7189 is annotated by van 

Aldenverelt van Rosenburgh as Aglaonema simplex 

f. inaeguale and bears the field note “Bloeikolf wit. 

Bloeischeede licht groen,” which must be the basis 

of the part of the description reading “Spatha pallide 

viridis. Spadix albus.” This taxon is not worthy of recognition. Leaves of most Aglaonema are unequal to a 

greater or lesser degree. 

The type of Aglaonema subarborescens is: Sum- 

[atra], W[ester] K[ust], G. Malintang, 1100 m, Bosch 

bij rivier, 5 August 1918, Biinnemeijer 4311 (lectotype: BO-100506; isolectotypes: BO-100507-100508, 

L) ; Sum[atra] W[ester] K[ust], G. Sago, 1150 m, Bosch 

rand, 7 August 1918, Biinnemeijer 4351 (syntype: 

BO) . I have designated Biinnemeijer 431 I (BO100506) as lectotype since it has the largest leaves, 32.5 

cm long and fully 24 cm wide. This exceptionally large 

size is approached in the genus only by Aglaonema 

densinervium, which differs in having the spadix over 

1.5 cm short of the spathe apex. In all characters this 

material resembles Aglaonema simplex. Other collections from the same general locality, Biinnemeijer 3631 

(which was determined by van Alderwerelt van Rosenburgh as A. simplex), Biinnemeijer 431 I, and Meijer 

5645 clearly indicate that the lectotype is, at best, only 

an extreme individual. It is desirable to be aware of 

these extremes, but I feel that formal recognition of 

them, even at an infraspecific level, will lead to confusion in such a variable species as Aglaonema simplex. 

Aglaonema alpinum Elmer (1938, p. 3611) is based 

on : Philippines, Sorsogon, Irosin, Mt. Bulusan [300C 

ft, wet humus-covered stony soil of densely wooded 

flats - fide description], July 1916, Elmer 16520 (BO, 

CAL, GH, NY) . Of the four duplicates of the type-collection known to me, only one (GH) has leaves 5.5 

cm wide as described by Elmer. This binomial is invalid under Article 36 (ICBN, 1966) since it was published without a Latin description. The material falls 

nicely into Aglaonema simplex with its short peduncles, 

short petioles, and nonvariegated leaves. Unfortunately, most of the Philippine collections of what I regard to be Aglaonema simplex are in fruit, and it is 

difficult to be certain whether or not the spadix equals 

the spathe as it should in A. simplex. The only flowering material is Zwickey 349 (GH) from Lanao. In two 

rather immature inflorescences the spathe seems to 

exceed the spadix by more than one centimeter but 

a third, rather passk, inflorescence appears to have the 

spadix equaling the spathe. 

14. Aglaonem cordifolium 

Aglaonema cordifolium Engler, 1915, p. 29, figs. 14A-C. 

Stem erect, to 1.2 cm thick. Internodes to 1 cm long. 

Petioles 7-9 cm long, 0.4 times as long as the leafblades. Sheaths membranous, 1 cm long, 0.1-0.3 times 

as long as the petiole. Leaf-blade broadly ovate, 10-21 

cm long, 7-16 cm wide, length/width ratio 1 : 1.3-1.5; 

base cordate; apex acuminate with apiculum; variegation none; venation differentiated into 5-14 primary 

lateral veins diverging from midrib at 45’-60”. Peduncles 2-3 together, 5-7 cm long. Spathe 5 cm long, 
NUMBER 1 47 

Herre 1179 from Negros Oriental, determined as A. 

philippinense, is very similar to A. cordifolium, at least 

in the appearance of the leaf-blades. 

15. Aglaonema marantifolium 

Aglaonema marantifolium Blume, 1837 [“1835”], p. 153, 

Arum aquaticum Rumphius, 1747, p. 312, PI. 108. 

Adpendix erecta Rumphius, 1747, p. 487, pl. 182: fig. 2. 

Calla oblongifolia Roxburgh, 1814, p. 65. 

Aglaonema oblongifolium (Roxburgh) Kunth, 1841, p. 55 

Scindafisus erectus Presl, 1851 [L‘1849”], p. 242. 

Aglaonema novoguineense Engler, 1898, p. 2. 

Stem erect or reclining in larger plants, 1-3 cm thick. 

Internodes to 2 cm long. Petioles (18) 21-31 cm long, 

(0.6) 0.8-1.0 (1.1) times as long as the leaf-blade. 

Sheaths with membranous margins, (1 1) 15-20 (25) 

cm long, (0.4 0.6-0.8 times as long as the petiole. Leafblades narrowly oblong, occasionally oblong, narrowly 

elliptic or oblanceolate, (22) 25-33 (40) cm long, 

(7.5) 10-15 (18) cm wide, length/width ratio 1 : (2.2) 

2.5-3.4; base obtuse to subrounded; apex acuminate, 

sometimes subrounded, obtuse or abruptly acuminate, 

often apiculate ; variegation none; venation undifferentiated to weakly differentiated into 5-8 primary 

lateral veins diverging from the midrib at (30”) 40”- 

50’ (65’) ; texture coriaceous. Peduncles 2-5 toget!ier, 

rarely solitary, (10) 13-19 (21 ) cm long, commonly 

about 0.5 the length of the petioles. Spathe green, 

turning yellow and withering in age, 4-7 (to 9 in fresh 

material) cm long, decurrent for 0.3-0.8 (1.0) cm. 

Stipe (0.7) 0.9-1.6 cm long. Spadix shorter than 

spathe by 0.3 the length of the spathe or more, 1.7-3.0 

(to 4 in fresh material) cm long; pistillate portion 

0.2-0.8 cm long, pistils 10-17 (22), the pistil with 

broad yellow stigma, the style distinctly contracted ; 

staminate portion 1.2-2.0 (to 3 in fresh material) cm 

long. Fruits becoming bright red, 1.5-3.0 cm long, 

0.7-1.7 cm wide. Seed clavate, to 2.5 cm long and 1.2 

cm thick in fresh material. 

TYPE coLLEcTIoNS.-Amboina, zippel s.n. (lectotype: L) ; Banda, Peitsch s.n. (syntype: L) . Blume 

(1837) noted that Zippel’s specimen was mutilated (a 

piece of a leaf and a small piece of stem with six inflorescences attached) . Peitsch‘s specimen is more 

complete but is sterile, and therefore I have selected 

Zippel’s specimen as lectotype. 

DISTRIBUTION.-MO~UCCS through New Guinea 

(Figure 2). 

pl. 66 [L‘marantaefolia”]. 

[not Schott, 18291. 

FIGURE 15.-Specimen of Aglaonema cordifolium Engler: 

Ramos and Edano, BS 36626 (US). 

2 cm wide, decurrent to 0.8 cm. Stipe 1-1.5 cm long. 

Spadix 2.5 cm long; pistillate portion 0.3-0.8 cm long, 

pistils ca. 20; staminate portion 2 cm long, 0.4 cm 

thick. 

TYPE COLLECTION.-Philippines, Mindanao, Davao, 

Mt. Dagapan, Ebenenwald, June 1888, Warburg 

14604 (holotype: B) . 

DIsTRIsuTIoN.-Known from only two collections 

in Davao and Zamboanga of southern Mindanao (Figure 8). 

This species appears to be most closely related to 

Aglaonema simplex with which it shares the following 

characteristics : petioles less than one-half the blade 

length, petioles less than 10 cm long, and petiolar 

sheaths less than 10 cm long. Perhaps future studies 

will indicate that A. cordifolium is not a distinct 

species, but the specimens seem to be distinctive by the 

broadly ovate leaf with a cordate leaf base and an 

exceedingly short petiolar sheath, characters that make 

it advisable to maintain the species. 
48 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

Calla oblongifolia is based on Arum aquaticum 

Rumphius. Most binomials in Roxburgh’s Hortus 

Bengalensis are nomina nuda but a footnote to Calla 

oblongifolia refers to the description and place of 

Rumphius. It is interesting to note that Roxburgh 

(1814) reports that his plant was introduced from the 

Moluccas by C[hristopher] Smith in 1798. Even Roxburgh‘s drawing, published by Wight (1844), is 

Aglaonema marantifolium and not A. nitidum as m’ost 

workers have assumed. 

The Roxburghian epithet was transferred to Aglaonema by Kunth ( 1841), thus creating a later homonym 

of the superfluous name Aglaonema oblongifolium 

Schott (1829) (see A. nitidum for details). 

Scindapsus erectus is based on Adpendix erecta 

Rumphius. 

Aglaonema marantifolium f. robustum Engler ( 1883, 

p. 291) is based on Beccari, P.P. 798 from Andai, site 

of a small village formerly about 20 kilometers south 

of Manokwari in West New Guinea. The name, however, was published without a description. Study of the 

original and other collections from the same general 

locality do not warrant validation of the forms, which 

Engler (1915) later ignored. 

The type of Aglaonema novoguineense is: Territory 

of New Guinea, Suor Mana [vicinity of Astrolabe 

Bay?], 700 m, 12 June 1896, Lauterbach 3.25 (lectotype: B) . In the original description six syntypes were 

listed, all at Berlin. Only one remains and I have designated it as lectotype. The segregation of this species 

is probably largely due to Engler’s overemphasis on 

the relative and subjective characteristic of strength 

of venation. He (1915) placed A. novoguineense in 

his classification as having subequal, thin, dense, subparallel primary venation and A. marantifolium as having primary veins slightly stronger (“paullum validiores”) than the secondary veins. This distinction does 

not exist and even the type of A. nouoguineense would 

key out to A. marantifolium either in Engler’s or my 

own key. 

Aglaonema marantifolium Blume is most closely 

related to A. philippinense, A. denrineruium, and A. 

commutatum. Study of herbarium material suggests 

that A. philippinense may be intermediate between A. 

marantifolium and A. simplex, that A. densineruium 

may be an overgrown intermediate between A. 

marantifolium and A. nitidum, and that A. commutatum may be a subapomictic complex of variegated 

forms in which A. marantifolium has participated 

FIGURE l6.-Aglaonema marantifolium Blume (1837, pl. 66). 

HABITAT.-shady, damp places in lowland rain forFLOWERING TIME.-Apparently nonseasonal. 

Both Arum aquaticum and Adpendix erecta are 

typified by the pertinent Rumphian plates. Rumphius 

described Arum aquaticum as being decumbent with 

half the stem lying on the ground and half erect. He 

placed Adpendix erecta among the climbing aroids but 

noted that it does not fasten itself to trees but lazily 

lies on neighboring bushes. There is no difference between them as Merrill (1917, p. 217) recognized when 

he stated: “The figures and descriptions of both the 

Rumphian species cited above agree closely with 

Aglaonema oblongifolium Kunth [=A. marantifolium] 

as currently interpreted.” 

ests. 
NUMBER 1 49 

along with other species. In the Moluccas and New 

Guinea all the specimens consistently have the spathe 

much longer than the rather aborted spadix, a shortly 

decurrent spathe, and a distinctly elongated stripe, 

usually much exceeding the length of the spathe decurrency. The specimens, however, from the Talaud Islands have rather short stripes, but they are longer 

enough than the spathe decurrency so that they can 

be included in A. marantifolium in its narrowest sense. 

Field work in the Philippines, Celebes, and Moluccas, 

particularly if combined with cytological studies and 

breeding experiments, would be extremely desirable 

to determine whether or not, or to what extent, A. 

marantifolium is capable of interbreeding with other 

taxa. 

16. Aglaonema commutatum 

Aglaonema commutatum Schott, 1856, p. 123. 

Stems erect, becoming decumbent in older and larger specimens, 20-150 cm tall, 0.5-6.0 cm thick. Internodes 0.4-2.5 cm long. Petioles 6-25 (35) cm long, 

(0.4) 0.5-1.2 (1.4) times as long as the leaf-blade. 

Petiolar sheaths membranous but occasionally scarious. 

Leaf-blades usually narrowly oblong-elliptic to lanceolate but occasionally ovate, elliptic or oblong, (10) 

13-30 (45) cm long, (2.5) 4-10 (20) cm wide, length/ 

width ratio 1 : (1.8) 2.3-3.7(6), base often unequal, 

broadly acute, obtuse or subrounded, occasionally subcordate; apex often apiculate, acuminate to gradually 

acuminate, rarely shortly acuminate or broadly acute; 

variegation confluent in bars along the primary lateral 

veins, rarely in irregularly scattered spots (var. commutatum) ; venation differentiated into 4-7 ( 11) primary lateral veins diverging from the midrib at (20") 

45"-70" (80") ; texture coriaceous. Peduncles solitary 

to 6 together, (4.5) 7-15 (20) cm long. Spathe light 

green, sometimes white, (3) 3.5-9.0 ( 12) cm long, (2) 

2.8-4.0 (4.8) cm wide, decurrent for (0.2) 0.4-1.2 

(2.0) cm. Stipe (0.2) 0.4-1.1 (2) cm long, rarely surpassing the spathe decurrency. Spadix thin-cylindric 

( 1.6) 2-6 (7) cm long, usually more than 1 cm short 

of spathe apex but occasionally equaling it; pistillate 

portion 0.3-1.0 cm long, pistils (7) 10-18 (22) ; staminate portion (1.1) 1.5-4.0 (6) cm long, 0.4-0.6 cm 

thick. Fruits turning yellow, then bright red, ellipsoidal 

to obovoid, 1.6-2.5 cm long, 0.4-1.7 cm thick. 

DISTRIBUTION (Figure 5) .-Philippines and northeastern Celebes. The ability of this species to naturalize 

makes it difficult to be certain which collections are 

from the wild and which are escapes. 

FLOWERING TIME.-June-September, the wet season 

in areas with pronounced seasons. 

This species is quite variable but, with the exception 

of the typical variety, is always variegated in the same 

barred pattern. In some of the cultivars extreme variegation and albinism occur but the basic pattern is still 

evident. This consistent variegation pattern, coupled 

with its relatively restricted distribution, has led me to 

accept the complex as a single variable species. 

The only hypothesis I can offer to explain the great 

variability in vegetative and reproductive characteristics is that it may have a hybrid origin. One of the 

most difficult areas in which to recognize and define 

species of Aglaonema is the area of the Philippines and 

Celebes. If all specimens from this general area are 

excluded from consideration, one has little difficulty in 

recognizing Aglaonema marantifolium (Moluccas and 

New Guinea), A. nitidum (most of western Malesia) , 

and A. simplex (most of western Malesia). The moment one tries to include and account for the Philippine 

and Celebesian specimens, these otherwise distinct 

species begin to show extremely confusing exceptions 

and intergrades. 

Aglaonema commutatum is one of the worst offenders in this regard. For instance, Aglaonema marantifolium has a long stipe and short spathe decurrency. 

These also occasionally occur in A. commutatum. 

Aglaonema nitidum has scarious petiolar sheaths. 

These also occasionally are found in A. commutatum. 

Aglaonema simpIex typically has very short peduncles, 

but these also may occur in A. commutatum. Each of 

these characteristics is virtually definitive for the three 

principal species. 

Why should the peripheral area of the Philippines 

and Celebes provide the exceptions? It seems reasonable to postulate a hybrid origin for the plants in this 

area, in particular for A. commutatum. This hypothesis is supported by what little cytological evidence is 

available. All chromosome counts of Aglaonema commutatum have been tetraploid or hexaploid and the 

plants were pollen-sterile and apomictic (see "Cytology," p. 6). 

A number of specific taxa have been proposed that 

appear to fall into this complex. After study of the 

specimens, I found it possible to accept all of these taxa 

by expanding their original concepts. This seemed 

preferable to completely submerging them or to 

proposing a whole series of new infraspecific taxa. 

It was surprising to me that some of the taxa are 
50 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

apparently restricted to central Luzon and northeast- studies, particularly if combined with breeding experiern Celebes with nothing in between. This is only one ments and cytological work, will clarify why simple 

of the many unsolved problems. Perhaps further field morphology does not solve all the taxonomic problems. 

Key to the Varieties and Cultivars of Aglaonema commutatum 

1. Petioles solid dark green. 

2. Variegation in irregularly scattered spots ............. 16a. var. commutatum 

2. Variegation in blot hy bars following the primary lateral veins. 

3. Petioles equalinc tn exceeding leaf-blades ........... 16b. var. warbi rgii 

3. Petioles shorter than leaf- lades 

4. Leaves large (more than 25 x 10 cm) with length/width ratio of 1 :1.8-2.5 ....... 

............................ 16c. var. robt,stum 

4. Leaves smaller, rarely over 25 cm long, less than 10 cm wide; length/width ratio 1 :2.’.-6.0 

5. Leaves less than 5 cm wide; length/width ratio 1 :4.5-6.0 .... 161. cv. ‘Tnubii’ 

5. Leaves usually over 5 cm wide; length/width ratio 1:2.4-4.3. 

6. Leaves over 20 cm long, base obtuse to acute ....... 16d. var. eleguns 

6. Leaves less than 20 cm long, base rounded to subcordate . . 16e. var. mucdutum 

1. Petioles variegated. 

7. Leaf variegation streaked ................... 16m. cv. ‘Tricolor’ 

9. Leaves over 50% variegated above ........... 16i. cv. ‘Parrot Jungle’ 

......................... 16h. cv. ‘Malay Beauty’ 

7. Leaf variegation in blotchy bars following the primary lateral veins. 

8. Petioles more green than white. 

9. Leaves less than 50% variegated. 

10. Sheath 0.5 times as long as petiole; petiole 0.9 times a long as blade ....... 

10. Sheath 0.9 times as long as petiole; petiole 0.5 times as long as blade . 16j. cv. ‘Pewter’ 

8. Petioles more white than green. 

11. Leaves tricolored above .... 

1 1. Leaves bicolored above. 

12. Leaves variegated below ... 12. Leaves solid green below ... 

16a. Aglaonema commutatum var. commutatum 

Aglaonema commutatum Schott, 1856, p. 123. 

A. marantifolium p commutatum (Schott) Engler, 1879, 

The following is a translation of Schott’s (1860, p. 

304) most complete description of Aglaonema commutatum: 

Petioles sheathing almost all the way to the apex, 6.5-8.0 

cm long. Leaf-blade oblong-lanceolate, base obtuse, margin 

weakly undulate, apex long-acuminate, on the upper side 

irregularly sprinkled with remote white-greenish spots. Veins 

spreading or erect-spreading, k5 with 3-4 secondary veins 

between. Peduncles longish, thin. Spathe large, lanceolateoblong, long-acuminate, whitish green outside, whitish inside. 

Spadix thin, slightly longer than spathe, the stipe longish, 

the pistils few, more or less in a single series. Stigma pale 

yellowish green, deeply excavated. Fruit elliptic, red. Seed 

elliptic. 

TYPE.-Unpublished drawing, Schott, Aroideae 38 

(neotype : W) . 

This variety is unknown to me except for several 

drawings in the Schottian herbarium in Vienna and 

photographs appearing in the various editions of Graf‘s 

p. 441 [sensu stricto]. 

.. ........ 16k. cv. ‘Pseudobracteatum’ 

.......... 16g. cv. ‘Fransher’ 

...... 16f. cv. ‘Albo-variegatum’ 

Exotica (1957, p. 71 ; 1959, p. 80; 1963, p. 118) as 

Aglaonema cuscuaria. 

The typification of this well-known species was different than expected and deserves comments. As typified here and described by Schott (1856, 1860), the 

leaves have irregular and scattered spots, not the usual 

pattern of blotchy bars following the primary lateral 

veins. An unpublished plate found among Schott’s reliquiae fits the description. A published plate by 

Schott (1858, pl. 59) does not show leaves that, in this 

case, are definitive. Therefore, I selected the unpublished plate (Aroideae Nr. 38) as neotype. 

16b. Aglaonema commutatum var. warburgii 

(Engler) Nicolson, new status 

Aglaonema warburgii Engler, 1915, p. 20. 

Petioles green, 13-24 (27) cm long, 1.0-1.4 times 

as long as the leaf-blade. Petiolar sheaths 7-14 cm long. 

Leaf-blades 10.5-24 cm long, 4-8.5 cm wide, length/ 

width ratio 1 : 2.2-2.8 (3.1 ) ; variegation barred, along 

primary lateral veins; venation differentiated into P-7 

primary lateral veins diverging from the midrib at 
NUMBER 1 51 

FIGURE 17.-Neotype of Aglaonema commutatum var. commutaturn: Schott, Aroideae Nr. 38 (W). 

(45") 50"-70". Spathe 3-4.5 cm long. Spadix 1.6-2.5 

cm long, shorter than spathe by 0.8 cm (in the type) 

to equaling it; staminate portion 1.1 cm long, 0.4 cm 

thick. 

TYPE coLLEcTIoN.-North Celebes, Menado [August-September 18881 Warburg s.n. (holotype : B) . 

The type material shows the usual barred variegation pattern of Aglaonema commutatum. It is distinct 

by its long petioles, which equal or exceed the length 

of the leaf-blade. A long petiole is also usual in A. 

marantifolium and a relationship to that species is also 

indicated by the fact that the stipe often equals the 

spathe decurrency. The material seems to be morphologically distinct enough to warrant maintenance at the 

varietal level for the-time being. 

16c. Aglaonema commutatum var. robustum 

Aglaonema commutatum var. robustum (van Alderwerelt 

van Rosenburgh) Nicolson, 1968, p. 125. 

A. robustum van Alderwerelt van Rosenburgh, 1922, p. 328. 

Petioles 15-35 cm long, 0.5-0.8 times as long as the 

leaf-blade. Petiolar sheaths 6-26 cm long. Leaf-blades 

21-45 cm long, 11-20 cm wide, length/width ratio 

1 : 1.8-2.5; variegation sometimes faint, along primary 

lateral veins; venation differentiated into 5-1 1 primary 

lateral veins diverging from the midrib at 65"-80". 

Spathe 7-12 cm long, 3.54.8 cm wide. Spadix 4.5-6.5 

cm long, tapered, equaling to 2 cm short of spathe 

apex; staminate portion 3.0-5.5 cm long, 0.4 cm thick. 

TYPE COLLECTION.-Java, Bogor Botanic Garden, 

XI. B. IX. 152, van Alderwerelt van Rosenburgh 358 

(holotype: BO) . Nicolson 955 is clonotype material. 

This variety was originally described as a species 

near Aglaonema commutatum. The variegation pattern is clearly the same. In habit the plants look like A. 

marantifolium, but the spathe decurrency is longer 

than the stipe, indicating that this material is not true 

A. marantifolium. 

Bunting 1545 represents a smaller form. The leaves 

are only 2 1 cm long; however, the presence of extremely 

broad leaves ( 1 : 1.8-2.4) fits this variety. 

16d. Aglaonema commutatum var. elegans 

Aglaonema commutatum var. elegans (Engler) Nicolson, 

A. elegans Engler, 1915, p. 28, fig. 11. 

Petioles 9-20 cm long, 0.44.7 times as long as the 

leaf-blade (equaling, fide Engler) . Petiolar sheaths 

(4) 9-13 cm long. Leaf-blades 17-30 cm long, 5-9.5 

cm wide, length/width ratio 1 : (2.7) 3-3.7; variegation in the form of bars following the main veins on the 

upper surface; venation differentiated into 4-7 primary lateral veins diverging from the midrib at (30") 

45"-70" (80", fide Engler). Spathe (3.5) 4.5-7.0 

(10.5) cm long, 2-3.9 cm wide. Spadix 2-7 cm long; 

staminate portion 1.'5-5.7 cm long, 0.4-0.6 cm thick. 

TYPE COLLECTION.-Java, Bogor Botanic Garden, 

Engler, Reise nach Java und Britisch Indien 19054 n. 

4070 (not found in B or BO). In the absence of a 

specimen, Engler's published illustration may be considered as type-material. It should be noted that Engler 

described the veins as diverging from the midrib at 70"- 

80" while the illustration shows the veins diverging 

at 45" or less. 

DIsTRIsuTIoN.-PhilippineS, central Luzon. Escapes 

from cultivation. 

Engler indicated that his taxon was close to A. 

commutatum but was distinguished by its larger and 

more elongate leaves and especially by its elongate1968, p. 125. 
52 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

oblong spathe. The illustration of the spathe is not 

particularly distinctive nor are the measurements 

given in the description: “6-7 cm. longa 1 cm. ampla” 

(Engler apparently measured width without unrolling 

the spathe and, thus, his “1 cm. wide” would correspond to 2-3 cm wide in a flattened spathe). The 

distinctive character of the leaves can be summarized 

as a length of over 20 cm and more than 2.9 times the 

width. 

The best approximation to Engler’s description is 

made by McGregor, Bur. Sci. 23077, which even has the 

veins diverging at 70” from the midrib. 

16e. Aglaonerna commutatum var. maculatum 

Aglaonema commutatum var. maculatum (Hooker f.) NicolA. marantifolium var. maculatum Hooker f., 1865, pl. 5500. 

A. commutatum var. commutatum of authors not Schott: 

Regel, 1865, p. 130, pl. 470.-Morren and deVos, 1882, 

p. 51.-Engler, 1915, p. 27, figs. lOG-P.-Birdsey, 1951, 

A. marantifolium sensu Boynton, 1924, p. 165, pl. 302 [not 

Petioles 6-13 (18) cm long, 0.5-0.9 times as long 

as the leaf-blade. Petiolar sheaths 3-1 1 cm long. Leafblades 10-20 cm long, 3.5-7.0 cm wide, length/width 

ratio 1 : 2.4-3.4( 3.7) ; variegation confluent in stripes 

following the primary veins on the upper leaf surface; 

venatibn differentiated into 4-7 primary veins diverging from the midrib at (35”) 450-60° (70”). Spathe 

3.5-6.5 cm long, 2.8-3.0 cm wide. Spadix 2-4.7 cm 

long, equaling to 1 cm short of spathe apex; staminate 

portion 1.5-4.0 cm long, 0.4-0.6 cm thick. 

TYPE coLLEcTIoN.-England, Chelsea, Royal Exotic 

Nursery, 25 March 1863 [without coIlector] (holotype: K). Loher 2445 is mounted on the same sheet 

with the holotype. 

son, 1968, p. 125. 

pp. 12-13. 

Blume, 18371. 

DISTRIBUTION.-Central LUZOn. 

According to the holotype, this variety was imported 

from Manila by Veitch. According to Morren and de 

VOS ( 1882, p. 51), Linden exhibited it in Brussels 

on 19 July 1863. Most authors have used the binomial 

A. commutatum to refer to var. mtaculatum, probably 

because the typical variety (var. commutatum) is quite 

rare and not nearly so well known and widely distributed as var. maculatum. 

This variety is apparently most closely related to 

var. elegans. As happens in that variety, occasional 

cultivated specimens show inflorescence characters of 

A. marantifolium, such as having the stipe more or less 

equaling the spathe decurrency. The similar length of 

FIGURE 18.-Holotype of Aglaonema commutatum var. maculatum (Hooker f.) Nicolson: without collector (K). (Holotype upper half onIy.) 

spathe decurrency and stripe suggests that A. marantifolium may be related, even though the overall sizes 

of the plants are widely different. 

The wild usually differs from cultivated material in 

the petiolar sheath length. In cultivated material the 

sheath is (0.6) 0.7-0.9 ( 1 ) times as long as the petiole 

length, but in the cultivated material it is only (0.3) 

0.5-0.6 (0.8) times as long. 

The Cultivars of Aglaonema commutatum 

The cultivar names known to me are discussed below 

in alphabetic order. 

Mr. Nat De Leon of the Parrot Jungle in Miami, 

Florida, has done a great deal of breeding and selecting in Aglaonema commutatum. He is responsible for 

the origin of most of the commercial cultivars. Un- 
NUMBER 1 

fortunately, time was not taken to study his material 

and to obtain information directly from him. I have 

relied on the information published by Graf (1963, 

p. 1526-1527). The hybrid origin noted for several of 

the cultivars may simply be the result of apomixis rather 

than hybridization since apomixis, pollen sterility, and 

high polyploidy are well established in this species. 

16f. Aglaonema commutatum cv. ‘Albo-variegatum’ 

Aglaonema commutatum albo-variegatum Graf, 1953, illustr. 

5: fig. 24 [“albo-uariegata”]; 1957, p. 70; 1959, p. 76; 

1963, p. 119. 

Graf described this cultivar as a sport. In the published photographs it looks exactly like A. commutatum 

var. maculatum except that the petioles are white, not 

green. 

16g. Aglaonema commutatum cv. ‘Fransher’ 

Aglaonema ‘Fransher’ Graf, 1963, p. 1526. 

This cultivar was noted by Graf as a hybrid, A. treubii X marantifolium tricolor [A. commutatum cv. 

‘Treubii’ X commutatum cv. ‘Tricolor’], and was 

named after Francis Scherr, founder of the Parrot 

Jungle in Miami, Florida. It is distinguished by variegated petioles, leaves heavily bicolored above, and 

speckled below along the midrib and lower main veins. 

16h. Aglaonema commutatum cv. ‘Malay Beauty’ 

Aglaonema ‘Malay Beauty’ Graf, 1963, pp. 117, 1527. 

Noted by Graf as a free-growing horticultural variation of A. commutatum ‘Pseudobracteatum’ and perhaps a return toward normal from the extreme albino 

form. Graf also notes that this cultivar is called ‘Pewter.’ 

In the material available to me there are two specimens that would fit Graf’s general description. Bunting 

1346 (Bamboo Nursery, Orlando, Florida, 3 April 

1962) has petiole 12 cm long (0.5 times as long as the 

leaf-blade) and the petiolar sheath 10.5 cm long (about 

0.9 times as long as the petiole). These measurements 

are similar to those of ‘Pseudobracteatum.’ I have 

identified this collection as ‘Malay Beauty.’ The other 

specimen with a much longer petiole was identified as 

‘Pewter.’ 

16. Aglaonema commutatum cv. ‘Parrot Jungle’ 

Aglaonema ‘Parrot Jungle’ Graf, 1963, pp. 117, 1527. 

This cultivar is distinguished by having most of the 

leaf variegated above and none below. It was noted 

by Graf as a hybrid, A. curtisii X pictum tricolor [A. 

nitidum f. curtisii X picturn cv. ‘Tricolor’]. The dis53 

tinctness of the putative parents suggests that they may 

have been misidentified. 

16j. Aglaonema commutatum cv. ‘Pewter’ 

Aglaonema ‘Pewter’ Graf, 1963, p. 1527 [under ‘Malay 

As discussed under ‘Malay Beauty,’ I have identified 

that cultivar with a specimen closely approximating 

A. commutatum cv. ‘Pseudobracteatum’ in its petiolar 

measurements. Another specimen, Bunting 1372 ( Ferrell’s Shack, Winter Park, Florida, 4 April 1962), that 

I identify with ‘Pewter,’ has petioles 35 cm long (longer 

than the leaf-blade) and the petiolar sheath 18 cm 

long (about half the petiole length). 

16k. Aglaonema commutatum cv. ‘Pseudobracteatum’ 

Aglaonema pseudobracteatum Graf, 1953, p. 5, fig. 35. 

A. ‘Pseudo-bracteatum’ Graf, 1957, p. 49; 1959, p. 39; 1963, 

This cultivar is distinguished by its massive variegation that eliminates most of the chlorophyll from both 

sides of the leaf and the petiole. The epithet is possibly 

a misapplication of A. robelinii lusus [sport] pseudobracteosum Engler (1915, p. 19, fig. 6). Engler applied 

this descriptive epithet to a plant with bractlike structures subtending the pistils, something not observed in 

living material nor in any of the descriptions or specimens of the cultivar under discussion. 

161. Aglaonema commutatum cv. ‘Treubii’ 

Aglaonema commutatum cv. ‘Treubii‘ Nicolson, 1968, p. 125. 

A. treubii sensu Graf, 1953, p. 6, fig. 38; 1957, p. 69; 1959, 

This cultivar is distinguished by its narrow leaves and 

long spathe (equaling the peduncle). Comparable 

spathe length is also found in A. commutatum cv. 

‘Pseudobracteatum’ and var. robustum. This cultivar 

approaches A. commutatum var. elegans in its leaf 

shape and its long spathe. Perhaps it is only a selection 

or sport of that variety. 

In the trade “treubii” is consistently used as a specific epithet for this cultivar, but it apparently represents a misidentification of A. treubii Engler (1898) as 

amended by Engler (1915). Engler’s (1915) description clearly does not agree with the cultivated material 

in the following points : he described the variegation 

as being along the midrib (“juxta costam”) and the 

leaf-blade as 7-12 cm wide (length/width ratio of 

1 : 2.5-2.9). It is true that there is confusion about the 

identification of A. treubii Enger (see “Dubious BiBeauty’]. 

pp. 37, 114. 

p. 79; 1963, p. 117 [not Engler, 18981. 
54 

nomials,” p. 60, no. 5) but it seems clear that the 

material in the trade is not truly A. treubii Engler. 

I have seen two collections of this taxon from the 

Philippines that may have been from the wild but that 

I suspect were from cultivation. Since the taxon is only 

kn30wn from cultivation, it seems best to treat it as a 

cultivar. 

16m. Aglaonema cornmutaturn cv. ‘Tricolor’ 

Aglaonema cornmutaturn cv. ‘Tricolor’ Nicolson, 1968, p. 125. 

A. marantifolium tricolor Graf, 1953, p. 5, fig. 30. 

There are two specimens from Mindoro, Philippines 

(Sulit and Conklin, PNH 16836, 17656) that apparently belong to this taxon; rather than validate the 

taxon as a botanical variety, however, I prefer to treat 

it as a cultivar. This cultivar is distinguished by its 

hazy to streaked leaf variegation affecting both sides of 

the leaf and its mottled to pink petioles. 

It should be noted that a usual character of A. 

marantifolium appears in the wild material of this 

cultivar; i.e., the stipe exceeds the spathe decurrency 

by 0.3 cm. The presence of this characteristic on Mindoro plants is exceptional as it normally is restricted 

to the Moluccas and New Guinea. This may be an 

indication that A. marantifolium had a part in the 

evolution of A. cornmutaturn. 

17. Aglaonema crispum 

Aglaonema crisfium (Pitcher and Manda) Nicolson, 1968, 

Schismatoglottis crispa Pitcher and Manda, 1892, p. 95 [as 

Aglaonema roebelinii Pitcher and Manda, 1894, p. 31. 

Schismatoglottis robelinii (Pitcher and Manda) Pitcher and 

Aglaonema robelinii lusus pseudobracteosum Engler, 1915, 

Stem erect, 40-120 cm tall, 0.8-3.0 [5, fide Alderwerelt] cm thick. Internodes about 1 cm long. Petiole 

8-25 cm long, 0.4-0.7 times as long as the leaf-blade. 

Sheaths (4) 6-12 (18) cm long, (0.4) 0.6-1.0 times 

as long as the petiole. Leaf-blades elliptic to narrowly 

elliptic 14-32 cm long, (5) 7-12 (16) cm wide, 

length/width ratio 1 : 2-2.9 (3.5) ; base obtuse, 

subrounded or broadly acute; apex acuminate; 

variegation covering most of the upper surface 

of each half of the leaf-blade, leaving the 

midrib and marginal areas free; venation rather weakly 

differentiated into (4) 6-8 (10) primary lateral veins 

diverging from the midrib at 40”-60”; texture 

coriaceous. Peduncles 2-5 together, 4-14 cm long. 

p. 126. 

“Schizmatoglottis”]. 

Manda, 1895, p. 141, pl. 138. 

p, 19. 

SMITHSONIAN CONTRIBUTIONS TO BOTANY 

Spathe fading from green to yellow, then black, 4-7 

cm long, 2.G2.5 cm wide, decurrent for 0.4-1.0 (1.5) 

cm. Stipe 0.3-0.7 (1) cm long. Spadix 2-3 cm long, 

0.7-2.0 cm shorter than spathe; pistillate portion 0.2- 

0.4 cm long (to 1 cm in fruit), pistils 10-17; staminate portion 1.6-2.6 cm long, 0.2-0.3 cm thick. Fruits 

becoming yellow, then red, 1.5-2.0 cm long, 0.9-1.5 

cm thick. 

NEoTYPE.-PhotOgraph labelled Schisrnatoglottis 

roeblinii on page 141 of Pitcher and Manda, General 

Illustrated Catalogue of Plants (1895). This same 

photograph also appears as Schismatoglottis crispata 

(not Hooker) in Vick’s Magazine (1899, vol. 23, p. 

71) and as Schisrnatoglottis picta (not Schott) in 

American Gardener (1898, vol. 19, fig. 122). 

DIsTRIBuTIoN.-Known only from Mt. Bulusan, 

Sorsogon Province, Luzon, Philippines, where it has 

been collected several times (Figure 5). 

HABITAT.-In deep shade of primary dipterocarp 

forest from 300 to 950 m. 

FLOWERING nME.-August. 

According to the label on a specimen of this species 

at Kew (without collector) it was given to Kew by 

Mr. Sander in 1881. In 1883 Linden’s firm began 

offering A. (or Schismatoglottis) roebelinii under a 

variety of spellings. I presume, as did Wocke (1895) 

and Engler ( 1915), that these nomina nuda pertain to 

this species. In 1891 Pitcher and Manda listed Schismatoglottis crispata, possibly the same as S. crispa 

Pitcher and Manda, although it is impossible to be 

certain since no description was given. 

FIGURE 19.-Neotype of Aglaonema crispum (Pitcher and 

Manda) Nicolson. 
NUMBER 1 

Schismatoglottis crisp. Pitcher and Manda was described thus: ‘Somewhat resembles Dieffenbachia in 

growth and has fine grayish green leaves, edged with 

dark green, the midrib being a lighter color.” This 

description can be followed through subsequent catalogs of Pitcher and Manda’s firm until it can be 

connected with the photograph published by Pitcher 

and Manda (1895) and called Schismatoglottis roebelinii, which I have designated as the neotype. 

Unfortunately, I have not been able to see any of 

Sander’s catalogs published between 1881 (when 

Sander material was made into a specimen at Kew) 

and 1892 when the hitherto obscure epithet “crkpa” 

was published with a description. Sander’s firm (still 

extant and run by a grandson) and Kew do not have 

catalogs. Until valid publication of A. roebelinii that 

antedates "crisps" can be demonstrated, I am obliged 

to adopt the oldest available epithet for this species. 

This is unfortunate because “roebelinii” is well known 

in the trade. 

At least nine different spellings have been published 

for the hitherto most commonly used epithet: roebelinii, rebellinii, robellinii, roblenii, rebelini, robelenii, 

robelini, robeleni, robellini. This epithet is believed to 

honor the probable collector, Carl Roebelen. According to van Steenis-Kruseman ( 1950, p. 444), Roebelen 

was collecting in the Philippines for Sander’s company between 1880 and 1882. Bellaire and St. LCger 

( 1900, p. 165), however, reported that “A. de Robelin” 

was being honored. I have no information on this 

person. According to Article 73 (ICBN, 1966), the 

original spelling of a name must be retained, except 

for correction of errors. Since there is some confusion 

about the correct spelling, it seems best to retain the 

original spelling. 

Elmer (1938, p. 3614) misapplied the binomial A. 

philippinense. His description is based on two specimens, Elmer 14993 and 15706, identified herein as A. 

crispum and A. simplex, respectively. The description 

is based on Elmer 14993. 

The relationship of A. crispum to other species is 

obscure. The long, scarious petiolar sheath is similar to 

that of A. nitidum. What is known of the cytology of 

cultivated material indicates that it is triploid (Pfitzer, 

1957). There is a possibility that apomixis contributes 

to the stability of the distinctive features of this taxon; 

however, it should be maintained as a species until 

further information is available. 

55 

18. Aglaonema stenophyllum 

Aglaonema stenophyllum Merrill, 1925, p. 452. 

Stem erect, 25-100 cm tall, 0.6-3 cm thick. Internodes 0.5-1 cm long. Petioles (6) 7-17 (22) cm long, 

(0.2) 0.4-0.5 (0.8) times as long as the leaf-blade. 

Sheaths (3.5) 6.5-17 (22) cm long, (0.5) 0.8-1.0 

times as long as the petiole. Leaf-blades sublinear to 

narrowly elliptic or lanceolate, (13) 20-35 (40) cm 

long, 2.9-7.5 (13) cm wide, length/width ratio 1 : (3.7) 

4-9 ; base narrowly acute, rarely obtuse ; apex gradually 

long acuminate; variegation usually none (but spots 

observable on lower leaf surface of Nicolson 789) ; 

venation more or less weakly differentiated into 4-12 

primary lateral veins diverging from the midrib at 

15”-30° (45) O. Peduncles 1-3 together, (5.5) 7.7-20 

cm long. Spathe 5.5-7.5 cm long, 3-3.5 cm wide, decurrent for (0.4) 0.7-1.5 cm. Stipe (0.4) 0.5-1.0 cm 

long. Spadix 4.2-5.0 cm long, 1-1.7 cm short of spathe 

FIGURE 20.-Lectotype of Aglaonema stenophyllum Merrill: 

Ramos, BS 30471 (US). 
56 

apex; pistillate portion 0.4 cm long, pistils 12-16 and 

slightly immersed, stigmas broad and styles contracted ; 

staminate portion 3.8-4.6 cm long, 0.5-0.6 cm thick. 

Fruits green, turning yellow and then red, 1.5-1.6 cm 

long, 0.6-1.1 cm thick. 

TYPE coLLEcTroNs.-Phi~ippines, Catanduanes, 

14 November-1 December 1917, Ramos, Bur. Sci. 

30471 (holotype; PHN [lost]; lectotype: US) : 

Samar, Catubig River, February 1910, Ramos, Bur. 

Sci. 24141 (paratype: US) ; Tayabas [Camarines 

Norte], east coast, Mt. Cadig, December 1916, Yates, 

Bur. Sci. 25413 (paratype: PHN [lost]; Biliran, June 

1914, McGregor, Bur. Sci. 18844 (paratype: PNH 

DISTRIBUTION.-Central Luzon to central MinHABITAT.-Damp places in primary forest to 900 m. 

This species is distinguished by its narrow leaves, 

long and scarious petiolar sheath, and long (usually 

over 10 cm) peduncles. Juvenile leaves of A. densinervium are rather small and elongate and have a 

surprising but superficial similarity to A. stenophyllum. Aglaonema stenophyllum is indistinguishable 

from A. philippinense so far as floral and inflorescence 

characters are concerned, but it differs in having a 

smaller petiole/blade-length ratio and a larger sheath/ 

petiole ratio. 

The general appearance of specimens of this species 

suggests a possible hybrid origin between A. nitidum 

(which has a long, scarious sheath) and narrow-leaved 

A. simplex. The short spadix (compared to the spathe) 

and the few pistils suggest that A. marantifolium may 

have been involved. 

19. Aglaonema densinervium 

Aglaonema densinervium Engler, 1906, p. 134. 

Stem erect, 90-200 cm tall, 1.2-6.0 cm thick. Internodes 1-2 cm long. Petioles (14) 20-35 (50) cm long, 

(0.6) 0.7-0.9 (1.2) times as long as the leaf-blade. 

Sheaths membranous, sometimes subscarious, (8.5) 

17-30 (45) cm long, (0.5) 0.7-1.0 times as long as 

the petiole. Leaf-blades elliptic, rarely narrowly elliptic, (22) 26-42 cm long, (9) 12-19 cm wide, length/ 

width ratio 1 : ( 1.8) 2-2.7; base rounded to obtuse, 

sometimes subtruncate or broadly acute ; apex abruptly 

acuminate ; variegation none ; venation undiff erentiated or weakly differentiated into (7) 10-12 primary 

lateral veins diverging from midrib at 45”-60°. 

Peduncles 1-7 together, (13) 20-32 cm long. Spathe 

6-11 cm long. 3-4 cm wide, decurrent for (0.5) 1-2 

[lost]). 

danao (Figure 8). 

SMITHSONIAN CONTRIBUTIONS TO BOTANY 

cm. Stipe (0.3) 0.5-1.2 (1.5) cm. Spadix 3-5.5 cm 

long, 1.5-3.5 short of spathe apex; pistillate portion 

0.3-1.0 cm, the pistils 6-15, with contracted style and 

broad stigma; staminate portion 2.7-5.0 cm long, 0.4- 

0.8 cm thick, tapered. Fruits turning red, 1-2 cm long, 

0.5-1.5 cm thick. 

TYPE COLLECTION.-Philippines, Mindanao, Davao, 

near sea level, March 1904, Copeland, Govt. Lab. 41% 

(holotype : B ; isotype : US) . DISTRIBUTTON.-Centra1 and southern Philippines 

and northeastern Celebes (Figure 5). 

HABITAT.-Near streams in disturbed or primary 

rain-forest up to 810 m. 

Specimens of this species have been referred to A. 

haenkei (Presl) Schott (see “Dubious Binomials,” p. 

59, no. 2). 

This species probably attains the greatest size of 

any in the genus, suggesting it may be related to the 

other two large species, A. nitidum and A. marantifolium. In many respects it combines characters of 

both: (1) the petiolar sheath may have a scarious or 

membranaceous margin, but it is nearly always membranaceous in A. marantifolium and scarious in A. 

nitidum; (2) the petiolar sheath is nearly equal to the 

petiole, as in A. nitidum, but the leaf-base usually is 

rounded to obtuse, as in A. marantifolium; (3) the 

spathe decurrency is rather long, like that in A. nitidurn, but the stipe is rather long, like that of A. 

marantifolium; (4) the staminate portion of the 

spadix is rather long, like that in A. nitidum, but the 

pistils are few, as are those in A. marantifolium; (5) 

its distribution is intermediate and peripheral to both 

species. 

Juvenile leaves of A. densinervium and unusually 

broad leaves of A, stenophyllum suggest a relationship 

that is unexpected when one compares the extremes 

of both. 

As one might suspect from the key, it may be difficult to separate large A. philippinense from smalI A. 

densinervium. Although these two species may be 

closely related, the characters given in the key will 

usually distinguish them. 

The Philippine material of Aglaonema is a rather 

confusing set of specimens that combine characteristics of A. marantifolium, A. nitidum, and A. simplex. 

To my mind, A. densinervium represents an overgrown 

intermediate between A. nitidum and A. marantifolium. It would be desirable to test this hypothesis 

with breeding and cytological studies. In the absence 

of such information, it is best to maintain it as a species. 
NUMBER 1 57 

20. Aglaonema philippinense 

Aglaonema philippinense Engler, 1898, p. 21. 

A. latifolium Engler, 1906, p. 134. 

Stem erect to decumbent in older plants, 50-75 cm 

tall, 1-3.2 cm thick. Internodes 0.4-2.5 cm long. 

Petioles (6.5) 14-25 (35) cm long, (0.5) 0.7-1.0 

(1.3) times as long as the leaf-blade length. Sheath 

membranous or scarious, rarely variegated (Nicolson 

829), (5) 8-17 (25) cm long, (0.3) 0.5-0.8 (0.9) 

times as long as the petiole. Leaf-blades elliptic to 

narrowly elliptic, rarely oblong to narrowly oblongelliptic, lanceolate or obovate, (11.5) 17-27 (39) cm 

long, (5) 8-13 (15) cm wide, length/width ratio 

1 : ( 1.6) 2.1-2.9 (3.8) ; base obtuse to rounded, rareIy 

subcordate or broadly acute ; apex often apiculate, 

acuminate, sometimes abruptly acuminate; variegation 

none ; venation undifferentiated to weakly differentiated into 5-9 (15) primary lateral veins diverging 

from midrib at 45"-60" (75"). Peduncles 1-4 (5) 

together, (7) 8-1 7 (25) cm long. Spathe often apiculate, 5-9 cm long, 2-3.4 cm wide, decurrent for 0.5-1.5 

cm. Stipe 0.3-0.8 (1.0) cm long. Spadix 1-2.5 cm short 

of spathe apex; 3-6 cm long; pistillate portion 0.2-0.4 

cm long, pistils 8-13, stigma broad, style contracted; 

staminate portion 2.8-5.2 cm long, 0.3-0.5 cm thick. 

Fruits green, turning yellow and finally red, 1.1-2.0 

(2.7) cm long, 0.6-0.9 ( 1.3) cm thick. 

TYPE COLLECTION.-Philippines, Tayabas [Quezon], 

Atimonan August 1904, Gregory, Govt. Lab. 86 

(neotype : B) . 

DIsTRIBUTI0N.-Philippines, Indonesia (Celebes) , 

and associated archipelagos (Figure 8). 

HABITAT.-Below 1000 m on slopes near streams in 

primary forests. 

FLOWERING TIME.-May-July, correlating with the 

onset of monsoon. 

Engler enumerated the following types in his original 

description of A. philippinense: Gaudichaud s.n. (B) 

from Manila; Warburg 12494 from Monte Alban 

[Montalban]; and Warburg 14604 from Mindanao. All 

of these were lost along with the Berlin herbarium except for Warburg 14604, which was later (1915) excluded by Engler and became the holotype of 

Aglaonema cordifolium Engler. Gaudichaud material 

from Manila does exist at Geneva, but it is annotated 

by Engler as A. marantifolium. Also, Engler (1915) 

cited Gaudichaud material from Manila in Paris as 

being A. marantifolium. 

FIGURE 2 1.-Neotype of Aglaonema philippinense Engler, 

holotype of Aglaonema latifolium Engler: Gregory, Govt. 

Lab. 86 (B). 

In Engler's 1915 treatment of A. philippinense, one 

additional specimen was cited, the one selected here as 

neotype. It must be noted that the neotype of A. 

philippinense Engler (1898) is also the holotype of A. 

latifolium, which Engler published .in 1906. Engler 

later, in 1915, indicated that he no longer regarded A. 

latifolium as distinct. In fact, he went so far as to redescribe A. philippinense (1915) with an almost exact 

copy of the original description of A. latifolium, in 

which the measurements are consistently larger than 

those appearing in the original description of A. 

philippinense. Since Engler and I are in agreement 

on the taxonomy, I have accepted the material introduced by Engler as neotype for A. philippinense 

although it is the holotype for A. latifolium. 

Syntype material of the nomen nudum A. simplex 

f. macrophyllum Engler (1898, p. 21) is placed here 
58 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

(Koorders 16126,16129, and 16155) ‘ The lectotype is 

A. simplex. 

Aglaonema philippinense Engler is similar to A. 

marantifolium in that it has relatively few pistils, the 

spathe distinctly exceeding the spadix, and petioles 

that are comparatively long compared to the leafblades; however, it lacks the combination of the short 

spathe decurrency and long stipe characteristic of A. 

marantifolium. The margin of the petiolar sheath is 

frequently scarious, suggesting A. nitidum, but many 

specimens have a membranous margin. The specimens 

as a whole do not seem to have any significant combinations of characters that suggest a strong relationship 

with either A. nitidum or A. simplex. 

It is surprising that the species is not found on 

Mindanao although it is found in all areas around 

Mindanao. This could be explained by a lack of collections from Mindanao or selection of wrong characters in the key resulting in an artifical grouping of 

specimens. I am inclined to suspect the latter hypothesis 

is true. This is part of the confusion involving all Philippine Aglaonema. 

21. Aglaonema pumilum 

Aglaonema pumilum Hooker f., 1893, p. 530. 

A. marmoratum Engler, 1915,~. 17. 

Stem decumbent ( ?) , often reddish, 0.3-0.5 cm thick. 

Internodes 0.3-1.3 cm long. Petioles 3.5-7.0 cm long, 

0.3-0.7 times as long as the leaf-blade. Sheaths 1-3.5 

cm long, 0.2-0.8 times as long as the petiole. Leafblades ovate to lanceolate, 8-13.2 cm long, 2.3-5.2 cm 

wide, length/width ratio 1 : 2-2.9; base often unequal, 

usually rounded, obtuse to truncate; apex gradually 

acuminate; variegation absent or ashy-blotched on the 

upper leaf surface; venation not or only weakly differentiated into 3-4 primary lateral veins diverging 

from midrib at 45” ; texture rather coriaceous. Cataphylls frequent, interspersed with leaves, like brown 

paper, 3-4 cm long. Peduncle solitary, 3-5 cm long, 

often equaling the petiole. Spathe 2-4.5 cm long, 1-1.5 

cm wide, decurrent for 0.4-1.0 cm. Stipe present( ?). 

Spadix thin-cylindric, 0.8-2.0 cm short of spathe apex, 

1.5-2.0 cm long; pistillate portion 0.2-0.4 cm long, 

pistils ca. 5; staminate portion 1-1.7 cm long, 0.2-0.3 

cm thick. Fruits 1-1.5 cm long, 0.5-0.8 cm thick. 

TYPE COLLECTION.-Burma Mergui, Grifith s.n. 

(lectotype: K) ; India, Assam, Lushai Hills, Chappedon, February 1827, Wallich, E. India Co. 8960B 

(syntype: not seen). There is sonz doubt about the 

identity of Wallich’s collectici since the locality, Chapa 

FIGURE 22.-Lectotype of Aglaonema pumilum Hooker f. : 

Grifith s.n. (K). 

Tong Hill, is about 700 miles away from what appears 

to be the range of A. pumilum. I was unable to find 

Wallich’s specimen at Kew, Calcutta, or any of the 

herbaria I visited. Since Griffith‘s specimen is in fruit, 

Wallich‘s specimen must be the basis of Hooker’s description of the spathe. 

DISTRIBUTION.-L~~~~ Burma and peninsular Thailand (Figure 6). 

HABITAT.-h evergreen forest. 

Among the Schottian drawings at Vienna, “Schott, 

Aroideae Nr. 68” is labelled “8900 Wallich Cat, Chappedong, Feb. 1827 Herb. Benth.” This label is incorrect. Study of Wallich, E. India Co. 89608 (not B) 

from Singapore clearly shows that this is the basis of 

“Schott, Aroideae Nr. 68.” This error is critical since 
NUMBER 1 

Wallich, E. India Co. 89604 is the holotype of A. minus 

(A. nebutosum f. nanum herein). 

The type of A. marmoratum is: Hort. Bot. Calcutta, 

18 April 1896, Prain 5.72. (holotype: CAL). Engler 

did not cite the collector’s name, but he did cite a 

specimen flowering in Calcutta in April 1896. This 

specimen meets those requirements and has been annotated by Engler as A. marmoratum. I can see no reason to maintain A. marmoratum, except perhaps as a 

variegated form of A. pumilum. In the original description of A. marmoratum, Engler (1915) commented: 

“Species imperfecte cognite et fortasse cum priore [A. 

pumilumj, cujus inflorescentia ignota, conjugenda.” 

My studies indicate that Engler’s hypothesis is correct. 

The relationships of this species are obscure. There is 

a possibility that it should be placed in the section 

Chamaecaulon in that it may be repent, there are 

cataphylls among the leaves, and it usually has very 

short petiolar sheaths; however, it may only be decumbent, the cataphylls do not clasp the petioles as 

they do in Chamaecaulon, and one specimen, Brown 

s.n., has sheaths 2-3.5 cm long. Gagnepain annotated 

the Thai collections as A. .costaturn var. virescens. The 

thin-cylindric spadix, however, much shorter than 

the spathe, is unlike that species. 

There is also a possibility that the spadix is sessile 

and these is no stipe, indicating a possible relationship 

with A. modestum or A. ovatum; this seems rather 

dubious and Brown s.n. does not appear to have a 

stipe. 

Dubious Binomials 

1. Aglaonema discolor 

Aglaonema discolor Hort. ex. Gentil, 1907, p. 11.-Engler, 

1915, p. 34. [Nomen nudum.] 

2. Aglaonema haenkei 

Aglaonema haenkei (Presl) Schott, 1859, p. 30. 

Caladium seguinum sensu Presl, 1827, p. 148 [not Ventenat, 

Scindapsus haenkei Presl, 1851 [“1849”], p. 240. 

18001. 

TYPE.-Philippines, Haenke s.n. (not seen) . 

According to Steenis-Kruseman ( 1950, p. 209)’ 

about 9000 of Haenke’s collections are in Prague, part 

in the National Museum, and part in the Charles 

(German) University. Through the kind efforts of Drs. 

A. Pilat and C. G. G. J. van Steenis, I had the onportunity to study all Presl types of Araceae from the 

Czechoslovakian National Museum. The type of A. 

59 

haenkei was not there. It may be at Charles University. 

A study of Presl’s description (1851) suggests that 

Scindapsus haenkei is not an Aglaonema and may belong to the Monsteroideae. The following is a translation of the description of the inflorescence and flowers : 

“Spadix cylindrical, a third shorter than the spathe, 

sessile, covered with bisexual [his “pseudohermaphroditus”] flowers: filaments 4, flat, fuscous; anthers cordate, bilocular; ovary quadrangular, cuneate, apex 

truncate and with a short stigmatic chink.” This description would fit a number of the Monsteroideae very 

nicely but none of the Philippine Aglaonema in which 

the spadix is not sessile, the flowers are clearly unisexual, the stamens are thick and never have flattened 

filaments, and the pistil is never quadrangular with a 

truncate apex. 

Despite this description and apparently without 

seeing the type of Scindapsus haenkei, Schott (1859, 

p. 30) transferred the epithet to Aglaonema. Study of 

the Schottian drawings in Vienna and Cuming’s specimens in Kew revealed that Cuming 863 (K) is the 

basis of Schott’s understanding of Scindapsus haenkei 

and not Haenke’s material. This material, as well as 

subsequent material cited by later authors as A. 

haenkei, is treated herein as A. densinervium. 

Cuming 863 (K), despite considerable usage as 

such, cannot even be considered as a neotype of 

Scindapsus haenkei since it does not agree with Presl’s 

description of the inflorescence. Without seeing typematerial I hesitate to treat A. haenkei as anything more 

than a dubious species although I strongly suspect it 

should be entirely excluded from Aglaonema. 

3. Aglaonema ?immaculatum 

Aglaonema ?immaculatum Hort. ex Engler, 1915, p. 33. 

TypE.-None cited, none found. 

Engler described his plant as having an abbreviated 

stem, a glaucous, purple petiole, and a blade that was 

glaucescent above and purple and glaucous below. 

The plant was of unknown origin and cultivated at 

the Berlin-Dahlem Botanic Garden. It had not 

flowered and Engler stated that he was not certain 

that the plant belonged to this genus. 

The information does not fit any Aglaonema known 

to me. It may be juvenile material of some species of 

Homalomena although glaucescence in that genus is 

not known to me. In view of the uncertainty, it is best 

to treat A. immaculatum as a dubious binomial. 
60 

4. Aglaonema maculatum 

Aglaonema maculatum Blume, 1837 p‘1835”], p. 154. 

Dracunculus Luzonis primus Camellus, 1704, p. 36. 

HOLOTYPE.-Plate 195 by Kamel (Camellus) in 

Hans Sloane MSS. No. 4082 (British Museum of Natural History, Bloomsbury) . This species began its career as a “species dubium” 

and has remained as such although Merrill interpreted 

it in 1923. His disposal, however, was not satisfactory. 

The only clues were given by Kamel ( 1704) : the plant 

came from Leyte and “Omauvana”; it was called 

“Gavay sin buaya” and “Panan quilon” ; the leaves are 

covered with white spots [“folia . . . albis conmaculata 

notis”] ; like “Pi16” and “Papaiya,” scrapings of the 

roots when mixed with a little salt are efficacious in 

treating skin problems caused by c’ontact with the 

irritating hairs [“pilosae”] of a plant called “Erucae 

Bad” 

SMITHSONIAN CONTRIBUTIONS TO BOTANY 

Merrill (1923) stated: 

For some years I thought that this [Le., A. commutatum] 

might be Aglaonema maculatum Blume. . . . I now suspect Aglaonema maculatum to be a form of Aglaonema 

oblongifolium Kunth [treated herein as A. marantifolium]. 

There is no evidence that A. commutatum Schott is of ancient 

cultivation in the Philippines. 

Merrill was not aware of the fact that A. commutatum is native to the Philippines. Also, Merrill did 

not realize that A. crispum, another species with 

variegated leaves, is native to the Philippines. 

Unfortunately, neither Kamel’s drawing n’or his 

description provide sufficient evidence for deciding 

whether A. maculatum really pertains to A. commutatum or A. crispum. 

5. Aglaonema treubii 

Aglaonema treubii Engler, 1898, p. 22; emended by Engler, 

1915, p. 19, fig. 5. 

LECT0TYPE.-Figure 5 in Engler (1915). 

In the original description of A. treubii, Engler cited 

two specimens, Koorders 16173 and 16146 from Minahassa, Celebes. He also noted : “Vide quoque specimina 

in horto Bogoriensi culta.” Later, Engler (1915) 

emended his description, excluded the specimens from 

the Celebes (transferred to A. haenkei) , and cited only 

“Bot. Garten Buitenzorg.” Unfortunately, the original 

material at Bogor is no longer extant. It seems logical 

to accept Engler’s (1915) drawing as a lectotype. The 

drawing of the inflorescence seems to me like A. 

simplex. The general aspect of the plant and the large 

number of pistils supports this disposition; however, it 

should be noted that Engler (1915) described the 

leaves as marked with irregular variegations along the 

midrib. Although this pattern is unknown to me in 

Aglaonema, except in A. vittatum Engler, it frequently 

occurs in Schismatoglottis and Dieffenbachia. 

The two syntypes of A. treubii, Koorders 16146 and 

16173, cannot be identified. The former collection 

(BO) has only seeds. The latter collection is only a leaf 

(L) and a sheet with labels (BO) . They do appear to 

belong to the genus Aglaonema. 

FIGURE 23.-Holotype of Aglaonema maculatum Blume. 

Excluded Species 

1. Aglaonema? cuscuaria (Aublet) Miquel, 1856, 

p. 217 [=Scindapsus cuscuaria (Aublet) Presl, 1851 

(“1849”), p. 2421. 

2. Aglaonema grifithii Schott, 1856, p. 123 [=Aglaodorum grifitlzii (Schott), 1858, p. 581. 
NUMBER 1 

3. Aglaonema mannii Hooker f., 1869, pl. 5760 [= 

Culcasia mannii (Hooker f.) Engler, 1887, p. 841. 

4. Aglaonema palustre Teijsmann and Binnendijk, 

1863, p. 399 [ = Aglaodorum grifithii (Schott) Schott, 

1858, p. 581. 

5. Aglaonema? pygmaeum Hasskarl, 1842, p. 10 

[ = Homalomena pygmaea (Hasskarl) Engler, 1898, 

p. 181. 

6. Aglaonema subundulatum Zollinger ex Schott, 

1856, p. 123 [ =&’chismatoglottis subundulatum 

(Schott) Nicolson, new combination]. 

Aglaonema subundulatum was provisionally published by Zollinger ( 1854, vol 1, p. 76) and was later 

validated by Schott (1856). A specimen of Zollinger 

I075 in the Drake herbarium (Paris) has Zollinger’s 

handwritten labels identifying it as Aglaonema 

subundulatum. It clearly belongs to the genus Schismatoglottis. Also at Paris there is a duplicate seen by 

Engler (unlike the specimen in the Drake herbarium) 

that lacks Zollinger’s identification and is the holotype 

of Schismatoglottis celebica Engler (1898, p. 19). 

Schott’s epithet clearly has priority over Engler’s and 

I, therefore, propose the new combination above. 

Identification List 

In this list there are ennumerated the fertile, numbered collections of Aglaonema that I have studied. 

Sterile specimens are usually omitted, as are specimens 

without a collector or number. The numbers in italics 

refer to the taxa as numbered in this work; types are 

indicated by “( T) ” after the collector’s number. 

The collector’s name is always used unless it did not 

appear on the duplicates (for example, “King’s Collector 10421” is used rather than “[Kunstler] King‘s 

Collector 10421”). Kunstler’s name does not appear 

on any of the duplicates of this particular number. Only 

by comparison of the date and locality of the collection 

with information given by van Steenis-Kruseman 

(1950, p. 282) can it be shown that Kunstler was Sir 

George King’s collector in this case. As another example, “Koorders 26731” is used rather than “Mangoenkromodiredjo, Koorders 26731” since the actual 

collector’s name appears only on the first duplicate. 

This policy means that any duplicate will immediately 

be recognizable from the citation (the purpose of citing 

specimens) although, in these very rare cases, the actual 

collector’s name is not cited. 

The number of a collection is cited directly after the 

collector’s name if the number belongs to a single numerical series belonging to the collector. If the collector, 

326-551 049-5 

61 

however, used several numerical series (i.e., Beccari 

used several series that were called “Piante Bomensi,:’ 

“Piante Papuanae” etc.) or the numerical series did 

not belong to the collector, then special abbreviations 

are interposed between the collector’s name and the 

number. These abbreviations are becoming standardized (Jacobs, 1968). Jacobs’ abbreviations have been 

adopted with the following additions : BGL=Bureau 

of Government Laboratories, Philippine Islands (this 

series was used by collectors such as Bolster and Elmer 

in the early 1900s). EIC=East India Company (this 

series was applied to specimens in the museum of the 

East India Company that were distributed and numbered by Wallich in the early nineteenth century ; the 

specimens were collected by many different collectors 

such as De Silva, Gomez, Griffith, Helfer, Porter, and 

Wallich in India, Burma, Pakistan, Nepal, and Malaya) . PGH = Philippine Government Herbarium 

(this series was used by collectors such as Copeland 

and Gregory in the early 1900s). REP=Reporter for 

Economic Products (this series was used in India, 

Pakistan, and Burma by collectors such as Burkill, Das, 

and Hooper, in the early 1900s). 

Abraham 3106: 16e; 3107: lb; 3108: 2b. Achmad 487 

(T), 1009 (T): 13. Adelbert 205: 13; 239: 15; 243: 13. 

Ahmad 89: 13. Ajoeb Exp. Jacobson 509: 6. van Alderwerelt 

van Rosenburgh 340: 2a; 244: 6; 245: la; 352: 17; 353: 6; 

355 (T) : 13; 356,357: 6; 358 (T) : 16c; 360,361 (T) : 13; 

368: 6; 369: 4; 372 (T): IOa. Alston 780: 17; 790: 6; 

792: 2b; 13564: lob; 13565,13566: 13; 13567: 26e; 13568, 

14017, 14601: 6;15396: 4;15398: 15;16053, 16431: 29; 

16477: 16b; 17072: 2a. Alvins 580: 13; 1887: 1Oa; 1974: 

7b; 2109, 2115: 13. Amdjah Exp. Genderen Stort 90: 13; 

Exp. Nieuwenhuis 384: 7b; 474, 482, 550: 1Oa. Anderson 

SAR 2812, 8023, 8056: 76; 11715: 13; 12331: 1Oa; 12533: 

13. Aiioneuvo PNH 13521 : 19; 13751 : 18. Asdat 53: 2Oa. 

Backer 7034, 14931, 21039, 23130, 23738, 24852, 30568: 

13. Bailey and Bailey 229: 16e; 230: 17. Baker 5575: 13. 

Bakhuizen van den Brink 1735, 2245, 3129, 4926, 6249, 

7691 : 13. Bakhuizen van den Brink Jr. 1339: 13; 2040: 16e. 

Balansa 266, 2049, 2052, 2064: 3. Bartlett 6649: 6; 6739: 

9.2; 6854: 16d; 7688, 7708, 8169: 6; 14663, 14889: 20. 

Beccari P.B. 159: 76; 1279: 1Oa; 1615: 13; 2051: IOa; P.P. 

798: 15. Beguin 1175: 15. Beum6e 1914, 2373, 3669, 5042, 

5684, 6750: 13. Bloembergen 4222: 20; 4230: 23; 4451: 

20; 4623: 13. Boerlage 522: 15. Bolster BGL 130: 20. 

Bon 2540, 3712: 3a. Borrsum Waalkes 263, 422: 23; 3107: 

15. Broadway 9180: 16e. Brooke 8450: 13; 8797, 8872, 

9382: 76; 9876, 10700: 13; 10717: 76. Biinnemeijer 513, 

630: 6; 1859, 2033, 2080: 13; 2102, 2105: 7b; 2215, 2232, 

3631: 13;4161,4283: 6;4311 (T): 13;4343: 6;4351 (T): 

13; 4499, 4636, 4671, 4992, 5477: 6; 5917: 23; 6601: 8; 

6876, 6953: 23; 7039 (T): 7b; 7044 (T), 7113: 13; 7161, 

7163: 8; 7179, 7189 (T): 13; 7345: 8; 7382: 13; 9105, 
62 

9261, 9391, 9510: 6; 7813: 13. Bunting 275: IOb; 276: 20; 

1346: 16h; 1369: 16m; 1370: 161; 1371: IOb; 1372: 16;; 

1457: 161; 1544: Ib; 1545: 16c; 1546: 161; 1548: 6. Burdick 4093: 17. Burkill SFN 912: 13; 3086, 3136, 6301: IOa; 

6524, 7018, 7648: 7b; R.E.P. 28046, 36623: 5; and Haniff 

SFN 12966: 13. H. Burkill 1862: 10a; 1888: 7b; and Shah 

186, 197: 13. Buwalda 4732, 5776: 15; 6338: 6; 6629: 8. 

CadiZre 27: 3a; 29: 4. Castro PNH 6498: 18; 6549: 20. 

Celestino and Ramos PNH 23072: 20. Chevalier 32003: 12; 

32011 (T): 11. Chun 6426: 4. Clarke 8266 (T) : 5. Clausen 

76: 4; 82, 85, 429: 16e. Clemens 1113: 15; 10003, 10442: 

13; 10490: 15;17640: 20;20686: 13;21950: IOa;22366: 

13; 22479: lob; 22555: 16d; 22556: 16e; 22581: 20; 

26142, 26703: IOa; 26820, 29133, 31149, 40985: 13. Cloun 

6426: 4. Collins 581, 766: 12. Copeland PGH 418 (T) : 19. 

Corner SFN 32462: 76; and Ngadiman SFN 37088a: 13. 

Cramer 20: 6; 52: 13; 53: 6. Cuming 863: 19. Curtis 1326: 

IOa; 1721, 1881 (at Kew) : 13; 1881 (at SING) : IOa; 2636, 

2679: 13; 2813 (T): 2a; 2834: 16d; 3820, 3821: 13. 

Dames 39: 6. Danser 7811: 13. Darnton 481: 13. Das 

REP 34952: 5. Derry 572, 613: 7b. De Silva EIC 4441: 5. 

Docters van Leeuwen 5384, 5412 (T) : 13. Dress 1667: I6e. 

Eberhardt 4196: 3. Edaiio BS 24790: 13; 24914: 19; 

26794: 20; 26856: 16e; 46049: ZO?; 76362: 16e; 77358, 

78290: 20; PNH 3589, 4118, 6731: 20; 15341: 18; 15544: 

19; 21909: 20; and Guttirez PNH 38450: 13. Elbert 3160, 

3169, 3207: 13. Elmer 6759: 16e; 10119: 19; 14993: 17; 

15706, 16520 (T) : 13; 17668: ZO?; 18182: 13; BGL 6433: 

20. Endert 4680: 13. Engler Exsicc. 73: 16e; 74 (T): 4; 

288: 13; Reise Java 4070 (T): 16d; 4080 (T): 2b; 4089 

(T) : 13. Evrard 478: 12. Eyma 2819, 3480: 15. 

Falconer 327: 13. FCnixBS 12585: 20; 26139: 19; 29899: 

18. Forbes 298, 1259: 13; 1685: 6. Forman 158, 321: 13. 

Fox PNH 4685, 4822, 4999, 8938: 20. Frake PNH 38043, 

38288,38338: 13. Franck 1377: 4. 

Galoengi 422: 13. Gamble 6766A, 7888: 5. Gibbs 6164: 

15. Godefroy 169: 12; 316: 11. Gomez EIC 8961B: 13. 

Gregory BGL 86 (T): 20. Griffith EIC 5981 (T): 13; 

5982 : 5; 5984,5985 : 13. Guillou 23 : 15. 

Haan 10, 48, 97: 76. Hallier 242, 676: 76; 1769: ZOa; 

2003: 7b; 3034, 3350: 10a. Hance 11459: 4. Haniff and 

Nur SFN 2906, 7159: IOa; 10124: 13; 10365: IOa. Harmand 1435 (Pierre 3347) (T) : 3. Helfer EIC 5992, 5993: 

13; 5994 (T) : IOc. Henderson SFN 18221, 18500, 20122: 

13; 21853, 22293, 22879, 29145, 29635: IOa. Henry 12287: 

3. Herre 1179: 20. Hewitt 11: 13; 13: IOa;43: 7b. Holttum 

9341, 10297: IOa. Hooker and Thomson 2131: 5. Hooper 

REP 25974: 5. Horsfield 193: 13. Hume 7245: 7b; 9260: 

IOa. Huttleston 1887: 13; 1907: 9. 

Iboet 244, 391: 13. Ilias Paie SAR 8320: 13. Irsan 34: 13. 

Jacobson 127: 6. Jaheri Nieuwenhuis Exped. 673 (T): 

7b; 1567: 13. Jelinek 196: 13. 

Kamphoevener 2578: 13. Kanjilal 6363: 5. Karta 75: 

16e. Kasin 607: 3. Keith 9419: 13; SFN 44593: 13. Kerr 

1160 (T): 4; 2070, 3005, 7010: 12; 7087: la; 7966: 11; 

9020, 10765 (mixed with 11): 12; 10864, 11558: IOa; 

12868: 11; 12893, 13968: IOa; 14369: 22; 14562: IOa; 

14658, 17581: 12; 18786: IOa; 18880, 19096: 12; 19365: 

21; 19440: 12; 19527, 19729: 11; 20370 (T): 36; 20525: 

SMITHSONIAN CONTRIBUTIONS TO BOTANY 

11; 20899: 3; 21263: 2b; 21776: 3. Kiah SFN 35006: IOa; 

35244: 12; 35294: 13; 36967 IOa. King’s Coll. [Kunstler] 

537 (T), 4023: 13; [Chaudering] 5652: IOa; [Kunstler] 

6158: 13; 10421 (T): 7b. Kjellberg 474: 20; 1768: 13. 

Kloss 55, 66: 13; 6724: IOa; SFN 14670: 13. Rondo and 

Edaiio PNH 38823: 20. Koorders 16126, 16129: 20; 16146 

(T) ;dub. bin. 5; 16155: 20; 16173 (T) : dub. bin.5; 16174: 

19; 16179: 13; 19434: 16b; 19646: 20; 20336, 20674, 

20700, 26105, 26240, 26241, 26731, 30001, 30105, 30809, 

30811, 31125, 31126, 32000, 33472, 33474, 33905, 34995, 

40678, 40698, 41196: 13. Kostermans 359: 13; 2881: 15; 

10522, 10569: 13; and Anta 244, 782: 13; et al. 218: 13; 

Kwae Noi River Exped. 359 (mixed with 3), 998: 2d; 

1129: 13. Kurz 240, 263, 1901, 2664: 13. Kuswata and 

Soepadmo 85: 15. 

Lahaie 931: 15. Lam 2625, 2988, 3155: 15; 3209: 20. 

Lauterbach K. Wilhelmsl. Exp. 325 (T) : 15. Leslie 306: 5. 

Liang 69877: 3a. Lister 22, 135, 239: 5. Larzing 4019: IOU, 

5043: 6; 5044: IOa; 5121: 2a; 5241: 13; 5352 (T): 10a; 

5357: 6; 5887 (T): IOa; 6278, 6987, 7338: 13; 11234: 6; 

12246: IOa;12537: 13.Loher2445: 16e;2447: 16b;2448: 

16d; 2449: 20; 2450: 2c; 14996: 161. Liitjeharms 3595: 

IOa?. 

Mabesa FB 26047: 20. McGregor BS 22847: 20; 23077: 

16d; 23084, 23088: 20; 32590: 19. Maliwanag 241: 20. 

Marcan 776, 1211: 12; 2659: 11. Meebold 15467: 13; 

15600: IOc; 17436: 13. Meijer 2392, 2425: 13; 3199: 6; 

4934, 5645: 13; 6431: 6; 6956: 13. Mendoza PNH 18422: 

20; and Convocar PNH 10286: 18. Mennega 373: Ib. 

Merrill BGL 3889: 16e; 4186: 20; BS 7347: 20; 11603: 19. 

Meyer FB 2486: 16e. Monchy 148: 13. Mondih 92: IOU; 

93: 13. Morse 374: 4. Motley 856 (T): 13; 1134: IOa. 

Mousset 838: 13. 

Nicolson 497: 4; 676: 6; 678: 17; 691: 20; 710: 19; 

705, 735: 18; 737: 19; 741: 16e; 742, 745: 19; 750: 16e; 

751: 20; 761: 17; 764: 18; 765, 766, 767, 768: 17; 783: 13; 

788, 789: 18; 790: 19; 795: 20; 797: 13; 801: 20; 801a: 

16b; 811, 812, 813, 824: 19; 829, 830, 833: 20; 955, 962: 

16c; 963a: IOa; 963b: 13; 964: 16e; 965, 984: 15; 986: 

13; 987: 6; 988: 13; 989: 17; 990, 991: 13; 992: 2a; 993: 

4; 994: IOa; 997: 2c; 1000: IOa; 1002: 13; 1004: 16d; 

1022: IOa; 1023: lob; 1094: 10a; 1114, 1115: 13; 1123: 

7b; 1134: 26; 1135: 13; 1136: I6m; 1137: 16d; 1146, 1147, 

11.57: 10a; 1179: 13; 1228: lob; 1238: lb; 1247: 16e; 

1254: 7b;1266: IOa;1291,1306: 13;1344: 7b;1345: IOa; 

1362: 13;1363: 7b;1406,1452,1478: 15;1603: 12;1607: 

4; 1608: 2b; 1615: la; 1616: 12; 1623: la; 1624: Ib; 1625: 

12; 1630: la; 1638: 12; 1660, 1681: 3a; 1692: 16k; 1697: 

4; 1698: 6; 1702: IOa; 1730: 9; 1745: 161. Nielsen 608, 

612, 694: 12. Nur SFN 18571, 34203: IOa; 47699: 2a; 

and Kiah SFN 7781 : 7b. 

Oostroom 12580: 13. 

Pancho 5287: 20. Peele 596: 13. Petelot 1371 (T): 3. 

Pierre 3350: 2b. Poilane 36: 3; 742, 8606: 12; 10746, 

10993,12010: la; 15290: 26; 15947: la; 17572: 2b; 20644: 

3; 22110: 12; 22899: la; 26752: 3; 27410: 12. Porter EIC 

4443: IOa. Purseglove 4537: 13; 5351: IOa; 5487: 13. Put 

616: 12;1065: 21;1214,1444: 12;1523: 21;1615,2749: 

12; 3050: 11. 
NUMBER 1 

Quisumbing, PNH 446 1 : 16e. 

Raap 216: IOa; 389: 13. Rahmat si Boeea 6030: 6; 7091: 

13; 10262: 6. Rahmat si Toroes 9, 173, 3481: 1Oa; 4982, 

4609: 13; 5031: 1Oa. Ramos 1670: 1Oa; BS 10882: 20; 

24141 (T), 30471 (T): 18; and Convocar BS 84010: 18; 

and Edaiio BS 36626: 14; 36751: 18; 36884: 19; 37359, 

48389: 13; 49059, 75580: 19; PNH 49829: IOa. Ridley 

1647: 13; 4604, 5107, 6002, 7667, 8081, 8082: 7b; 9508 

(T), 10144: 13; 11224: 76; 12416: 1Oa. Robinson 6362: 

13; BS 18139: 20. Rock 499: 12. Rostados 211: 7b. Royerr 

3519: 15. Russell 36: 21. Rutten 38: 13. 

Santos 5324: 20. Sarip Exped. R. Maier 102: 13. Saxton 

3004: 16d. Schlechter 16764, 18490: 15. Schmidt 90 (T), 

451a (T): 12; 675a (T) la. Schwarz 442: 4. Scortechini 

218: 1Oa; 255 (T): 7b; 548: 13; 1049: 1Oa; 1196: 13. 

Seidenfaden 2780, 2940: 12. Shah 344: IOU. Sinclair 5026: 

7b. Slooten 2141: 13. Smith 530: 13. Smitinand 1029: 1Oa; 

6778: 3a. Spire 174: 3a; 726 (T) : 4. van Steenis 647, 701, 

721: 13; 915: 7b; 1108, 3959, 5672: 13; 5979: 6; 6131: 

63 

13; 6407: 6?; 6414,8887,9899: 6; 12054: 13. Steiner PNH 

22929: 19; 36379: 13?. Sulit PNH 2806: 13; 3620, 3685: 

17; 6278: 18; and Conklin PNH 16836, 17656: 16m. 

Symington SFN 22669, 78552: IOa; and Agama 9335: 13; 

44523: 13. 

Taleon PNH 33970: 20. Thompson 9: 17. Thorel 1174: 

12. 

Umpol3 : 12. 

Valeton 183: 13. Vesterdal 60: 13. Voogd 2040: 13. 

Vidal 1241: 12; 1786: la; 1852: 2d; 1853: 12; 2508: 3; 

3970: 19; 3971: 20. 

Wallich EIC 8960A (T) : 7b; 8060B (T) : 21; 8961A: 5; 

8965A (T), 8965B: 1Oa. Wang 79372, 80747: 3. Warburg 

14604 (T): 14. Wenzel 277: 13. Williams 12: 16e; 2429: 

19. Winkler 2727 (T) : 13. Wray 2698: 1Oa; 2828,3123: 13; 

3585: 1Oa. Wyatt-Smith SFN 63169: 7b; 71016: 13; 71900: 

7b; 76462,7651 1 : 13. 

Yates 1228: 16d. 

Zollinger 531,695: 13. Zwickey 349: 13. 

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Index 

(New names and new combinations in italics; page num’bers of principal accounts in italics) 

Acorus, 6, 14 

Adpendix erecta, 5,47,48 

Aglaodorum, 3-4, 7, 19 

griffithii, 4, 7, 60, 61 

Aglaonema acutispathum, 26, 27 

alpinum, 46 

angustifolium, 7, 39,41,43-44,45 

birmanicum, 39,43 

borneense, 39,4 1,44 

brevispathum, 8, 11, 14, 17 (key), 19 (key), 20 (map), 

calamus, 9, 14 

var. undatum, 39,41,44 

22-23, 22 (fig.), 24, 25 

f. brevispathum, 19 (key), 23 

f. hospitum, 19 (key), 22-23 

brevivaginatum, 39, 41, 45 

brownii, 45 

cambodianum, 37 

clarkei, 28-29 

cochinchinense, 18 (key), 20 (map), 28, 34, 37 

commutatum, 7-10, 12-14, 16, 18 (key), 21 (map), 48, 

49-54,50 (key), 51 (fig.),60 

cv. ‘Albo-variegatum,’ 50 (key), 53 

cv. ‘Fransher,’ 50 (key), 53 

cv. ‘Malay Beauty,’ 50 (key), 53 

cv. ‘Parrot Jungle,’ 50 (key), 53 

cv. ‘Pewter,’ 50 (key), 53 

cv. ‘Pseudobracteatum,’ 14, 50 (key), 53 

cv. ‘Treubii,’ 50 (key), 53-54 

cv. ‘Tricolor,’ 50 (key), 53,54 

var. commutatum, 50 (key), 51 (fig.) 

var. elegans, 50 (key), 51-52,53 

var. maculatum, 14,50 (key),52 (fig.) 

var. robustum, 50 (key), 51,53 

var. warburgii, 50-51,50 (key) 

cordifolium, 18 (key), 21 (map), 4647 (fig.), 57 

costatum, 7-8, 11, 14, 17 (key), 18-20 (map), 23-25, 

23 (key), 28 

cv. ‘Foxi,’ 6, 7 

f. concolor, 23 (key), 24-25 

f. costatum, 23-24, 23 (key) 

f. immaculatum, 6,23 (key), 24 

f. virescens, 23 (key), 24 

var. foxii, 23,24 

var. immaculatum, 24 

var. inornatum, 24 

var. lineatum, 24 

Aglaonema costatum-Continued 

var. maculatum, 23,24 

var. virens, 24 

var. virescens, 24, 59 

var. viride, 19, 22, 26,27 

crispum, 7, 13, 14, 18 (key), 21 (map), 34, 54-55, 54 

curtisii, 53 

cuscuaria, 50, 60 

decurrens, 26 

(fig.), 60 

densinervium, 7, 16, 18 (key), 21 (map), 34, 46, 48, 

56-57, 59 

discolor, 59 

elegans, 51 

elongatum, 39, 41, 44,45 

emarginatum, 39,41,45,46 

fallax, 43 

gracile, 29 

grande, 39,41,45,46 

griffithii, 60 

haenkei, 7,56,59,60 

helferi, 37 

hookerianum, 8, 17 (key), 20 (map), 25, 27, 28-29, 41, 

hospitum, 7, 22 

immaculatum, 59 

integrifolium, 5, 17, 34,35 (fig.) 

laoticum, 26,27,28 

latifolium, 57 (fig. ) 

latius, 39, 41, 4546 

liangii, 26 

longicuspidatum, 39, 43 

maculatum, 5, 60 (fig. ) 

malaccense, 39,41,4243,45 

mannii, 61 

marantifohm, 5, 8, 11, 13, 14, 16, 18 (key), 20 (map), 

26, 34, 36, 41, 4749, 48 (fig.) 51, 52, 54, 56, 58, 60 

43 

var. obtusatum, 19,22 

f. robustum, 48 

var. commutatum, 50 

var. maculatum, 52 

var. tricolor, 53,54 

marmoratum, 58, 59 

minus, 13,30,31,46,59 

var. nanum, 31 

var. scortechinii, 31 

67 
SMITHSONIAN CONTRIBUTIONS TO BOTANY 

Aglaonema-Continued 

stenophyllum, 18 (key), 21 (map), 34, 55-56, 55 (fig.) 

subarborescens, 39,41,46 

subfalcatum, 38,39 

subundulatum, 61 

tenuipes, 18 (key), 21 (map), 22, 25, 28, 38-39, 41, 44 

treubii, 7, 53,60 

versicolor, 7, 9-10, 29 

vittatum, 13, 14, 18 (key), 21 (map), 31, 32, 33, 60 

warburgii, 50, 51 

section Chamaecaulon (see also Chamaecaulon), 4, 18-19, 

Aglaonema section Aglaonema, 17 (key), 25 

59 

Aglaonemateae, 3 (key) 

Alocasia, 1 1, 14, 16 

Amauriella, 3, 4 

Ambrosinia, 6 

Amorphophallus, 14 

Anadendrum marginatum, 35 

Anchomanes, 3 

Anubiadeae, 3 (key) 

Anubias, 3, 7, 19 

Araceae, 3-6,10, 14 

Arum, 42 

aquaticum, 5,47, 48 

integrifolium, 5, 17, 34,35, 36 

ovatum, 5, 41 

Caladium 

princeps, 34, 36 

seguinum, 59 

simplex, 39 

angustifolia, 34 

humilis, 34 

nitida, 5,33,34-35 

oblongifolia, 5, 35, 36,47, 48 

picta, 5, 29 

Cannaceae, 10 

Chamaecaulon, 1, 4, 11-13, 17 (key), 18-19, 22, 33, 59 

Colletotrichium, 16 

Colocasieae, 3 

Colocasoideae, 11 

Culcasia mannii, 61 

Calla 

Aglaonema-Continued 

modestum, 7-10, 12, 16, 17 (key), 20 (map), 25, 26-28, 

27 (fig.), 41, 59 

cv. ‘Variegatum,’ 26, 28 

nanum, 30,3 1 

nebulosum, 8, 18 (key), 21 (map), 30-32, 31 (key) 

f. nanum, 31-32,31 (key), 59 

f. nebulosum, 31 (key, fig. ) 

nicobaricum, 39,41,43,44-46 

nieuwenhuisii, 39, 41, 46 

nitidum, 5, 8-10, 13-18 (key), 20 (map), 29, 33-37, 34 

(key), 35 (fig.), 41, 48, 49, 55, 56, 58 

var. helferi, 18, 34 (key), 37 

var. nitidum, 34 (key), 36 

f. curtisii, 14,, 35 (key), 36-37 

f. nitidum, 34-36,34 (key) 

novoguineense, 47, 48 

oblanceolatum, 34, 36 

f. maximum, 34,36 

oblongifolium, 5, 17, 34, 35-36, 35 (fig.), 44, 47, 48, 60 

var. curtisii, 36, 37 

obovatum, 3 1,32 

ovatum, 17 (key), 20 (map), 25-26, 25 (key), 27, 28, 

38, 39, 41, 43, 59 

f. maculatum, 25 (key), 26 

f. ovatum, 25 (key), 26 

var. maculatum, 26 

palustre, 61 

philippinense, 18 (key), 21 (map), 34, 44, 47, 48, 55, 56, 

57-58, 57 (fig. ) 

pictum, 7-10, 12-14, 

cv. ‘Tricolor,’ 53 

var. nanum, 3 1 

var. scortechinii, 3 1 

var. tricolor, 29, 53 

pierreanum, 38-39 

princeps, 34, 36 

18 (key), 21 (map), 29-30, 31 

_. ~ 

propinquum, 39,41,43,45 

pumilum, 18 (key), 19, 21 (map), 25, 58-59, 58 (fig.) 

pygmaeum, 61 

ridleyanum, 19,22 

robustum, 51 

roebelinii, 7, 14,54 

lusus pseudobracteosum, 53, 54 

var. pseudobracteosum, 14 

rotundum, 11, 13, 18 (key), 19, 21 (map), 25, 31, 32- 

schottianum, 39,40,41-42,45 

33 (fig.) 

var. brownii, 39,41,45 

var. genuinum, 39 

var. malaccense, 39,45 

var. winkleri, 39,41,44,45 

f. angustifolium, 39, 44-45 

scortechinii, 31-32 

siamense, 22, 23, 38, 39,44 

simplex, 5, 7-11, 13, 14, 17, 18 (key), 20 (map), 26, 28- 

30, 34, 36, 39-46, 40 (fig.), 47-49, 55, 56, 58, 60 

f. inaequale, 40, 46 

f. macrophyllum, 44, 57 

f. typicum, 39 

Dieffenbachia, 3, 4, 6, 7, 11, 14, 15, 17, 54, 60 

Dieffenbachieae, 3 (key) 

Dracunculus Luzonis primus, 5,60 

Enallagma, 15 

Gleosporium graffii, 16 

Heteropsis, 11 

Homalomena, 4, 7, 11, 14, 15, 59 

angustifolia, 34 

brevispatha, 19 

humilis, 34 

pygmaea, 61 
NUMBER I 

Lagenandra ovata, 5,41 

Lasioideae, 3, 11 

Liliaceae, 16 

Monsteroideae, 13,59 

Montrichardia, 3, 11 

Nephthytis, 3, 14, 17 

Peltandra, 3,4, 7, 11 

Peltandreae, 3 (key) 

Philodendreae, 3 (key) 

Philodendroideae, 3 (key), 4,6,11, 13 

Philodendron, 3,4, 7, 10, 11, 15, 16, 17 

Piperaceae, 10 

Pistia, 3 

stratiotes, 5 

Plesmonium, 3 

Porphyrospatha, 14 

Pothoideae, 3 

Pothoidium, 11 

Pothos, 11, 14,42 

marginatus, 35 

Pseudohydrosme, 3 

Pythium, 16, 17 

splendens, 16 

Rhapidophora, 17 

Richardieae, 3 

69 

Schismatoglottis, 4,7, 11,60 

celebica, 61 

crispa, 54-55 

crispata, 54 

picta, 54 

robelinii, 54, 55 

siamensis, 22,23 

subundulatum, 61 

Scindapsus, 26 

cuscuaria, 60 

erectus, 47,48 

haenkei, 59 

tonkinensis, 26 

Spathiphyllum, 6, 14 

Symplocarpus, 6 

Syngonium, 16 

Tilia, 15 

Typhonium, 6,45 

Typhonodoreae, 3 (key) 

Typhonodorum, 3, 11 

Xanthosoma, 6 

Zantedeschia, 4, 7, 11, 16 

Zantedeschieae, 3 (key) 

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