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Full text of "Phytologia"






vol. 1? 
1Q68 




PHYTOLOGIA 

Desigtied to expedite botanical publication 



\o\. 17 J"ly> 1968 No^ 



CONTENTS 

GUNN, C. R., >iotes on the use of Ca'^^ in determining leaf 

thickness ^ 

MOLDENKE, H. N., Additional notes on the genus Vitex. \III. ... 8 



LIBRARY 

AUG 25 19C9 

NEW YORK 
BOTANICAL GARDEN 



Published by Harold N. Moldenke and Alma L. Moldenke 

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Price of this number, Si; per volume, S6.75, in advance, 
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NOTES ON THE USE OF Ca*^ 5 in DETERMINING LEAF THICKNESS 
Charles R. Gunn l/ 

Abstract 

The usefulness of leaf thickness as taxonomic character 
caji be enhanced by a simple, rapid method of determining dry- 
leaf lamina thickness. Several experiments were conducted with 
Calcium 5 testing the principle that thicker laminae absorb 
more soft beta particles than thinner laminae. By measuring 
the chajiges in particle intensity, mass is obtained; to the de- 
gree that thickness correlates with mass, one has determined the 
thickness. Shards of Magiolia grandiflora L., Asimina triloba 
(L. ) Dunal, suad Vicia americana Willd. were used. Results of 
these tests indicate that there is a correlation between the a^ 
mount of beta particles passing through the laminae of dry 
leaves and the thickness of the laminae. 

Introduction 

Leaf characters are commonly used in taxonomic treatments. 
Leaf thickness, if reported, usually is determined by inspec- 
tion. Aside from this empirical method, there is apparently no 
simple method of ascertaining thickness. Thickness has been 
determined by 1) use of cursory examination, 2) use of either 
freehand or microtome sections in conjunction with an optical- 
measurement system, 3) use of a micrometer or, k) use of a 
punch-weigh system. The section-optical-measurement system is 
not simple; the micrometer method results in large error; and 
the punch- weighing system is time consuming and cannot be ap- 
plied to leaves with narrow laminae. The usefulness of leaf 
thickness as a taxonomic character would be augmented if objec- 
tive measurement techniques were available. 

Radioactive isotopes, as the soirrces of beta particles for 
determining thickness, have had practical application in indus- 
try. Zumwalt (195^) reported two common uses of beta particles 
in industry. These are the determination of the thickness of 
continuously moving materials, and the concentrations of solu- 
tions. Measurements are based on two principles. The absorp- 
tion of the beta particles as they pass through a material 



1/ Research Plant Taxonomist, Crops Research Division, 
Agricultural Research Sei^ice, U.S. Department of Agriculture, 
Beltsville, Maryland 20705 

1 



2 PHYTOLOGIA Vol. 17, no. 1 

provides an indication of the thickness of the material. The 
degree of backscattering of beta pajrtides from a substance 
also ceui be used to determine thickness. 

The principle of absoj*ption of beta particles as they pass 
through a material is applicable to leaf studies. The few in- 
vestigations that have been published are concerned with water 
content of leaves, leaf thickness being a complicating variable 
(NaJtayama and Ehrler, I96U; and Yamada, et al., I956). No ref- 
erence has been found to studies undertaken from the teuconomic 
point of view. 

When a leai" is irradiated with beta particles, the inten- 
sity of the rays decreases as a result of interaction with the 
leaf. Thickness may be calc\ilated from the equation 

1=1 e-^^ 

o 
where d is the weight per unit area and }i the absorption coef- 
ficient (a constajit, which is determined only by the maximum 
energy of the beta particles and is peculiar to the nuclide 
used); for Ca^5 it is 0.128. The logarithmic constant e is 
2.718, I is the unmodified intensity, and I is the modified 
intensity. When d=0, 1=1 . Therefore when thickness is meas- 
ured by using beta particles, the expression is not in units of 
distance, but in units of weight per unit area, e.g., milligrams 
per square centimeter. The resiilts can also be reported as 
counts per unit of time from a standardized source. By insert- 
ing the leaf between the radiation source and the Geiger-Muller 
(GM) tube, the intensity of the radiation changes proportion- 
ately to the leaf mass. Thus by measuring such intensity 
changes, mass is obtained; to the degree that thickness corre- 
lates with mass, one has determined the thickness. 

Not all beta soxirces are amenable to herbarium leaf stud- 
ies. Soxrrces such as Strontium^^, Yttrium^*^, and Radium D&E 
(all used in industry) are not useful in leaf studies because 
of the strong penetration capacity of their beta particles. 
Sources such a^ Calcium 5 and Sulpher35 are applicable because 
of the weaker penetration capacity of their beta rays. The for- 
mer have been labelled hard sources, the latter soft sources. 

Materials and Methods 

A Radiation Coiinter Laboratories Scaler-ratemeter, model 
2032U was used to measure and record the beta radiation (Fig. 
1). The Itad shields covering the plastic planchets (Fig. 2) 
holding the sources contained 300 milligrams of lead per square 
centimeter. This thickness of lead absorbed the radiation from 
all soxorces tested. The holes drilled through the shields were 
1/16 inch in diameter. These holes allowed the passage of beta 



1968 Gunn, Determining leaf thickness 3 

rays from the source through the shield and test material to the 
GM tube. Since the isotopes were not uniformly distributed in 
the matrices, the shields were taped to the planchets. The 
distance between the soiirce and the test material wa^ 2.5 mm.; 
the distance between the tested material and the GM tube, 11 
mm. The background count averaged 9.2 counts per minute with a 
range of 11.8 to 6.3 counts per minute. The high voltage var- 
ied from 810 to 830, usually holding steady at 820, a setting 
recommended by the manufacturer. 

Sixty herbarium sheets of Vicia americana and its varieties 
(Gunn, 1968) were selected to represent the variation of leaf- 
let thickness in its North American remge. The leaflets were 
selected at random from these sheets. The co\int per minute 
from the open source was i 2700. 

A single leaflet of Vicia americana Willd. was tested for 
one half hour; readings were taken every minute using a 1-hole 
plate. When the resulting information was analyzed by means 
of maximum curvature, it was found that a 3-minute count inter- 
val was sufficient. 

The thickness of V. americana leaflet shards was also meas- 
ured by using a compound microscope equipped with an ocular 
micrometer and an oblique above-stage microscope light. The 
measurements were recorded in increments of 11.1 microns, round- 
ed to the nearest whole niimber. 

Results and Discussion 

Saran Wrap with Car'^ as the beta producing isotope vas used 
to test the equation I=lQe~^ . In a sequence of tests, layers 
of Saran Wrap were added (from 1 to 13) to the top of a one-hole 
plate, and readings were taken every 3 minutes. In Fig. 3 the 
layers of Saran Wrap were plotted against the log of the counts 
per minute producing nearly a straight line. These results il- 
lustrate that thickness cem be determined by counting the beta 
particles that are not absorbed by the test material. The line- 
ar arrangement of the averaged counts per minute in Fig. 3 
proves this point. The extension of this concept from a homo- 
geneous material (Saran Wrap) to a hetergeneous material (leaf 
laminae) was tested. 

Radium D&E, Carbon 1^+, and Calcium 1+5 were surveyed with 
shards of two test leaves taken from herbarivim (dry) material 
possessing obvious differences in thickness. Magnolia grandi - 
flora (magnolia) and A simina triloba (pawpaw). Of the three 
isotopes used, only Ca 5 gave results which were commensurate 
with the 8.1 thickness ration of dried meignolia and pawpaw. 
The results from the Radiim D&E test were inconclusive, since 
there was more intra- than inter-leaf variation. Readings 



U PliYTOLOGIA Vol. 17, no. 1 

obtained from C-'-'* were too close to the background count to be 
usable . 

Before testing CaT^ on the other leaves, lead shields with 
1, 2, and 3 holes (Fig. 2) were used with the magnolia and paw- 
paw leaves. In Fig. h each dot represents five, 3-mlnute counts 
averaged. Ba^ed on the results of the 3-hole test when compared 
to the 1-hole test, the Car^ concentration was trebled In Vlcla 
eunerlcana leaflet tests. This increased the 1-minute count 
through a l/l6 Inch diameter aperature from ± 365 to i 2700 
counts per minute, a seven- fold Increase. 

Magnolia shards with the red Indumentum of hairs Intact 
absorbed as much beta radiation as the same shards when denuded. 
This Indicates that pubescence Is not a factor affecting the 
outcome of this type of thickness determination. 

When the leaflets of Vlcia amerlcana were introduced Into 
the system, the counts remged from 846 (the thickest leaflet) to 
1874 (the thinnest leaflet). These coiints were converted to 
logs and plotted against the measurements recorded in microns 
obtained from the optical system. These resiolts are given in 
Fig. 5. The larger dots represent the 95 percent confidence 
limits of the population means. The means are represented by 
the smaller dots. The decrease in the co\ints per minute with 
the increase of leaflet thickness indicates a direct relation- 
ship between leaflet thickness and the amount of absorbed beta 
particles. A comrparison of Figs. 3 and 5 reveals that while 
the leaflet means are more variable than the Saran Wrap megms, 
the test did measure leaflet thickness. An analysis of the 
leaflet data indicates that 57 percent (r^=56.9'+) of the vari- 
ation in the counts per minute can be attributed to the thick- 
ness of the leaflets. 

The measurements in microns are at best an estimate. 
Therefore, the 57 percent correlation figure may be low because 
of errors in the measurement system. Additional tests on other 
leaves using other standards would help to establish the cor- 
rect correlation between true leaf thickness and the amount of 
absorbed beta particles. 

Literature 

Gunn, C. R. I968. The Vlcia americana complex (Leguminosae) . 
la. Jour. Scie. >2(3): 171-21 i+. 

Nakayama, F. S. and W. L. Ehrler. 196^. Beta ray gauging 

techniques for measuring leaf water contents changes and 
moisture status of plants. Plant Physiology 39(l):95-98. 



1968 Gunn, Determining leaf thickness 

Yamada, Y. S. Tamai, and T. Miyaguchi. I96I. A-I9. The 
measurement of thickness of leaves using s35. AEC-tr- 
kk&2. Translation Series. U.S.A.E.C. 

Zumwalt, L. R. 195^*^- The best performance from beta gauges. 
Nucleonics 12(1): 55-58. 



Mention of material by trade-name implies no preference 
over similar equipment made by other manufacturers. 



PHYTOLOOIl 



il w 



Vol. 17, no. 1 





Fig. 1. Radiation Counter Laboratories Scaler-ratemeter, model 
25321*. 





t^ 



--a 



Fig. 2 . A 2-hole lead shield taped to a planchet and carrying 
slide with a Vlcia amerlcana leaflet. 



1968 



Gunn, Determining leaf thickness 




liAHT THtCKNeiS n MKRONS 



a^^aloat tbe loga of tha count pi 
Klaut* vu t 270C. Tfa* l«rg«r ( 
coDfldsacB llalta of ttw populfti 
r«pr«a«Qt tba populatloo aaHia. 



Tba awtUar dota 



ADDITIONAL NOTES ON THE GENUS VITEI . VIII 
Harold N. Moldenke 



VITEX KUYLENII Standi. 

/Additional bibliography: Moldenke, Phytologia 16: ^02. I968 . 

Additional ciUtiona: GUATEMALA: Izabal: Jones & Face/ 3500 
(W— 2^65868); Jones , Proctor , & Facey 3031 (W— 2^6^867) . BRITISH 
HONDURAS: Gentle $$$1 (Mi). 

VITEX KWEICHOWENSIS P'ei 

Additional bibliography: Moldenke, Phytologia 15: 2^1. 1967. 

The Tsiang $831 collection, cited below, is marked "paratype" 
on its label, but the original description of the species by P'ei 
plainly designated Tsiang 6317 as the type collection. I see no 
valid reason for giving any other collection a type designation. 

Additional citations: CHINA: Kweichow: Tsiang $831 (W— l$75l$U) . 

VITEX LANUGINOSA Mohl 

Synonymy: Vitex laiiuginosua Mohl, Beitr. Anat. & Physiol. Geir. 

8$. I33li. 

Bibliography: Uohl, Beitr. Anat. Sc Physiol. Gew. 85. I83lii 
Kohl, Ann. Sci. Nat. Bot., ser. 2. 3- 319. 1335; Selling, Bishop 
Uus. Spec. Publ. 38: 27li, 275, & Ull. 19U7. 

I know nothing about this plant beyond what is given in the 
bibliography above. It seems most probable that the binomial is 
the resxilt of a typographic error or an error in copying. 

VITEX LEUCOXILON L. f ., Suppl. PI. 293. 1781 [not V. leucoaylon 
Blanco, 1895, nor Naves, 1918, nor Roth. 1956, nor Roxb., 
l8Ui, nor Span., 1856, nor Schau., 1893 J . 

Additional synonymy: Vitex leucoaylon Willd. ex Roxb., Fl. 
Ind., ed. 2 [Carey], 3t 7U— 75. 1832. 

Additional & emended bibliography: J. F. Qael. in L., Syst. 
Nat., ed. 13, pr. 1, 2: 963 (1789) and pr. 2, 2: 963. 1796; Pera., 
Sp. PI. 3: 361. 1819; Steud., Nom. Bot., ed. 1, 888. 1821; Roxb,, 
Fl. Ind., ed. 2 [Carey], 3: 7U & 75. 1832; Gamble, Man. Ind. 
Tlmb., ed. 1, 298. 1881; Watt, Econ. Prod. India $: 29h (I883), 
6: 191 (1883), and 7: 255. 1883; Gamble. Man. Ind. Timb., ed. 2, 
5U2. 1902; Prain, Beng. PI., ed. 1, 2: 832 & 833. 1903; Gamble. 
Fl. Presid. Madras 2: 1102 & 1103 . 1921^; Stapf, Ind. Lond. 6: U78 
& li89. I93I; H. F. MacMillan, Trop. Plant, k Card., ed. 5, pr. 3, 
197, 193, St 529. 1962; Prain, Beng. Fl., ed. 2, 2: 621, 622, & 
1012. 1963; Sen St. Naskar, Bull. Bot, Surv. India 7: 60, 1965; Se- 
bastine & Ramamurthy, Bvill. Bot. Stitv. India 8: 130. I966; Molden- 
ke, Phytologia 15: 253 & 316 (I967) and 16: 500 & 501. 1968. 

Jain (1963) records this species from Madhya Pradesh, India, 
while Sebastiiie & Ramamurthjy found only a "few" in Madras, citing 
a National Herbarium number I6096. Prain (I963) records it frco 



1968 Moldenke, Notes on Vitex 9 

Orlssa, but comments "on islands in the river Mahanadi; perhaps 
only introduced". He cites the first of the Watt references 
given by me in the bibliography above as •♦£, D, ^: 160", but 
this appears to be a paragraph reference, not a page referencel 
An additional vernacular name recorded for the plant is "kaddu- 
nochchi" . 

Additional citations: CULTIVATED: India: Herb, Drake s.n. 
[Hort. Bot. Calcutt.] (W--2li97125) . 

VITEX LIMONIFOKLA Wall. 

Additional synonymy: Vitex Hmonifolia "Wall, ex Kxirz" apud 
Deb, Bull, Bot, Surv. India 3: 315. 1961, Vitex aminifolia Wall,, 
in herb. 

Additional bibliography: Gamble, Man, Ind, Tim.b,, ed, 1, 296 
(1881) and ed, 2, 5Ul. 1902j A. Chev., Cat. PI, Jard, Bot, Saigon 
36. 1919; Deb, Bull, Bot, Surv, India 3: 315 . 1961; Moldenke, 
Pl^ologia 15: 253--25U. 1967. 

Chevalier (1919) records this species as cultivated in Viet- 
nam under the coimnon name of *'binh linh ving'* , In Burma it is 
called "kyungaukmre" , In Thidland the name "tin nok" is applied 
both to this species and to V^ peduncularls Wall, Deb (I96I) 
says of the plajit "shoots hairy or wooly, petiole broadly -ninged, 
panicles long branched, fulvous hairy" and cites Mukerjee 29U3 » 

Banterngsuk describes the plant as a large tree, common in 
dry deciduous forests in Thailand; Rock also refers to it as a 
cosranon tree in that covmtry. It has been collected in anthesis 
also in July and December. The corollas on Banterngsuk 6 are de- 
scribed as having been "purple". 

Additional citations: BURMA: Herb, Burma Forest School 22 
(W— 17l66Ui); Huk 8,n, [Bunna, I89O] (W— 7389I) . THAILAND: 
Banterngsuk 6 [Herb, Roy, Forest Dept, 1991] (W— 2061^782) ; Rock 
U66 (W— 1171368, W— 1171369), 

VITEX LONGISEPALA King & Gamble 

Additional bibliography: Moldenke, Phytologia 15: 251i — 255 & 
325. 1967. 

VITEX LUCENS T. Kirk 

Additional bibliography: Allan, Fl, N, Zeal, 1: 9$9 — 960, 
1961; D, Price, Contrib, N, S. Wales Nat. Herb. 3: 19U. 1961; 
Seikel, Chow, & Feldman, Phytochem. $: h39 — li55. 1966; J, S. 
Beard, Journ. Ecol, $^: 277. 1967; Seikel, Chow, & Feldman, Biol, 
Abstr, U8: 9U50, 1967; Moldenke, Phytologia 15: 255—256 (1967) 
and 16: 501, I968. 

Seikel and her associates (I966) report that the wood of this 
species is a rich source of glycoflavonoid (C-glycosylflavonoid) 
compovinds , In addition to the previously described apigenin de- 
rivatives vitexin (U',^»7-trihydro2yl-8-C-glTicopyranosylflavone) 
and isovitexin (the 6-C-gluco8yl isoner), the corresponding lu- 
teolin derivatives orientin and isoorientin have been discovered. 
Compounds of vitexin and orientin, which have xylose attached to 



10 PHYTOLOGIA Vol. 17, no. 1 

the 8-gluco8yl group, are also present. The moat unioaual constit- 
uents are wight compounds which appoaj: to be 6,3-dl-G-glycoByl 
derivatives of apigenin and luteolin. Several conpounds in each 
series are inter-convertible in hot acidic solution. 

Beard (196?) speaks of a V^ glabrata which is one of the main 
members of the broadleaf tree level in Australia alone with Eu- 
calyptus and Termlnalla . He is undoybtodly here referring to V. 
lucena . 

VITEX MADIEN5IS Oliv. 

Additional bibliography: A, Chev,, Sudanla 1: 11. 1911j Mol- 
denke, Phytologia 1$: 257-r-260. 1967. 

VITEX MADIEWSIS var, BAUMII Weper 

Additional bibliography: Moldenke, Phytologia 15: 208 — 259. 

1967. 

Additional citations: ANGOLA: Ble-Cuando-Cubango : E^ £. Mendes 

2632 (Rf). 

VITEX MADIENSIS var. MILAN JIENSIS (Britten) Pleper 

Additional bibliograohy: Moldenke, Phytologia 10: 2$9-— 260. 

1967. 

The corollas on Lewalle 1115 are described as having been 

"rose violac5 clalr", on his 1328 as "bleutS", on 2296 as 
"blanc sale", and on 2355 as "blanc et bleu". This collector 
has encountered this plant growing at 900 meters altitude. 

Additional citations: BURUNDI: Lewalle U03 (Ac), 1115 (Ac), 
1328 (Ac, Rf), 2296 (Ac), 2355 (Ac, Rf ) . ANGOU: Huila: Goss - 
weller 13UU; (Rf ) . 

VITEX MASONIANA Pittler 

Additional bibliography: lloldenke, Phjrtologla 15: 260. 1967. 

Recent collectors describe this plant as a tree, 50 — 75 feet 
tall [Allen says "50 m.", but surely In error], with a trunk di- 
ameter of 6—15 Inches at breast height, coarse leaves, and fruit 
brown and "fruity In odor", green when immature, grovdng at the 
edge of roads, at 15— UOO metei^ altitude. In anthesis also in 
February and March, In immature fruit In June and In mature fruit 
In October. Allen describes It as "Infrequent" in Darl^n. The 
corollas are described as having been 'hrhlte" on Jjj^ A. Duke 8387 , 
"lavender" on P. H. Allen 265, and "blue" on ?. H. Allen li588 and 
J. A. Duke 978U. Vernacular names for the tree are reported as 
"cuajado" , "kwidl machl", and "pu-pu-chlru" . The specific epithet 
is often uppercased. Duke assures us that the tree is not used 
by the Chocol Amerinds in Panama, 

The H. Pittler 660U, distributed as V. masonlana , is actually 
V. florldula Duchass, gc 'iTalp, 

Additional citations: PANAMA: Darlen: P, H, Allen 265 (E— 
1191569), U588 (E— 1572218); J. A. Duke Q'^SlT^TT iyil^ (Ac), 
11^639 (Ac, E— 1909076)} Stem , Chanbers , Dwyer , & Ebinger 299 (E— 



1968 Moldenke, Notes on Vitex 11 

1757555), 903 (£—1757560). Panami: J_. A. Duke UlM? (E~ 
1909075) . COLOMBIA: Choc6: £. A. Duke 9WU (Oh) . 

VITEX MEGAPOTAMICA (Sprang.) Moldenke 

Additional bibliography: Schnitzl., Icon. Fam. Nat. Reg. Veg. 
137. 1856; Rosengurtt, Estud. Prad. Nat. Urug. 5: 39U. 19U6: Rios 
de Moura Baptista, Anais XV Congr. Soc. Bot. Bras. 200. 196U; 
Dcmbrowski & Kxiniyoshi, Araucariana 1: lU. 1967; Anon,, Biol, Ab- 
str, li8 (20): S.I8I, 1967} Rimpler & Schulz, Tetrahed, Lett. 22: 
2033—2035. 1967 J Rljnpler & Schulz, Biol. Abstr, U8: 9253. 1967; 
Moldenke, Phytologia 15: 261—263. 1967. 

Recent collectors describe this plant as a tree, 8 m, tall, 
growing in foi-ests and at forest margins, at 500 to 1000 meters 
altitude, with the vernacvilar names "flor anil" and "tarxiffla". 

Additional citations: BRAZIL: Parani: Hatschbach 15363 (W — 
256U72li). Rio Grande do Sul: Rambo 37993 (B), Ult520 (B), 
1;9270 (B), 51795 (B) . Santa Catarina: Smith & Klein IklSh (N) . 
ARGENTINA: Misiones: k^ G. Schulz 7151 (N) . 

VITEX MEGAPOTAMICA f . ALBIFLORA Moldenke 

Additional bibliography: Moldenke, Phytologia 15: 263. 1967. 

Additional citations: BRAZIL: Parand: Hatschbach 13392 (W — 
256U667); Hatschbach & GuimarSes I5l5l (W— 2563953, Z) . 

VITEX MICRANTHA Gttrke 

Additional bibliography: Cave, Ind. PI. Chromosome Numb. 1: 
5U. 1958; Moldenke, Phytologia 15 : 263— 26U & 31ii. 1967; N. H. 
A. Cole, Bull. Inst. Fond. Afr, Noire 30: 107, 1968, 

Cole (1968) reports that this species grows among trees in 
matured secondary forests on slopes in Sierra Leone, flowering 
in February and March. Cave (1958) reports the diploid chromo- 
some number for the species as 32. 

VITEX MOLLIS H.E.K., Nov. Gen. & Sp. PI., ed. folio, 2: 199. 
1817. 

Additional & emended synoriymy: Vitex mollis Humb, & Bonpl, 
apud Steud., Nom. Bot., ed, 1, 888. 1821. Vitex trifolla Sess6 
& Moc. ex Moldenke, Prelim. Alph, List Invalid Names $2. in 
syn, I9U0 [not V. trifolia Graham, I966, nor Kemsl., 19u9, nor 
L,, 1753, nor L. f., 1895, nor Moon, 1895, nor Vahl, 19li, nor 
"sensu Matsumura ^ Hayata", I963] . 

Additional & emended bibliography: H.B.K,, Nov. Gen. & Sp. 
PI., ed. folio, 2: 199 (1817) and ed. quart., 2: 2ii5. I8I8; 
Steud., Nom. Bot., ed. 1, 888, 1821; Bamhart, Bull, Torrey 
Bot. Club 29: 590. 1902: Moldenke, Phytologia 15: 26U— 267 
(1967) and 16: 1;95. 1968. 

It should be noted that the H.B.K. reference dates given a- 
bove have been authenticated by consultation of the work on 
this subject by Bamhart (1902). 

The corollas are described as having been "purple" on J_. 



12 PHYTOLOGIA Vol. 17, no. 1 

Rzedowski 2207 and the plant waa collected in a deciduoua tropi- 
cail forest. A note on the sheet states that "one digit [is] mis- 
sing in fthe] collection no. given". What that missing digit ia 
has not been determined. 

Additional citations: MEXICO: Chiapas: F. Miranda 818U (W— 
2508351). Mexico: J_. Rzedofwski 2207 (Mi). Moreloa: Pringle 6993 
(Ms— 309U8). 

VITEX MOMBASSAE Vatke 

Additional bibliography: Watt & Breyer-Drandwijk, Med. L Poi- 
son. PI. S. Afr., ed. 2, 10^5 & lli5U. 1962; Moldenke, PhTtologia 
1$: 266—267. 1967; Friedrich-IIolzhannner in Mersm,, Prodr. Fl - 
SUdw. Afr. 122: 10. 1967. 

Additional citations: ANGOLA: Huila: E. £. Mendes 162$ (Rf ) . 
PORTUGUESE EAST AFRICA: Mozambiqu*: M. F. Correira 119 (Rf ) . 

VITEX NEGUNDO L., Sp. PI., ed. 1, 638. 1753 [not V. negundo Cur- 
tis, 1832, nor Lour., 193U, nor Noronha, 1790]. 

Additional & emended synonymy: Vitex negunda Willd. ex Rox±)., 
Fl. Ind,, ed. 2 [Carey], 3: 70. 1832 [not V. negwxia Mill., 1768]. 
Vitex leucoxylon Blanco apud Jacks, in Hook, f , & Jacks., Ind. 
Kew,, pr. 1, 2: 12lU, in syn. 1895 [not V. leucoxylon L., 1829, 
nor L. f ., I78I, nor Roth, 1956, nor Roxi)., 1811;, nor Schau., 
1893, nor Span., 1856, nor Wall., 18U7, nor ?^illd., I832] . Vit«x 
negundo L. f. apud Naithani, Bull. Bot. Surv. India 8: 260. I966. 
Vitex trifolla Graham ex Chavan & Oza, Mahar. Savaj. Univ. Baroda 
Eot. Mem. 1: 18?, in syn, I966 [not V. trifolla Hemsl., 19U9, nor 
L.. 1753, nor L. f., 1895, nor Moon, 1895, nor Sess4 & Moc,, 
19U0, nor Vahl, I9UI, nor "sensu Matsumura & Hayata", I963] . 

Additional & emended bibliography: J. F, Gmel. in L., Syst, 
Nat., ed. 13, pr. 1, 2: 963 (1789) and pr. 2, 2: 963. 1796; Pers., 
Sp. PI. 3: 361. 1819; Steud., Norn. Bot., ed. 1, 888. 1821; Roxb., 
Fl. Ind., ed. 2 [Carey], y. 70 & 71. 1832; Hook. & Am., Bot. 
Beech. Voy. 206. I836; Schnitzl., Icon. Fan. Nat. Reg. Veg. 137. 
1856; Gamble, Man. Ind. Timb., ed. 1, 297. 1881; Watt, Econ. Prod. 
India 5: 29U (1883) and 7: 255- 1883; Vidal, Phan. Cuming. Philip. 
I3I4. 1885; Watt, Diet. Econ. Prod. India 6 (li) : 2li8— 250. 1893; 
Gamble, Man, Ind. Timb., ed. 2, 539 — %0, 1902; Prain, Beng. PI., 
ed. 1, 2: 832 & 833. 1903; Duthie, Fl. Upper Gang. Plain 2: 22ii. 
1911; R. N. Parfcer, For, Fl. Pvmjab 39U. 1918; A. Chev., Cat. PI. 
Jard. Bot. Saigon 36. 1919; Gamble, Fl. Presid. Madras 2: 1101 & 
1102. I92I1; Hosokawa, Joiim. Soc. Trop, Agr. Taiiran 6: 206. 193iii 
Selling, Bishop Mus, Spec, Publ, 38: 27i4, 275, &: Ul. 19li7; Li & 
Keng, Taiwania 1 (2 — k)'. 127. 1950; Kuck & Tongg, Mod, Trop. 
Card. 77 & 236. 1955 J Encke, Pareys Blumengartn., ed, 2, Ui6, 
I96O; Cave, Ind, PI. Chromosome Numb, 2: 137. 1961; Deb, Bull, 
Bot, Surv, India 3: 315. 1961; H, F, MacMillan, Trop, Plant. & 
Gard., ed. 5, pr. 3, 198 & 366. 1962; Prain, Beng. PI., ed. 2, 
2: 621, 622, & 1012. 1963; Sharma & Mukhopadhyay, Joum. Genet. 
58: 359, 366, 376, 383, & 539, pl. 11, fig. 30. 1963} Maheshirari, 



1968 Moldenke, Notes on Yltex 13 

Fl. Delhi 281, 1963; A, Banerjee in Lahiri, West Beng. Forests 
56. I96U; Puri, Jain, Mukerjee, Sarup, & Kotwal, Rec. Bot. Surv, 
India 19: 107. 1961; j Cave, Ind. PI. Chrcmosome Numb. 2: 331 (1961;) 
and 2: U38. 1965 J Banerji, Rec. Bot. S\irv. India 19: 1$. 1965} 
Sen & Naskar, Bull. Bot. Surv. India ?: 60. 1965; M, S. Mani, 
Bull. Bot, Surv. India ?: HU. 1965; B, C, Stone, Micronesica 2: 
132, 1966; S, V, Ramasirami, Study Flow. Pi. Bangalore [thesis] 
xxix, 1029—1039, & lli67. 1966; Panigrahi, Bull. Bot. Surv. India 
8: 3, U, & 11. 1966; Panigrahi & Joseph, Bull. Bot. Surv. India 
8: 151. 1966; Matthew, Bull. Bot. Surv, India 8: l61i. I966; Bala- 
pvire, Bull, Bot, Surv. India 8: 190 & 19li. I966; Jain & De, Bull. 
Bot. Surv, India 8: 2k7 > 1966; Naithani, Bull. Bot, Surv. India 
8: 260. 1966; Rao & Rabha, Bull. Bot. Surv. India 8: 301. I966; 
J. L. Ellis, Bull, Bot. Surv. India 8: 337. 1966; Santapau, Bull. 
Bot. Surv, India 8: 39, 1967; Moldenke, Biol. Abstr. U8: 10560. 
1967; R. R. Stewart, Pakistan Joum. Forest. 17: 515. 1967; Mol- 
denke, Phytologia 15: 30U— 311 (1967) and 16: U93--lt95, 500, & 
501. 1968; Moldenke, Biol. Abstr. h?-' 851 (1968) and k9 (2): S. 
72 & S.186. 1968. 

It should be noted here that the Yitex trifolia of Hemsley, 
referred to in the synonymy above, as well as that of "sensu 
Matsumura & Hayata", is a s3monym of V. trifolia var. simplici- 
folia Cham,, that accredited to Moon is V. altissima L. f ,, that 
accredited to Vahl is V. tri flora Vahl, that of Sess6 & Mociflo 
is V, mollis H.E.K., while that of Linnaeus is a valid species, 
with the homonym accredited the Linnaeus the younger as a syno- 
nym. The V. leucojcylon of Linnaeus the younger is a valid spe- 
cies, with the homonym accredited to Linnaeus the elder as a 
synonym, as well as that ascribed to Willdenow, while the V, 
leucoxylon accredited to Schauer is V^ glabra ta R, Br,, that 
ascribed to Roth and to Roaburgh is V_, glabrata var. bombacifo- 
lia (Wall,) Moldenke, that accredited to Spanoghe is V. parvi- 
flora A. L, Juss. and that ascribed to Wallioh is V. leuco:ylon 
L. f. 

Aggarwal & Mukherjee (I963) state that this plant, along 
with Clerodendrum inerme , Cyperus stoloniferus , and Sporobolus 
maderaspatanus , play an important r8le in stabilizing dunes on 
Rameswaram Island and Krusadi Island, but surely the typical 
form of the species is not here being referred to — probably it 
is V. trifolia var, simplicifolia Cham, to which reference is 
here being made. 

Balapure (I966) records V. negundo from Madhya Pradesh, iriiere 
he fovind it growing in moist shady places along riverbanks and 
"very common" in waste places, along roadsides, and on riverbanks. 
Rao & Sastry (1961;) also refer to it as common along watercourses 
in that state. ELlis (I966) records it from Andhra Pradesh and 
cites a National Herbarium no. 15911. Jain (I963) found it in 
Gujerat, Rao & Rabha (I966) in Assam, and Santapau (I967) in Sau- 
rashtra. Bhattacharyya (1961i) reports it as "ccsnmon" in Uttar 
Pradesh. Panigrahi and his associates (1961i) found it to be "a- 



Hi PMYTOLOGIA Vol. 17, no. 1 

bundant on river banks" in Orisaa; Lau describee it as abundant 
in dry sandy soil on Hainan Island. 

Jain & De (I966) report that in West Bengal, where it is known 
as "begna" and "ichur", a decoction is made of the leaves w^iich 
is given with Andrographis and/or hyoscyamua to euro coughs, gout, 
and symptoms related to colds and tho leaves are uaed for fumiga- 
ting huts to remove flies and mosquitoes, citing Jain 7903 « Muk- 
erjee (1965) also avers that V. negundo is a conunon shrub in the 
villages of West Bengal. Kuck 1^ Tongg (195^) point out that it 
is wind-resistant and grows rapidly and irregularly. Janardhanan 
(1963) found it to be scarce in Uaharashtra, where it is called 
"nirgvid" or "nirgundi*', and where the leaves are used as a tonic 
and vermif\ige and the leaf-juice by the local population to re- 
move fetid discharges and worms from ulcers. 

Joseph (1963) found the plant "fairly conmon near streams" in 
Kadhya Pradesh, while Malick (I966) describes it as "common" in 
West Bengal and cites Chatterji 3. Panigrahi ^ Joseph (1966) 
claim that it is "abundant" in Nefa and cite a National Herbarium 
no. 16788. Panigrahi (I966) reports it as "abundant on dry open 
flat hilltops" and on hill slopes in Bihar and cites no. II89I. 
RamasweUJ^r (196ii) encountered it growing along riverbanks with 
Phyla nodi flora under a thick growth of Sallx tetrasperma in 
Bangalore and also in hedges there. Naithani (I966) refers to It 
as "rare" and cites no. 23873. Vidal (1885) cites Cuning I886 . 
Deb (1963) reports that the species inhabits moist and dr;' decid- 
uous forests. 

Maheshwari (I963) describes V. negundo as it occurs in Delhi, 
India: "Flower clusters lax, in a widely spreading panicle; leaf- 
lets mostly broader [than in V. agnus-castus ] .... Jl shrub or 
small tree. Branchlets quadrangiilar, densely white-tomentose . 
Leaflets 3 — 5, 10 — 17 x 2.5 — h cm., petiolulate, lanceolate, 
acuminate, white-tomentose beneath. Flowers lavender to blue, in 
loose clustei^, surranged in a large terminal panicle. Drupes 
black . Planted in gardens , lawns and along railway lines . Com- 
mon in the Bangar tract on raised bunds along the roads . The 
■warmed leaves are applied to painful and rheumatic swellings; the 
macerated ones are used as cooling medicine on the forehead in 
headache. Local name: Sambhalu. Flowers: Major part of the yevr J' 
He cites Maheshwari 118 & 689. Banerji (1965) cites his no. 536 . 

Prain (I963) records the species from Bihar said Chota Nagpur 
and gives the following additional vernacular names for it: 
"begunia" and "sandbhalu" . Banerjee (I961i) records the name 
"sindubara" for it, while Stewart (1967) tells us that the species 
is very conmon "in graveyards and near streams" in Swat, Pakistan, 
while Chevalier (1919) reports it cultivated in Vietnam under the 
name "cSy ngu triu" . 

Cave (1961, I96I4, 1965) reports the haploid number of chromo- 
somes as 12 and the diploid as 26 and 3U. 

MacMillan (I962) reiterates that in India the leaves and bark 
of this plant are used in the treatment of toothache, rheumatism, 



1968 Moldenke, Notes on Vltex 15 

and eye diseases, as weU as for a tonic, carminative, and vermi- 
fuge. Watt (1893) includes V. bicolor 7/illd. and V. arborea Desf. 
in the synoi^omiy of V. negundo. The latter, however, is a synomym 
of V. negundo f . alba P«ei, while the former is V. trifolia var. 
bicolor (Willd.) Moldenke 

It should perhaps be noted here that the V^ neg\indo accredited 
to Curtis is actually V. negundo var. heterophylla (Franch.) Rehd., 
that accredited to Loureiro is V. quinata (Lour.) F. N. Will., 
while that of Noronha is V_. pinna ta L, 

Encke (I96O) describes V, negundo as follows: "Indien bis Ost- 
asien und Malesien. Juli — ^August. Bis h m hoher, baumartiger 
Strauch. Blatter fingerfOrmig-^zShlig, mit linealisch-lanzett- 
lichen, gezShnten, S — 10 cm langen Blattchen. Blttten in end- 
standigen, 1^ — 20 cm langen zusammengesetzten Rispen, lila oder 
lavendelblau." 

The species has been collected in fruit in July as well as in 
the months previously recorded. Ching states that it is "common" 
along roadsides in Kwangsi, while Rodin describes it as "common 
along streaabanks" in Swat. The corollas are described as having 
been "pink" on Liang 6I466I , "purplish" on R. C. Ching 5U50, 
"blue-purple" on Koelz 10-37 , "blxiish" on Taam 1728 . and "blue" on 
Rodin 5U27, Tsang s.n, [Herb. Lingnan Univ. 16629], and E^ H. 
Wilson 10972 . A note accompanying Clemens L Clemens 380ii indi- 
cates that the species occurs wild and also in cultivation in 
Annam. 

Material has been widely misidentified and distributed in 
herbaria under the names V, agnus-castus L., V, incisa Lam,, V. 
negundo var. cinnabifolia (S, & Z.) Metcalf, V, trifolia L., and 
even Buddleia asiatica Lour, 

There seem definitely to be at least two fonns of what is cur- 
rently being regarded as the typical form of this species. One 
of these bears a striking similarity to the typical fonn of V, 
trifolia L. [ e.g. , Babu Ram 99, Hafizthan s.n, [Balakoli], Rodin 
gU27 , S_. N. Singh 18 [3,8. 2U & 11.1.2^], and R. R. Stewart 17067] . 

The other form is more typical of what I regard as V, neg^lndo 
in the strict sense. Examples are Barchet 5^6 , H. H. Bartlett 
6267, R. C. Ching $l;gO , Clemens & Clemens 380ii , Fraser I96, A. 
Henry 11U2 &; 97^0 , Koelz 3^7 , C, 0^ Levine s.n. [Herb. Canton 
Chr. Coll. 376], Liang 6U66I, Nagazawa s.n. [July 1928], Peng , 
Tak, & Kin s.n. [Herb. Canton Chr, Coll, 12670], Poilane 8I3O, 
Taam 1728, Tanaka 97, Tanaka & Shimada 17878, E. H. Wilson 1697 , 
and Ying 1263 . The complex needs further study. It is very 
possible that V, negundo and V, trifolia hybridize where they 
grow together. 

Barchet $$6 appears to be a mixture with var, cannabifolia 
(Sieb, & Zucc) Hand,-Mazz. The Rock 388O, $0$$ , & 90lUi , dis- 
tributed originally as V_, negundo, are all cotypes of f , alba 



16 PHYTOLOGIA Vol. 17, no. 1 

P'el; Ling 1721 ^ 3.n. [Herb. Univ. Nanking 9386], as well as 
Herb. Univ. Nanking 9366 , apjjear to be var. heteroph/lla (franch.) 
Rehd.; and Chiao 270U , Fu C. Chine 2U29, Farges 8.n. , Fung 21196 , 
Herfa. Canton Chr. Coll. 89U & 11^20 , Herb. Univ. Nankin£ 1726 , 2387 , 
lliO^a , Ut^ao , ^ 1877U , Petelot 1170 , Rock 698I, Tsang 27733 ^ 
278U3 , Taiang ^ P'ei £725, Tsui 303 , E. H. ffllaon 790 &■ 2702 , and 
Zimmermann 2 appear to represent var. intermedia (P'ei) Moldenke. 
It i3, however, very obvious that the named varieties grade into 
each other in most conTusing fashion. 

I am not at all certain of the tnie identity of the Herb. Post 
a.n. [Hamath, Aug. I88U] , cited below. It was originally identi- 
fied and distributed as V^ agnus-castus L., but most certainly 
cannot be that species in its restricted sense. It may be a ma- 
ture fiTiiting specimen of V, agnus-castus var. pseudo-negtmdo 
Hausskn., but it also greatly resembles V. negimdo . 

Additional citations: SYRIA: Herb. Post s.n» [Hamath, Aug. 
188U] (W— 805058). PAKISTAN: Swat: Rodin ^27~(W—22li2322) . 
INDIA: East Punjab: Koelz 3^7 (W— 1667937), Ul37 (W--1607992) . 
Mussoorie: R. R. Stewart I7067 (W— 1992176) , Siwalik & Jaunaar: 
Babu Ram 99 (W—ll 70327) . Uttar Pradesh: Crovalti 81 [July] (W— 
1372659), 81 [November] (W— 1372659)} Mohammed s.n. [13.7.29] (W- 
17166U5), s^n. [5.11.29] (W— 17l66Ii5); K. Singh 80 (w— 13U7706)} 
S. N. Singh 18 [3.8.2U] (W— 13li77U5) , 18 [11.1.25] (W—13U77U5) . 
CEYLON: Fraaer I96 (W--73890) . CHINA: Chekiang: Barchet 556 , in 
part (W— 596II8). Hupeh: E. H. Wilson 12U1 (W— 596717) , 2701 
(W— 777U69). Kiangsi: E. H. Wilson 1697 (W— 77729ii) . Kwangsi: 
R. C. Ching 5U50 (W~12li8670) . Kwangtung: C_. 0_. Levine s.n. 
[Herb. Canton Chr. Coll. 376] (W— 778695)} Peng , Tak , & Kiii a.n. 
[Herb. Canton Chr. Coll. 12670] (W--12U7923) ; Ying 1263 (W— 
1513156) . Ytlnnan: A. Henry 9750 (W— U57296) . CHINESE COASTAL 
ISLANDS: Hainan: S^ K. Law 298 (W~162916U); Liang 6U661 (W— 
1671297). Lantau: Tsang s.n. [Herb. Lingnan Univ. 16629] (W-— 
12li9326). HONGKONG: Taam 1728 (W— 22U4633) . INDOCHINA: Annam: 
Clemens & Clanens 38OU (W— ll;27683, W~ll;2768ii) j Poilane 8I3O 
(W— 239U576). WESTERN PACIFIC ISLANDS: FORLJDSA: H. H. Bartlett 
6267 (W~12li8580) J Nagazawa s.n. [July 1928] (W—2063380) j Naka- 
hara s.n. [1905] (W— 1053769); A. Henry 111^2 (W— 1;55567)} Tanaka 
97 (W— 1528110) } Tanaka & Shimada 17878 (W— I7OO296); E. H. Wil - 
son 10972 (W— 1051i28l) . CULTIVATED: India: Voigt 272 (W— 
2126892). LOCAKETY OF COLLECTION UNDETERMINED: Hafizthan a.n. 
[Balakoli] (W— 1239953). 

VITEX NEGUNDO f. ALBA P'ei 

Additional bibliogr^hy: Watt, Diet, Econ. Prod. India 6 (U): 
2U8. 1893; Moldenke, Phytologia 15: 308. I967. 

This plant has been collected at altitudes of 3000 to 10,000 



1968 Moldenke, Notes on Vitex 17 

feet, flowering in August. The corollas are descidbed as having 
been "blue" on Rock 10l;6$ , "bluish" on Rock ^0^, "pale-purple" 
on Rock 3880 , and "lavender-blue" on Rock 90141^ . It is therefore 
evident that the statement made by me in Phytologia 1$: 308 
(1967) concerning V. arborea Fischer and V. arborea Desf . belon- 
ging here because the represent white-flowered plants is entirely 
incorrect. If Schauer and Jackson are correct in placing these 
binomials in V. negundo, then they appear to represent a white- 
flowered fomTfor which I am proposing the name V, negundo f . 
albiflora Moldenke. 

Additional citaUons: CHINA: Yttnnan: J. F. C. Rock 3880 (¥— 
1332136— CO type) , |055 (W— 1332137— cotype) , 90141 (W— 13321138— 
cotype) , 10U6^ (W— 1332139) . 

VITEX NEGUNDO f . ALBIFLORA Moldenke, nom. nov. 

Synonymy: Vitex arborea Fischer ex Desf., Cat, Hort. Paris, 

ed. 3, 391—392. 1829 [not V. arborea Br^on, 1955, nor Brown, 
1806, nor Ro±b,, I81U] . Vitex arborea Desf. apud Schau, in A. 
DC., Prodr. 11: 685, in syn. 18U7. 

BibUography: Deaf., Cat. Hort. Paris, ed. 3, 391—392. I829i 
Schau. in A. DC., Prodr. 11: 685. 18U7} Watt, Diet. Econ, Prod. 
India 6 (U): 2U8. 1893} Jacks, in Hook. f. & Jacks,, Ind. Kew., 
pr, 1, 2: 1213 (1895) and pr. 2, 2: 1213. 19U6; Moldenke, Phyto- 
logia 55 li86. 1957; Jacks, in Hook. f. & Jacks., Ind. Kew., pr. 
3, 2: 1213. I96O; Moldenke, Phytologia 15: 308. I967. 

This form differs from the typical form of the species in 
having white corollas. 

As yet I have not seen the type of this taxon, doubtless 
preserved in the Paris herbarium, but I am assuming that Schauer, 
Watt, and Jackson are correct in placing it in V, negundo. 

It should be noted here that the V, arborea ascribed to Br^on 
belongs in the synor^yTijy of V. beraviensis var. ac\ipinata , that 
accredited to Brown belongs in the synonynor of V^ heptaphylla A. 
L. Juss,, while that of Roxburgh is Y. pinnata L. 

VITEX NEGUNDO var, CANNABIFOLIA (Sieb. & Zucc.) Hand.-^Iazz. 

Additional synonymy: Vitex negundo var, cinnabifolia (S. & Z,) 
Metcalf ex Moldenke, Phytologia 17: 15 & 17, in syn, I968 

Additional bibliography: Kitamura & Okamoto, Col, LLlustr. 
Trees & Shrubs Japan 221. I96OJ Moldenke, Phytologia 15: 308 
(1967) and 17: 15 & 17. 1968. 

An additional vernacular name recorded for this plant is 
"nindinboku" . Material of this variety has been widely misiden- 
tified and distributed as V. incisa Lam, and V, negundo f . inter - 
media P'ei, Zimmennann hh2 appears to be a mixture with var, in- 
termedia — at least, on most specimens the leaf serration seems 
to be far too uniform for var. intermedia . 

Additional citations: CHINA: Chekiang: Barchet 556, in part (W- 



18 PHYTOLOOIA Vol. 17, no. 1 

596117). Shantung: Zljmaennann hh2 , in part (W— 79$l?0) . HOICKDIO: 
C. Wright 8.n. [Hong Kong] (W--Ui?l6). WKoTEItH PACIflC I3LA!iI6: 
JAPAN: IIon3hlu: Collector undetermined 36$ (V^— 73902), 3.n. [Sept. 
1, 1890] (ff--206l82), s.n. [Yanaka, Muaashi, 18 August 1910] 
(W— 1178281) i £. Matsumura a.n. [Tokio, Octob. 13, 1879] (W— 
11^7605) i yaxljnowicz 8.n. [Yokohama, 1862] (7f— 73892) . 

VITEI NBGUNDO var. HETEHDPHYLLA (Franch.) Rehd. 

Additional (Sc emended synonymy: Vitex sinuata P.aeusch. ex 
Steud., Nom. Bot., ed. 1, 888. 1821. Vitex negundo Curtis ex 
Roxb., Fl. Ind., ed. 2 [Carey], 3: 72, in syn. 1832 [not V. ne- 
gundo L., 1753, nor Lour., 193U, nor Noronha, 1790, nor Royle, 
1919, nor Willd., 1918], Vitex incisa Y/illd. ex Roxb,, Fl. Ind., 
ed. 2 [Carey], 3: 72. 1632. Vitex chinensis Banks ex Ro:d)., Fl. 

Ind., ed. 2 [Carey], 3: 72, in syn. 1832. 

Additional bibliography: J. F, Gmel. in L., Syst. Nat,, ed. 
13, pr. 1, 2: 963 (1789) and pr. 2, 2: 963. 1796; Pers., Sp. PI. 
3: 360—361. 1819; steud.. Nam. Hot., ed. 1, 888. 1821; Roxb., 
Fl. Ind., ed. 2 [Carey], 3: 72—73. 1832; Watt, Diet. Scon. Prod. 
India 6 (U): 2U8 & 2^1. 1893; Bonstedt, Pareys BlumengSCrtn . , ed. 
1, 278. 1932; Encke, Pareys BlumengStrtn., ed. 2, 2: UU6. I960; 
Moldenke, Phytologia 15 : 307 & 309—311. 1967; Moldenke, Biol. 
Abstr. U9 (2): S.186. 1968. 

Encke (I960) says of this variety: "In Kultur wohl nur durch 
die strauchige var. heterophylla (Franch.) Rehd. (syn. var. in- 
cisa (Bunge) Clarke; V. incisa Bunge) . Nord- und Centralchina, 
Mandschurei, Philippinen. Mit eingeschnitten-gezSthnten oder 
fast fiederspaltigen, 2—8 cm langen Blattchen. — IM 1750. EJi. 
36U; N.K. Hi: 12; B.C. III:3U8l. (g) llur im Weinbauklima bedingt 
■winterhart. In kalten Y^intem auch dort immer wieder zurtlck- 
frierend, aber an einjSChrigen Trieben im gleichen Herbst noch 
bltlhend. SchOne Kerbstbltlher zur Verwendung in der NShe des 
Hauses auf der Gajrtentenrasse oder in Verbindung mit andem 
tropischen Blattpflanzen. Am besten ist frx>stfreie Uberwinter- 
\ing voad Mitte Mai Pflanzung ins Freie . Bei gute Pflege und Er- 
nShrung machen sie in wenigen Monaten lange SchOsslinge, die in 
warmen Sommern fast immer noch zur Bltfte koramen. Vermehrung 
durch Aussaat und ausgereifte, krautige Stecklinge Im Sommer." 

The varietj-- has been collected at 200 meters altitude in 
Shantting. An additional vernacular name for it is "mu chin". 
The corollas are described as having been "bluish" on Chiao 
3052 and as "lavender" on K_. H. Beach lli5 . 

The Herb. Hort. Bot. Petrop . s.n. , originally distributed as 
this variety, appears to be V. negundo var. heterophylla f . mul - 
tifida (Carr.) Rehd., £. 0_^ Levine s.n. [HeA. Canton Chr. Coll. 
250 & 1585] , Fung 21196 , J. E_. Norton 1558 . and W. T_. Tsang 
230U7 are V. negundo var. intermedia (P'ei) Moldenke, C^ 0^ Le- 
vine s.n. iHerb. Canton Chr. Coll. 376] is V. negundo L., and 



1968 Moldenke, Notes on Vitex 19 

CO, Levine s.n. [Herb. Canton Chr, Coll, 7U6] is V. s ampsonl 
Hance. The Zimmenaann Ui2 , originally distributed as this varie- 
ty, seems to be mostly var, cannabifolia (Sieb. & Zucc) Hand.- 
llazz., although it was identified by P'ei as his f , intermedia 
and at least one specimen of it has been so cited by me. Cer- 
tainly the collection is not var, heterophylla l 

Additional citations: CHIMA: Chahar: Kozlov 71 (W— 16585U9) . 
Hopeh: K. H. Beach U;$ (W— 2070711^) , 228 (W~2070785)i Chiao 227 • 
[Herb. Univ. Nanking 2138U] (W— l5^U26l) j Comiry s.n. [Vicinity 
of Peking, 1919] (W— 1051760); H. J. Sheehan 98 (W— 1576691) . 
Shansi: Ling lU67 [Herb. Univ. Nanking 9113] (W~1370U52), 1721 
[Herb, Univ. Nanking 9366] (W— 1370l;53), s.n. [Herb. Univ. Nan- 
king 9386] (W— 1370U5U) . Shantung: Chiao 30^2 (W— 1576506) . 
Province undetermined: Bunge s.n. [Chin. bor. I83O] (W— 2[i97090) . 

VITEX NBGUNDO var. IIETEROPHYLU f . ALBA (Carr.) Moldenke 

Additional bibliography: Moldenke, Phytologia 15: 310. 1967} 
Moldenke, Biol. Abstr. U?: 851. I968. 

VITEX NEGUNDO var. KETEROPHYUA f. MULTIFIDA (Carr.) Rehd. 

Additional synonymy: Vitex dissects Vasey ex Moldenke, Phyto- 
logia 17: 19, in syn. 19^. 

Additional bibliography: Moldenke, Phytologia 15: 310 — 311. 
1967. 

Material of this form has been distributed in herbaria under 
the name V. incisa Lam. 

Additional citations: CULTIVATED: District of Columbia: Vasgy 
s.n. [Greenhouse, 1881] (W~7389ii) . Russia: Herb. Hort. Bot. 
Petrop. s.n. (W— 73895) . 

VITEX NBGUNDO var. INTERMEDIA (P'ei) Moldenke 

Additional bibliography: S. V. Ramaswami, Stu(^ Flow. PI. 
Bangalore [thesis] IO3O— IO3I & lii67. I966j Moldenke, Phytologia 
15: 307 & 311. 1967. 

Recent collectors have found this plant growing on slopes. 
Tsang reports it as "fairly cooimon" and "abundant scattered 
shrubs" in Kwangsi, Norton refers to it as "common on open hill- 
sides" in Fukien, and Ching found it in "open thickets on stream 
banks" in Chekiang. 

The corollas are described as having been "lavender" on Chiao 
27OU & s.n. [Herb. Univ. Nanking U4O5U] and Koelz 1;592 , "bluish" 
on Rock 6981 and Tsiang & P'ei 5725, "bluish-purple" on R. £. 
Ching 2U29, "blue" on J. B. Norton 1558 and Tsang 27733 , "pink" 
on Tsang 278U3 , and ••white" on Tsui 303. 

Herbarium material has been identified and distributed under 
the epithet V. negunda L., in addition to the epithets previously 
recorded. On the other hand, the £. Wright s.n. [Hong Kong] and 
the Zlmmermann hh2 and Barchet 556, cited by P'ei or so identified 



20 PHYTOLOGIA Vol. 17, no. 1 

by him, seem to me to be better placed as var. cannabilolla (bieb. 
& Zucc.) Hand.-llazz. 

Additional citations: INDIA: East Punjab: Koelz kt>92 (W— 
I6679ii9). CHINA: Anhwei: Herb. Univ. Nanklng~1726 (W--13U$!>70) . 
Chekiang: Barchet s.n. (W— 597^86, W— 59759U) ; Chiao s.n. [Herb. 
Univ. Nanking liiO^li] (W— 11^26576), s.n. [Herb. Univ. flanking 
1U580] (W— 11^26962)} R. C. Ching 2U29 (W— 12U7250) ; A. N^ Steward 
s.n. [Herb. Univ. Nanking 2387] (W— 13115971) . P^OcLen: J. B^ Nor- 
ton 15$8 (W— 117273li) . Hupeh: E. H. Wilson 790 [7/07] (TT— 7771i;3), 
790 [12/07] (W— 77711i3), 2702 [6/07] (W— 777U70), 2702 [8/07] (W— 
777U70). Kiangsi: Chiao s.n. [Herb. Univ. Nanking 1877U] (W— 
155U01U). Kwangsi: Fung 21196 (W— I70U6II) ; W. T. Tsang 27733 
(W— 1757177), 2781i3 (W— 17^7^8), 28017 (W— 17^7132 ) . Kirangtung : 
C. 0. Levine s.n. [Herb. Canton Chr. Coll. 89U] (W— IO9I67U), s.n. 
[Herb. Canton Chr. Coll. ll;20] (W— 877508), s.n. [Herb. Canton 
Chr. Coll. 1585] (W— 877507), 3. n. [Herb. Canton Chr. Coll. 3Uh2] 
(W— 1270970) J Tsui 303 (W--175U587) . Kweichow: Tslang & P'ei 
5725 (W~1575153) . Shantung: Chiao 270U (W— 1553816, W--1595051) . 
Szechuan: Farges s.n. (W— 2li97126) . Yttnnan: J^ F. C. Rock 698I 
(W— -1212126) . Province xindetermined: Schoch U27 (7/--117U976) . 
CHINESE COASTAL ISLANDS: Honan: £. 0^ Levine s.n. [Herb. Canton 
Chr. Coll. 250] (W— 778606). THAILAND: Zimmemann 2 )W— 595002) . 
INDOCHINA: Tonkin: P^telot 1170 (¥—171701271 

VITEX ORINOCENSIS H.B.K., Nov. Gen. & Sp. PI., ed. folio, 2: 200. 
1817. 

Additional synonyiny: Vitex orinoccensis Humb. & Bonpl. apiid 
Steud., Nan. Bot., ed. 1, 688. 1821. Vitex orineceasis Ruber, 
in herb. 

Additional & emended bibliography: H.E.K., Nov. Gen. & Sp. 
PI., ed. folio, 2: 200 (1817) and ed. quart., 2: 2h7 * 1818; 
Steud., Nom. Bot., ed. 1, 888. 1821; Barnhart, Bull. Torrey Bot. 
Club 29: 590. 1902; Veillon, Revist. Forest. Venez. 5: 59. 1962; 
Uoldenke, Phytologia 15: 312—313. 1967. 

It shovild be noted that the H.B.K. reference dates given above 
have been authenticated by consultation of the work by Barnhart 
(1902) on this subject. 

VITEX ORINOCENSIS var. MULTIFLORA (Mq.) Huber 

Additional synoriyrny: Vitex orineceasis var, multiflore (Mig) 
Huber, in herb. 

Additional bibliography: Moldenke, Phytologia 15: 313. 1967. 

Breteler describes this plant as a tree, 13 m. tall, the trunk 
3^ cm. in diameter at breast height, branched from low down, the 
bark shallowly and finely fissured, brownish-gray, the leaflets 
papery, slightly glossy and medium-green above, paler and dull 
beneath, the corolla pale-purple (on his no. 3662 ), the fruit 
subglobose, glossy, smooth, black at maturity, and growing at 350 



1968 Moldenke, Notes on Vitex 21 

meters altitude. 

Additional citations: VENEZUEIA: Barinas: Breteler 3662 (W — 
21^65602), 3907 (W— 2U65856). 

VITEX OXrCUSPIS J. Q. Baker 

Additional bibliography: Moldenke, Phytologia 1^: 311i. 1967j 
Anon., Biol. Abstr. 19'- 390. 1968. 

VITEX OXrcUSPIS var. MOSSAMBICENSIS Moldenke 

Additional bibliography: Moldenke, Phytologia 1^: 311i — 315. 
1967 i Anon., Biol. Abstr. U9: 390. 1968, 

VITEX PARVIFLORA A, L. Juss. 

Qnended synonyB^r: Vitex leucoaylon Span, ex Miq., Fl. Ind. Bat. 
2: 863. 1856 [not V. leuco^lon Blanco, 1895, nor L., 1829, nor 
L. f., 1781, nor Naves, 1918, nor Roth, 1956, nor Roxb., I8lli, 
nor Schau., 1893, nor Wall., 18U7, nor Willd., 1832]. 

Additional bibliography: Pers., Sp. PI. 3: 360. 1819 j Steud., 
Norn. Bot., ed. 1, 888. 1821; Vidal, Phan, Cuming. Philip. 13U. 
I885j Moldenke, Phytologia 15: 316—317 & 320 (I967) and 16: 500 

& 501. 1968. 

It should be noted that the V. leucoscylon of Linnaeus the 
younger is a valid species, with the hcmonyms ascribed to Lin^ 
naeus the elder, to Wallich, and to Willdenow as synonyms, that 
of Blanco and of Naves is V. negimdo L., that ascribed to Schauer 
is V. glabra ta R, Br,, and that accredited to Roth and to Rox- 
burgh is V^ glabrata var. bombacifolia (Wall.) Moldenke. 

The corollas are described as having been "light-blue" on 
Seibert 1535 » This collector describes the plant as a tree, 6—8 
meters tall, with blue-black fruit in August, He states that it 
is cultivated along the riverbank at Farm No. 5, ALnirante, in 
the Changuinola District, by the United Fruit Company, in Panama, 
where it was originally introduced because "the wood is good for 
railroad ties". 

Vidal (1885) cites Cuming Uhh , 1365, & 18 30 for this species. 
Herbarium material has been misidentified and distributed as V. 
floridula Duchass. & Walp. 

Additional citations: CULTIVATED: Hawaiian Islands: Degener & 
Degener 30092 (Ms— U9581) . Panama: Seibert 1535 (E— 1570765) . 

VITEX PEDUNCULARIS Wall. 

Additional synoEyn^r: Vitex peduncularis "Wall, ex Schau." apud 
Deb, Bull. Bot. Surv. India 3: 315. 1961, 

Additional & emended bibliography: Watt, Diet. Econ. Prod, In- 
dia 6 ik)i 250. 1893; Gamble, Man. Ind. Timb., ed. 2, 5Ul. 1902; 
Prain, Bengal PI., ed. 1, 2: 832 & 833. 1903; Gamble, Fl. Presid. 
Madras 2: 1102 & UO3. 192li; Deb, Bull. Bot, Surv. India 3: 315. 
I96I; Prain, Bengal PI., ed. 2, 2: 621, 622. & 1012. I963; R. C. 
Ghosh in Lahiri, West Beng, Forests 197. 1964; Sen Sc Naskar, Bull. 
Bot, Surv. India 7: 60, 1965; Jain & De, Bull. Bot. Surv, India 



22 P H Y T L G I A Vol. 17, no. 1 

8: 217. 1966; Rao F^ Rabha, Bull. Dot. Surv. India 8: 301. I966; 
Moldenke, Phytologia 1^': 319—320. 1967 . 

Prain (I963) desciribea this species aa a tree, 20~IiO feet 
tall, and records it from Bihar, Chota Magpur, and Orissa. Deb 
(1961) says "leaflets densely covered with minute yellow glanda 
beneath, panicles axillary", and cites Ueebold ST39- R^o *£ Rabha 
(1966) record the species from Assam, while Jain ^ De (I966) tell 
us that in V/est Bengal it is known as "bhadu", the ripe fruits 
are eaten, the wood is used to make agricultural implonents, eind 
the leaves are eaten aa a vegetable in the treatment of ophthal- 
mia. Ghosh (I96U) encountered the species at 150 meters altitude 
in the foothills of West Bengal. It has been foxind in flower and 
frxiit in July. 

An additional vernacular name recorded for V. peduncularis is 
"kyelyo", while the name, "tin nok", previously recorded for it, 
is said to be applied also to V, limonifolia Wall in Thailand. 

Additional citations: INDIA: West Bengal: C^ B^ Clarke 11733c 
(W — 802339). BURMA: Upper Burma: Anno on s.n. [Herb. Bunra Forest 
School 93] (W— 17166U3) ; Prazer 7 (W~712906), 73 (W— 712957). 
THAILAND: Native collector A.33 [Herb. Roy. Forest Dept. 5883] 
(W— 2O6U8O6) . INDOCHINA: Cochinchina: Thorel IOO6 (W— 2U97093) . 

VITEX PEDUNCULARIS var. ROXBURGHIANA C. B. Clarke 

Additional bibliography: Roxb,, Fl, Ind.. ed, 2 [Carey], 3- 

72. 1832; Watt, Eoon. Prod. India 7: 25U. 1883; Watt, Diet. 

Econ. Prod. India 6 (U): 250. 1893; Gamble, Man. Ind. Tirab., ed. 

2, 5hl. 1902; Prain, Bengal PI., ed. 1, 2: 832 & 833 (1903) aixi 

ed, 2, 2: 621 «c 622. 1963; Moldenke, Phytologia 15: 320. I967. 
Prain (1963) records this variety from Bihar and from Chota 

Nagpur, and adds the vernacular name "marak' " , In his 1903 work 

he cites the Watt reference given above as "E. D. 5 J 17U". 

VITEX PHAEOTRICHA Mildbr. 

Additional bibliography: Moldenke, Ph(ytologia 15: 311i & 321 — 
322. 1967. 

VITEX PIERREI Craib 

Additional bibliography: Moldenke, Phytologia 15: 323. 1967. 

Additional citations: THAILAND: Mrs. D.J. Collins 706 (W — 
1700656) . 

VITEX PINNATA L., Sp. PI., ed. 1, 638. 1753 [not V. pinnata 
Lo\ir., 18U7, nor •T.our. ex Schau.", I963] . 
Additio n al & emended synonymy: Vitex negundo Noronha, Verh. 
Batav. Gen, 5, ed. 1, art. U: 86. 1790 [not V. negundo Curtis, 
1832, nor L., 1753, nor L. f., I966, nor Lour., 193li, nor Royle, 
1919, nor Willd., I9I8] . Vitex arborea Roxb., Hort. Beng. l^, 
hyponym. l8li;; Fl. Ind., ed. 2 [Carey], 3: 73. 1832 [not V. ar- 
borea Br^on, 1955, nor Brown, I8O6, nor Desf ., 18U7, nor Fischer, 
1829] . Pistacia vitex L. ex Watt, Diet. Eeon. Prod. India 6 (U): 



1968 Moldenke, Notes on Yitex 23 

2^0, in sjm, 1893, 7itex pubescens -var, genuina Hochr,, Candollea 

5: 191. 193ii. 

Additional & emended bibliography: J. F. GmeL, in L,, Syst. 
Nat., ed. 13, pr. 1, 2: 963 (1789) and pr. 2, 2: 963. 1796; Pers., 
3p. PI. 3: 360 & 361. I8l9i Steud., IJom. Bot., ed. 1, 888. 1821; 
Roxb., Fl. Ind., ed. 2 [Carey], 3'- 73~7ii. 1832; Gamble, Man. Ind. 
Timb., ed. 1, 297—298. 1881; Watt, Econ. Prod. India 7: 255. 
1883; ¥att. Diet. Econ. Prod. India 6 (U): 250. 1893; Gamble, Man. 
Ind. Timb., ed. 2, 5Ul. 1902; Prain, Beng. PI., ed. 1, 2: 832 k 
833. 1903; Gamble, Fl. Presid. Madras 2: 1101—1103. 192U; C. 
Coster, Ann. Jard. Bot, Buitenz. 38: pi. 6, fig. 2. 1928; Hochr,, 
Candollea 5: 191 — 192. 193U; M, R, Henderson, Ccnmon Malay, 
Wildfls. 39. 1961; Prain, Beng. PI., ed. 2, 2: 621, 622, & 1012. 
1963; Santapau & Wagh, Bull. Bot. Siirv, India 5*. 109. 1963; Dcmk, 
Trav. Lab. Mat. M5d. Pharm. Gal. Paris 50: 1 — 26U. 1965; Sen & 
llaskar. Bull. Bot. Surv. India 7: 60. 1965; M. S. Mani, Bull. 
Bot. Surv, India 7: HU. 1965; Anon., Ind. Bibliog. Bot, Trop, 3 
(2): 15. 1966; Moldenke, Phytologia 15: 323—325 (1967) and 16: 
U95. 1968; Moldenke, Biol, Abstr, U9: 851. 1968, 

Additional illustrations: G, Coster, Ann, Jard, Bot, Buitenz. 
38: pi. 6, fig. 2. 1928. 

It should be noted here that the V. negtuido of Linnaeus the 
elder is a valid species, with the homonyms accredited to Lin- 
laeus the younger, to Rqyle, and to Willdenow as synonyms, 
while the V. negundo ascribed to Curtis belongs in the synonymy 
of V. negunao var. heterophylla (Franch.) Rehd. and that ascribea 
to Loureiro is V. quinata (Lour.) F. N. V^ill. The V, pubescens 
ascribed to Heyne is a synonym of V^ altissima L, f , The V, 
arborea accredited to Brion is a sjoionym of V. beravienais var, 
acuminata Moldenke, that accredited to Brovm is V. heptaphylla 
A. L. Juss., while that ascribed to Desfontaines and to Fischer 
is V, negundo var. albiflora Moldenke. 

Santapau & Wagh (I963) feel that the name, V. pinna ta Lour., 
should always be written "V^ pinnata Lour, ex Schau.", but my 
contention has always been that such a double citation is de- 
sirably ONLI in a formal synonymy where complete bibliographic 
references are given. It is too cumbersome to give such a 
double credit citation on identification labels or in the text 
of a paper where it would be of little, if any, added value. 

Banternsuk dascribas V. pinnata as a "medium tree common in 
dry deciduous forests" in Thailand. The corollas are described 
as having been "purplish" on his no. 13. Mani (1965) reports a 
plant gall found on this species, made by Eriophyes cryptotrichus 
Nalepa. It is an epiptQrllous hemispheric venrucose pouch-gall 
0.5 — 5 nm, in diameter, and is his gall no. 29. 

The bibliographic reference "Gamble 772" is sometimes given in 
literature for this species, but has not as yet been located by 
me. 

The Burma Forest School 22, distributed as V_, pinnata , is ac- 
tually V. limonifolia Wall, 



2U P H Y T L G I A Vol. 17, no. 1 

Additional citations: INDIA: West Bengal: Helfar 132 (W— 
1669076). BUKliA: Tenaaserim: Gallatly 1012 (W— 26307BT. THAI- 
LAMD: Banternauk 13 [Herb. Hoy. Forest Dept. 2010] (W— 206U733); 
Hansen ^ Smitinand 12186 (Rf ) . 

VITEX PINNATA var. ALATA Uoldenke 

Additional bibliography: Uoldenke, P^^ytologia 1$: 32ii 4 325. 
1967; Moldenke, Biol. Abstr. U9: 851. 1968. 

Additional citations: IlffilA: Khasi States: Hooker & Thomson a . 
n. [Mont. Khasia] (■ff--2U97073) . 

VITEX PINNATA f . ANOMALA Moldenke 

Bibliography: Uoldenke, Phytologia h' 181;. 1953} Uoldenke, 
Biol. Abstr. 27: 2026. 1953; Uoldenke, Phytologia 6: 79—80. 
1957 J Uoldenke, R6s\an6 198 & U78. 1959. 

VITEX PINNATA var. PANTJAfiZNSIS (Hochr.) Uoldenke, comb. nov. 

Synonymy: Vitex pubescens var, pantjarensis Hochr., Candollea 
5: 191—192. 193li. 

Hochreutiner's originitl description of this taxon is as fol- 
lows: "Flores ochroleuci, calyx profundius dentatus, inflorescen- 
tia majus elongata thyrsoidea, folia 5-foliolata, sed ut in typo 
pubescentia et nervata. Java, Goenoeng Psintjar, 4 1*E. de Bui- 
tenzorg au pied de la montagne. formant de grsinds arbres especis 
dans la brousse et haute de ±8 m. alt. ca. 350 m., 17 septembre 
I90U, fleurs jaunStres (n. I8U6) . Conme on le voit, c'est line 
vari^tS tres distincte du tjnpe. D'aucuns y verront une espece 
sp^ciale. Toitefois, conme les specimens hindous du V. arborea 
RojdD. — considlr^s comme synonymes — ont le temps a autre h et 
peut-Stre 5 folioles, on peut consid4rer ce caractere coujae 
varietal . " 

VITEX POBEGUIM Aubr^v. 

This taxon has recently been shown to be conspecific with V. 
madienais Oliv. and should therefore be deleted from ny list 
of valid and accepted taxa. 

VITEX POOGEI Gtlrke 

Additional bibliography: Durand & Jacks., Ind. Kew. Suppl, 1, 
pr. 1, U57 (1906) and pr. 2, U57. 19^1; Uoldenke, Phytologia 6: 
80. 1957; Uoldenke, RSsuml lii3 & U78. 1959; Durand & Jacks., Ind. 
Kew. Suppl. 1, pr. 3, U57. 1959. 

VITEX POLYGAMA Cham. 

Additional bibliography: Bocq., Adansonia 3: [Rev. Verbenac.] 
253. 1863; Jacks, in Hook. f. & Jacks., Ind. Kew., pr. 1, 1: Ukl 
(1893) and 2: 1211i. 1895; Sampaio, Bol. Uus. Nac. Rio Jan. 13: 
258. 1937; Jacks, in Hook, f . & Jacks., Ind. Kew., pr. 2, 1: Ui7 
(I9U6) and 2: 121ii, 19U6j Le Cointe, Amaz. Bras. Ill Arv. fit 
Plant. Uteis, ed. 2, 292. 19U7; Angely, Fl. Paran. 7: 13. 1957; 
Uoldenke, Phytologia 6: 80 — 89. 1957; Jacks, in Hook. f. & Jacks., 



1968 Moldenke, Notes on Vitex 2$ 

Ind. Keir., pr. 3, 1: UU7 (I960) and 2: 1211i. I960; Moldenke, Phy- 
tologia 8: 75. 1961; Moldenke, R6sum6 Suppl. 12: 5. 196^. 

Recent collectors refer to this plant as a "touceira com di- 
versos caules, 2 m.", with red anthers and white pollen, growing 
in sandy soil, flowering in August, and known as "graiina". The 
corollas are described as "violet" on H. F. Martins 2U2 and as 
"violacea com tubo floral maisclaro" on Mattes & Mattos 8382 » 

A cotype collection, in fruit, Sellow 3.n» , deposited in the 
herbarium of the Botanisches Museum at Berlin, was photographed 
there by Macbride as his type photograph number 17565 (in part), 
but is now destroyed. 

According to Sampaio (1937), the name "maria preta", recorded 
for Vitex polygama , is also applied to Blanchetia heterotricha P. 
DC,, Cordia curassavica Roem, & Schult., Melanojylum bra\ina 
Schott, and Zollemia ilicifolia Vog, 

The Schwacke s,n, [Mani] , distributed as V, polygama , is actu- 
ally var, hirsuta Schau, 

Additional citations: BRAZIL: Guanabara: Alston & Lutz lli2 
(Ja--llli096, Ja); Hans s.n. [30-10-19U6] (Ja— U3757, Ja); B. Lutz 
919 (Ja — 29U89); H, Tj_ Martina 2U2 [Herb. Cent. Pesq. Florest. 
I0II8] (Ac); Rosa 59 (Ja— 523U2, Ja, Ja); N. Santos 5268 [23^2] 
(Ac, Ja), 5300 [237-2] (Ac, Ja), 5373 [2Uh-3] (Ja, Ja). Minas 
Gerais: A. Castellanos 25U21 [Herb. Cent. Pesq. Florest. U229] 
(Ac); Heringer 7257 (B) . Rio de Janeiro: Glaziou 386O (Ja — 
5959). SSo Paulo: Mattos Sc. Mattos 8382 (W— 2i;li5l91) . State un- 
determined: Heringer 359U (B); Sellow s.n. [Brasilia; fructifera; 
Macbride photos 17565, in part] (W— photo of cotype). 

VITEX POLIGAMA var. BAKERI Moldenke 

Additional & emended bibliography: Moldenke, Phytologia 6: 83 
& 86—87 (1957) and 8: 75. 1961. 

VITEX POLYGAMA var. DUSENII Moldenke 

Additional & emended bibliography: Jacks, in Hook, f . & Jacks., 
Ind. Kew., pr. 1, 2: 1211* (1895) and pr. 2, 2: 121U. 19U6; Angely, 
Fl. Paran. 7: 13. 1957; Moldenke, Phytologia 6: 83 & 87—88. 1957; 
Jacks, in Hook, f . & Jacks., Ind. Kew., pr, 3, 2: 121ii, I960; Mol- 
denke, Phytologia 8: 75 — 76. I96I; Moldenke, R5sum^ Suppl. 12: 5. 

1965. 

A specimen of G. Gardner 582, deposited in the herbarium of 

the Conservatoire et Jardin Botaniques at Geneva, was photographed 
there by Macbride as his type photograph number 21^703, but is not 
a type collection of ai^r sort. 

The original description of V. laciniosa by Turczaninow (I863) 
is as follows: " Vitex ( pyrostoma y laciniosa . V. tota pilis rufes- 
centibus tecta, ramis ccanpresso-tetragonis; foliis longe petiola- 
tis 5foliolatis, foliolis obovato-oblongis , basi longe attenuatis 
petiolulatis, apice obtusis macronulatis vel acutiusculis integer- 



26 PHYTOLOOIA Vol. 17, no. 1 

rinds aut subrejMincila inconapicue denticulatia, supra pilia ad- 
preasis acabria, aubtus praeaartim ad nenratlonea denaius piloaia 
cinereia; cymia axillaribua petiolo duplo brevioribua blfidis, 
cum llore aolitario in dichotomlaj bracteia linearibua riorea ex- 
cedentibus; calycia dentibua tubum aequantibua, tubo corollae 
parum brevioribua. Bahia, Kegel No 12319- V. polygama Cham, et 
Schl. huic valde aimilia, differt tonento, praesertim in tergo 
foliorum multo denaiore, atque corollis calycem duplo superanti- 
bus." 

Additional citations: BRAZIL: Gvianabara: A. Caatellanoa 2U027 
[Herb. Cent. Pesq. Florest. 286U] (Ac); H. F. Martins 337 [Kerb. 
Cent. Pesq. Florest. 2870] (Z). Rio de Janeiro: G. Gardner $82 
[Macbride photos 2U703] (N — photo, W— photo) . 

VI TEX POLYGAMA var. GUZIOVII Moldenke 

Additional bibliography: Moldenke, Phytologla 6: 82, 33, & 88. 
1957; Moldenke, R6sum5 111 5c li78. 195^9. 

VITEX POLYGAMA var. HirtSUTA Schau. 

Additional &; emended bibliography: Moldenke, Phytologla 6: 83 
& 87—89 (1957) and 8: 76. I96I. 

A cotype specimen, Sellow s.n, , deposited in the herbarium of 
the Botanisches Museum at Berlin, was photographed there by Mac- 
bride as his type photograph number 17565 (in part), but is now 
destroyed. 

The corolla is described as "blue" on Schwacke s .n. , and the 
plant has been found in anthesis in December. Material has been 
misidentified and distributed in herbaria as typical V. polygama 
Cham. 

Additional citations: BRAZIL: Espirito Santo: Sellow s.n. 
[Macbride photos 17565, in part] (W — photo of cotype) . Rio de 
Janeiro: Schwacke s.n. [Man4] (Ja — 5968). 

VITEX POLYGAMA var. WARMINGII Moldenke 

Additional bibliography: Moldenke, Phytologia 6: 83 & 89. 
1957 i Moldenke, R6sum5 112, 379, & U78. 1959. 

VITEX POQARA Corbishley 

Additional bibliography: A. V/. Hill, Ind. Kew. Suppl. 6: 219. 
1926; J. Hutchinson, Botanist in South. Afr. 29U. 19U6} Moldenke, 
Phytologia 8: 76. 196lj Watt & Ereyei^Brandwijk, Medic. & Poison. 
PI. S. Afr., ed. 2, 1055 & lU51i. 1962; C. A. Sm., Common Names S. 
Afr. PI. 2U3, 37U, U38, li39, U98, & 601. I966. 

Snith (1966) records the vernacular names "hardekool" , 
"poeraboom", "poerasbooa", "stinkbessie", "stinkbos", 
"stinkbossie", and "weelxiisbessie" for this species — the first 
of which is also applied to Cambretxim . He reports that the ripe 
drupes are black and have the offensive smell of bedbugs or 
"weeluls", but this does not deter the natives and Europeans 
from eating the fruit. Hutchinson (19U6) cites his no. 1877. 



1968 Moldenke, Notes on Vltex 27 

VITEX PSEUDOCHRYSOCARPA Pieper 

Additional synonyny: Vitex pseudo-chrysocarpa Pieper ex Wors- 
dell, Ind. Lend. Suppl. 2: 500. 19la. 

Additional bibliography: A. W. Hill, Ind. Kew. Suppl. 8: 2h9» 
1933; Moldenke, Phytologia 6: 90—91. 1957; Moldenke, R^sum^ 133, 
138, 1U3, 382, & U78. 1959; Ruber in Hutchinson & Dalz., Fl. W. 
Trop. Afr., ed. 2, 2: iJiS. 1963; Moldenke, Phytologia 15: 95* 
1967; Moldenke Risum4 Suppl. 15: 25. 1967. 

Huber (1963) reduces this species to V, chrysocarpa Planch., 
but fails to cite the type collection, which is probJibly Dalziel 
771 [not "Dabjiel" as stated in error previously], and, being a 
collection by one of the co-authors of the work in -which Huber 
was writing, should have been available to him for examination. 
He does, however, cite Barter 121 Ij , a collection also cited by 
Pieper, so therefore doubtless bases his opinion on this specimen. 

VITEX PSEUDOCUSPIDATA Mildbr. 

Additional bibliography: A. W. Hill, Ind. Kew. Suppl. 7: 252 
(1929) and 8: 2U9. 1933; Moldenke, Phytologia 6: 91. 1957; Mol- 
denke, R^sum^ 139 & U78. 1959. 

VITEX PSEUDOLEA Rusby 

Additional bibliography: A. W. Hill, Ind. Kew. Suppl. 8: 2U9. 
1933; Moldenke, Phytologia 6: 91—93. 1957; Moldenke, R5sum4 85, 
115, & U78. 1959; Soukup, Biota 5: 137. 196U; Moldenke, R6sum5 
Suppl. 15: 5. 1967. 

Fenreyra describes this plant as a tree, 10 — 12 m» tall, with 
"violet" corollas, known locally as "palo de perro", the wood 
being used for timber. 

Additional citations: PERU: San Martin: Ferreyra 14829 (W~ 
1998617). BOLIVIA: Cochabamba; R. F. Steinbach U6U (S) . EI 
Beni: 0. E. Ylhlte 767 (G — isotype) . 

VITEX PDBERULA J. G. Baker 

Additional bibliography: K. Schum. in Just, Bot, Jahresber. 
28 (1): U97 & U98. 1902; Thiselt.-Dyer, Ind. Kew. Suppl. 2: 19h, 
I90U: Moldenke, Phytologia 6: 93. 1957; Moldenke, RSsianS lii8, 383, 
& li78. 1959. 

VITEX PULCHRA Moldenke 

Bibliography: Moldenke. Phytologia 3: hk5—Ul6, 1951; Moldenke 
in Humbert, Fl. Madag. 17U: 76, 132—133, 135, & 273, fig. 21, 1 
& 2. 1956; Moldenke, Phytologia 6: 93 — 9ii. 1957; Moldenke, R^sum^ 
157 & U78. 1959; G. Taylor, Ind. Kew. Suppl. 12: l5l. 1959. 

Illustrations: Moldenke in Hxaabert, Fl, Madag. 17Us 135, fig. 
21, 1 & 2. 1956. 

VITEX PYRAMIDATA B. L. Robinson 

Additional synonymy: Virex pyramidata Robins, ex Moldenke, R4- 
sum6 Suppl. 6: 11, in syn, 1963. 



28 PHYTOLOGIA Vol. 17, no. 1 

Additional & omondod bibliographer: lAirarid L. Jacks., Ind. Kew. 
Suppl. 1, pr. 1, li57. 1906: P. C. Standi., Contrib. U. S. Hat. 
Herb. 23: 123^ & 1236. 192/4; H. D. Davis, Life L V/orka Prln^le 
lis, 28U, 668, & 669. 1936; Durand fi. Jacks., Ind. Kew. Suppl. 1, 
pr. 2, IxSl (I9UI) and pr. 3, U57. 1959; Holdenke, Phrtoloeia 8: 
76. 1961 ; Langnan, Select. Guide Lit. Flow. PI. Uez. $96 L 1010. 
I96I4; L'oldenke in Shrove L Wiggins, Veg. 4 Fl. Son. Etes. 2: 
1261—1262. 1961ii Moldenke, Phytologia 15 : 265. 1967. 

Recent collectors describe this plant as a troe, to 5 m. tall, 
with fragrant flowers, fruiting in November and December, known 
locally as "capulln" or "jupari", and ascending from 100 to lUOO 
meters altitude. It has been found growing in matorral with 
Byrsonima sp. or with B_. eras si folia and Curatella sp., in dis- 
turbed matorral, or in open woods and pastures, "in rocky soil in 
association with Bursera , Erythidna , etc., in regular abundance" 
in Morelos . Fedderoa reports it "coianon" on savannas with Bryso - 
nima and Curatella , as well as in cleared areas, in Nayarit. In 
the same state it is said by McVaugh to be "occasional" with Bro - 
simum, Platymiscium , and Sapium , or to be "abundant" in rocky 
disturbed woodlands. 

The corolla is described as "blue" on J_. Rzedowski 15267 , 
"lavender" on R. Q. Abbott li^l , and ''b right-purple" on R. McVaugh 
15223. In Shreve & Wiggins (I96U) the distribution of the spe- 
cies is given as "On rocky hillsides, prairies, and basaltic 
mesas, in arrqyos, and at edge of craters. Lower Sonoran to 
Tropical Zones, Sonora to Yucatdn. Einployed by the natives for 
food, fuel, and construction. "kVhen burned, the ash is blue." 

The G, F, Gaumer 607 , distributed as V, pyr ami data , is actu- 
ally the type collection of V, gaumeri Greenm., Arguelles s.n. 
[San Bernardo, 12 Agosto 1958] is V. mollis K.E.K., and Janzen 
s.n, [29 Kay I96I1.] is not verbenaceous , 

Additional citations: MEXICO: Guerrero: R. Q. Abbott lljl (Ip); 
Hinton 10002 (Rf ) , 10005 (Rf), 112l4i4 (Rf); Paray 1915 (Ip). Jal- 
isco: Herb. Univ. Kans. Mex. Exped. W.57 (W— 2088629); A. R. Mol- 
denke 1823 (Rf); Pringle 14;29 (Ms— 309U9~isotype); J. Rzedowski 
15267 (Du— 513631, Ip). Mdxico: Hinton U086 (Rf ) ; R. V. Moran 
10159 (Du— Ii98l5ii). Morelos: Cox & Guzmin MCC .63I (Ip). Nayar- 
it: Feddema 877 (Mi), 13l;3 (Miyr "2632 (Mi); R. McVaugh 15223 
(Mi), 19089 (Mi); J. Rzedowski ll;396 (Ip), 17861t (Ip, Mi). Sin- 
aloa: J_. Gonzalez Ortega 793 (Ip) . Sonora ; Arguelles s.n. [San 
Bernardo, 18 Octubre 1958] (Rf). 

VITEX QUINATA (Lour.) F. N. Will. 

Additional & emended synonymy: Vitex hetrophylla Roxb . apud 
Kawakami, List PI. Formos. 85, sphalm, I9IO. Vitex quinata 
(Lam.) F. N. Will, apud S. Sasaki, List PI. Formos. 353, sphalm. 
1928. Vitex negundo Lour, ex Crevost & P^telot, Bull. Econoni, 
Indo-chine 37: 129U, in syn. 193U [not V. negundo Curtis, 1832, 



1968 Moldenke, Notes on Vitex 29 

nor L., 1753, nor L. f ,, 1966, nor Noronha, 1790, nor Royle, 1919, 
nor Willd., 1918]. Yltex quinata Dop ex Fletcher, Kew Bull. Misc. 
Inf. 1938: U3U, in syn. 1938. Connutia quinata Lour, apud Li, 
Wood. Fl. Taiwan 83!;, in syn. I963. Vitex quinata Lour, ex Mol- 
denke, R6sum6 Suppl. 15 : 2$, in syn. 1967. Vitex quinaria (Lour.) 
F. N. Will., in herb. 

Additional & emended bibliography: Steud., Nom. Bot., ed. 1, 
228. 1821; Roxb., Fl. Ind., ed. 2 [Carey], 3: 75. 1832} Hook. & 
Am., Bot. Beech. Voy. 206, pi. UQ* I836} Bocq., Adansonia 3: 
[Rev. Verbenac.] 253. l863j Gamble, Kan. Ind. Timb., ed. 1, 296. 
1881; Jacks, in Hook, f . & Jacks., Ind. Kew., pr. 1, 1: 6I9 (1893) 
and 2: 1213 & 12lU. 1895; Gamble, Man. Ind. Timb., ed. 2, 539. 
1902; Prain, Beng. PI., ed. 1, 2: 832 & 833. 1903; Prain, Ind. 
Kew. Suppl. 3: I89. 1908; Kawakami, List PI. Formos. 85. 1910; 
Dunn & Tutcher, Kew Bull. Misc. Inf. Addit. Ser. 10: 20U, 1912; 
A. W. Hill, Ind. Kew. Suppl. 6: 219. 1926; S. Sasaki, Ust PI. 
Fomos. 353. 1928; Stapf, Ind. Lond. 6: li78 & Ii79. 1931; P'ei, 
Sinensia 2: 70 & 73. 1932; E. D. Merr., Comm. Lour. 33U. 1935; 
Backer, Tectona 29: 686. 1936; Kanehira, Form. Trees, rev. ed., 
652, fig. 608. 1936; Fletcher, Kew Bull. Misc. Inf. 1938: U32 & 
li3U. 1938; Jacks, in Hook, f . & Jacks., Ind. Kew., pr, 2, 1: 619 
(I9U6) and 2: 1213 & 1211; . I9I16; Neal, In Gard. Hawaii, ed. 1, 
61i3. I9U8; Anon., Kew Bull. Gen. Index 1929-1956, 8U & 293. 1959; 
Jacks, in Hook, f . & Jacks., Ind. Kew., pr. 3, 1: 619 (i960) and 
2: 1213 & 1211; . I96O; Moldenke, Phytologia 8: 76. I96I; Liu, H- 
lustr. Nat, & Introd. Lign. PI. Taiwan 2; 1230, pi. IO38. 1962; 
U, Wood. Fl. Taiwan 16, 832, 83U, & 973. 1963; Prain, Beng. Pi., 
ed, 2, 2: 621, 622, & 1012, 1963; Srinivasan & Agarwal, Bull, 
Bot, Surv. India 5s 68. 1963; Panlgrahi, Chowdhury, Ra3u, & Deka, 
Bull. Bot, Surv, India 6: 255. 19611; Smitinand, Govt. Sarawak 
Sympos. Ecol. Res. Humid Trop. Veg. lil & 1^3. 1965; Mukerjee, 
Bull. Bot. Surv. India 7= 135. 1965; Backer & Bakh., Fl. Java 2: 
606. 1965; Sen & Naskar, Bull. Bot. Surv. India 7: oO. 1965f 
Hatuslma, Mem. Fac. Agr. Kagoshima Univ. 5 (3): 16 & U7. I966; 
Moldenke, R^sum^ Suppl. 15: 8, 9, & 25. 1967; Moldenke, Phyto- 
logia 15: 210; & 307. 1978. 

Additional illustrations: Hook. Sc Am., Bot. Beech. Voy. pi. 
1*6. I8I1I; Kanehira, Form. Trees, rev. ed., fig. 6O8. 1936; Liu, 
Ill\istr. Nat. & Introd. Lign. PI, Taiwan 2: pi. IO38. 1962, 

It should be noted here that the V. negundo of Linnaeus the 
elder is a valid species, with the homonyms ascribed to Linnaeus 
the younger, to Royle, and to Willdenow as synonyms, while that 
accredited to Curtis is V, neg\indo var. heterophylla (Franch.) 
Rehd. and that ascribed to Noronha is V. pinna ta L. 

The Hooker & Amott reference given in the bibliography a- 
bove is scsanetime erroneously cited as "181A", but actually pages 
193 to 288 and plates UO to 59 in this work were issued in 18 36, 

Recent collectors describe this plant as a tree, 12 — 21 m, 
tall, the trunk 15 cm, to 2 m, in diameter, the bark grayish- 
brown, the immature fruit green or yellow, and the mature fruit 



30 PHYTOLOGIA Vol. 17, no. 1 

puri)li3h-black to black, growing in ravines, the edges of cleared 
ravines, mixed loreets, open moist wood-margina, anc; dry ground 
beside forests . 

The corollas are described as having been ••whdte" on Lau 153, 
Lei nh, and W. T_. Tsang 178 . "white-purplish" on Wang 337^2 , 
"blue" on Lau 16, "pink" on Taam 1532, and "yellow*' on W. T. Tsan^ 

719. 

Backer & Bakhuizen van den Brink (1965) describe the species 
as follows: "Leaflets 3 — 5, pellucid dotted (by the presence of 
cystoliths. In dried materials the upper surface of the leaves 
often shows a whether or not [=more or less?] circumvallate shal- 
low depression near each cystolith.), petioluled, oval-elliptic- 
obovate-oblong, mostly acuminate, herbaceous or thdnly coriaceous, 
pubescent on the neirves when young; median one 5 — 13 cm by 2 1/2 - 
6 cm, on a petiolule 1 1/2 — 3 cm long, the other ones smaller, 
on shorter petiolules; petiole 2 — 10 cm. Panicles terminal and 
often also in the upper leaf-axLls, 5 — 25 cm long; cymes 1/2 — 3 
cm (inclusive of 2 — 10 mn petiole); calyx 3 — U ma, with broad 
teeth; corolla sordidly violet; tube 5 — 7 nm, inside glabrous or 
(from the insertion of the stamens up to the base of the lower 
lip) with few to many hairs; filaments glabrous or basally spar- 
ingly hairy, shortly exserted; drupe subglobose, 3/U — 1 cm di- 

am. Tree not too dry forest." They certainly meant to 

say "peduncle" rather than "petiole" in their description of the 
length of the cymes. They include V_. sumatrana liiq. and V^ vel- 
utina "K. & V," in the synonymy of V, quinata . 

Hatusima (1966) gives the distribution of the species as "In- 
dia to S, China, Formosa, Malaysia". Srinivasan ^ Agarwal (I963) 
record it from West Bengal, Assam, and East Bengal. Panigrahi 
and his associates (I96U) refer to it as "abundant" in Orissa, 
but Hatusima (I966) tells us that it is "rare" on 5a tan Island. 
Mukerjee (1965) states that this tree "helps with smfficient 
moisture to convert a deciduous forest to evergreen". 

Vernacular names recorded for it include "five-leaved chaste- 
tree", "hu'kham", ••kaazlib", "ka wariba-nimzinboku", "kuburasi", 
"n8a-4", "6-ninjin-baku", "oo-nimzinboku", "o-tin", "patt'ttu", 
"po-kiU", "poorasu", "pvr-kiang" , and "soa-po-kiwn" . 

Li (1963) cites Faurie 1021 , A. Henry 1182 , 1182 A/3 , & 1182 
C, Kawai s.n. , Kawakami & Mori 7, Keng 1369, Makino s.n. , Uatuda 
359 & s.n. , Oldham 38U, Owatari s.n. , Suzuki 20503 , and E. H. 
Wilson 10019 & 11127 from Formosa, 

Material has been misidentified and distributed in herbaria 
as Ardisia sp . On the other hand, the R. £. Ching 5552 , 0_. Deg- 
ener IUU8I , Greenwood 3lUta , A. Henry 1182 & ll32c, Keng K.I369 , 
Liang 62780 , Peng , Tak , & Kin s.n. [Herb. Canton Chr. Coll. 
I26I3] , Pgtelot 963, A. C. Smith U307 & 6295 , and F. K. Ward 
37559, distributed as typical V, quinata , are actually var. ^- 
berula (H. J. Lam) Moldenke, while E. H. VJilson 11127 is the type 
collection of var. serrata Moldenke, Herb. Canton Chr. Coll. 



1968 Moldenke, Notes on Vitex i""- 

12882 and E. H. Wilson I^OS are V. canescens Kurz, £. Wright s ji. 
[Hong Kong] is V. negiindo var. cannabifolia (Sieb, & Zucc.) Hand,- 
Mazz,, and Clemens & Clsmens 339h is V, tripinnata (Lour.) Merr, 
Additional citations: CHINA: Chekiang: R. C. Ching 198? (W— 
13U68U6). Kwangsi: R. C_. Ching 7309 (W~12U8677) . Kwangtung: C. 
0. Levine s.n, [Herb. Canton Chr, Coll. 10] (W— 778511), s.n. 
[Herb. Canton Chr. Coll. 999] (W~779l62), s.n. [Herb. Canton Chr. 
Coll. 1206] (W--II7313I), s.n. [Herb. Canton Chr. Coll. 1807] (W— 
1U2869U), s.n. [Herb. Canton Chr. Coll. 1876] (W— 13U7890) ; Tsi- 
ang 1066 (W— 1513168); Ying 770 (W— 1513078). Yttnnan: A. Henry 
12638 (W~U59211) . HONGKONG: W. Y. Chun 5177 (Ws) j Taam 1532 (W- 
20638I9), 18U6 (W— 2072690). CHINESE COASTAL ISLANDS: Hainan: 
Chun & T3oT395i^ (Bi), lil;673 (W--l675U22)i Fung 20U20 (Mi)} F. C. 
How 70570 (Bi), 70858 (Bi)} How & Chun 7021^8 (W— I669U2U); Lau 16 
(W— 1629005), 15^ (W— I62922I); Lei 66 (W-.1753851), Vh (W— 
l65U279)i Liang 62069 (W~l6 70785); F^ A. McClure 786 [Herb. Ling- 
nan Univ. I8320] (W— I666U92); W. T. Tsang 178 [Herb. Ungnan U- 
niv. 15677] (W--1250000) , 223 [Herb. Lingnan Univ. 15722] (W— 
I2U98O9), 719 [Herb. Lingnan Univ. 17U68] (W— I672609), 868 [Herb. 
Lingnan Univ. 16 367] (W— 12U9U97), 9ldi [Herb. Lingnan Univ. 
I6l4li3] (vr— 12U9327); W^g 33752 (¥/— 1670257), 3U267 (W— 1670370). 
Honam: Herb. Canton Chr. Coll. I66 (¥■— 77856U) . IflESTERN PACIFIC 
ISUNDS: Mindanao: Wenzel 2523 (Mi), 2912 (Bi) . 

VITEX QUINATA var. PUBERULA (H.J. Lam) Moldenke 

Additional synonymy: Vitex mindanaenais Merr. ex Moldenke, R6- 
sum6 Suppl. k'' 21, in syn. I962. 

Additional bibliography: Thiselt .-EJyer, Ind. Kew. Suppl. 2: 
I9U. I90U; Maun, Philip. Joum, Forest. 16: IO8. I96O; Moldenke, 
Phyliologia 8: 77—78 (I96I) and 15: 2UU & 307. 1967. 

Merrill based his V. mindanaenais on an unnumbered collection 
made by B. Rafael and S. S. Ponce in Butuan Subprovince, Mindanao, 
Philippine Islands, in September or October, 1913 [ Herb. Philip . 
Forest Bur. 207U6] , deposited in the United States National Her- 
barium at Washington. 

Elmer describes the variety as a stocky tree, 25 feet tall, the 
trunk 12 inches thick, the wood moderately soft, whitish, soon 
discoloring to a dirty-white, odorless and tasteless, the bark 
thick, grayish-white, finely checked, the branches numerous above 
the middle, forming a dense elongated crown, the twigs ascending, 
greenish-brown, with elongated lighter-brown lenticels, the peti- 
oles green and ascending, the leaflets horizontally recurved, 
strongly conduplicate on the upper sublucid and darkeivgreen sur- 
face, thinly coriaceous, the inflorescence erect, greenish, 
slightly fragrant, the corolla creamy, the upper segment purp- 
lish-streaked, the filaments whitish, and the anthers purplish- 
brown. Keng describes the bark as pale-gray and furrowed. 



32 P II y T L G I A Vol. 17, no. 1 

The variety has been found groning on fertile soil of open 
grasslands at altitudes of 1^ to lOOO meters. Additional vernac- 
ular names for it are "shok wong king", "tai wong muk", and 
"topaa". 

The corollas are described as "violet" on F, K_. Ward 37$3'9 « 
Material has been misidentified and distributed in herbaria under 
the names V. glabrata R. Br., V, heterophyllum Roxb., _V. negundo 
L., V^ quinata (Lour.) F. N. Will., V^ pentaphylla Uerr., Teijs - 
manniodendron coriaceum (C. b. Clarke) Kosterm., and Araliaceae . 

Additional citations: CHINA: Kwangsi: R. C. Ching $$$2 (W— 
12U8671) . Kwangtung: Peng , Tak , ^ U£ s.n. [Herb. Canton Chr. 
Coll. 12613] (W— 12U8220). Yttnnan: Feng 13396 (A). CHUffiSE 
COASTAL ISLANDS: Hainan: Liang 62780 , in part (TF— 1670920) . 
THAILAND: Smitinand hS$9 W^ F. K. Ward 37^59 (S) . IinX)CHINA: 
Tonkin: P6telot 963 (Vf--1759227) . LIALAIA: Perak: Comer 3162$ 
(N). WESTERN PACIFIC ISLANDS: FORMOSA: A. Henry 1182 (W— 1^55592), 
1182c (W— U55593)i Keng K.1369 (W--2035969, W— 2035970) . PHILIP- 
PINE ISLANDS: Mindanao: Elmer 11602 (Bi, N)j RaTael L Ponce s.n. 
[Herb. Philip. Forest Bur. 207U6] (W— 900566) . INDONESIA: GREAT- 
ER SUNDA ISLANDS: Celebes: Laleno U9 [Boschproefat. B. b. 19iiUi4] 
(Bi)j Waturandang 6l9 [Boschproefst. Cel/V.335] (Bi) . Sarawak: 
M. Jacobs 5llit (W— 2377357). Sumatra: Yates I6O9 (Mi) . MELANE- 
SIA: YASAWA FIJI ISLANDS: Viti Leva: 0_. Degener llilt8l (Bi); J. W. 
Gillespie 2953 (Bi, Bi), IjlSh,! (Bi, Bi). ii691.8 (Bi); Greemrood 
3hka> (Bi) ; A. C. Smith U307 (Bi) , 6295 (Bi) . 

VITEX QUINATA var. SERRATA Moldenke, var. nov. 

Haec varietas a forma typica special foliolis grosse serratis 
recedit. 

This variety differs from the typical form of the species in 
having its leaflet-blades coarsely serrate along the margins a- 
bove the middle. 

The type of the variety was collected by Ernest Henry Wilson 
(no, 11127) in forests along the upper Pinan River, province of 
Pinan, Formosa, on November 17, 1918, and is deposited in the 
United States National Herbarium at Washington. The collector 
describes the plant as a tree, to 60 feet tall, with a spread of 
15 feet. The type is in full fruit, so the serrate character of 
the leaflets cannot be ascribed to the specimen being fron a 
watersprout, as might otherwise be said. It was originally dis- 
tributed as V. heterophylla Roxb. and annotated as V, qoinata 
(Lour.) F. N. Will by Hui-lin Li in 1951. 

Citations: WESTERN PACIFIC ISLANDS: FORMOSA: E. H. Wilson 
11127 (W— IO52UOO— type). 

VITEX RADUU Mildbr. 

Synonymy: Vitex robynsi DeWild., Plant. Bequaert. $: 13 — lii. 
1929. 

Additional & emended bibliography: DeWild,, Plant. Bequaert. 



1968 Moldenke, Notex on Vltex 33 

$: 13 — Hi. 1929 J Moldenke, Known Geogr. Distrib. Verbenac, ed. 
1^ Ii9~5l & lOU. 19l;2} H. N. & A. L. Moldenke, Plant Life 2: 79. 
I9U8; Moldenke. Known Geogr. Distrib, Verbenac, ed. 2, 11^, 117, 
120, & 202. 19U9; Moldenke, PhTtologia 6: IO8 & 116. 1957; Mol- 
denke, R6sum« 1U3, 1U5, liio, 150, & U78. 1959; Moldenke, Phyto- 
logia 8: 78. I96I; Moldenke, R^surafe Suppl. 13: 3, 5, & 7. 1966. 

The binomial, Vitex radula Mildbr., appears to have been pro- 
posed first as a hyponym on July 1, 1928, and validated on May 30, 
1929. The binomial, V. robynsl De^iD.d., was also validly publish- 
ed in 1929, but as yet I have not been able to ascertain the ex- 
act month or day. I am therefore tentatively reducing it to 
synonomy under V. radula , since the two taxa are apparently con- 
specific . The type of V. robynsi , as has been stated previously, 
was collected by my good friend. Prof, Dr. Frans Hubert Edouard 
Arthur Walter Robyns ( no. 1913) in a shrubby savanna at Kasenga, 
at an altitude of about 970 meters, in the Republic of the Congo, 
on April 8, 1926, and is deposited in the herbarium of the Jar- 
din Botanique de I'Etat at Brussels. 

Recent collectors describe V, radula as semi-climbing or as a 
shrub, 3.5 m, tall, growing in rather wet or sandy soil, on shrub- 
by or on secondary woody savannas, in deciduous forests, secon- 
dary evergreen forests, dense Brachystegia-Pterocarpus forests, 
or Brachystegia forests with groups of Oxytenanthera abyssinica in 
black sandy soil with granite boulders, at 8OO to 1100 meters al- 
titude, called "bebesuco" or ••linuna-nuna" , flowering in February, 
and f mi ting from February to April and in June. The corollas 
are described aa '•white" on Barbosa 1037 . 

The Torre 1268 , distributed as V. radula , is actually V. thyr - 
si flora J, G. Baker, 

Additional & emended citations: CONGO LEOPOLDVILLE : Robyns 
1913 (Br, Br, N— photo, Z~photo). ZAMBU: Bredo UOO8 (Br, N) , 
PORTUGUESE EAST AFRICA: Cabo Delgado: Torre & Paiva 1200^ (Ul) . 
Manica e Sofala: Andrada 10^9 (Ul) ; Barbosa 1037 (Ul, Z), 1^83 
(Ul) ; Torre U3U0 (Rf, Ul) . Niassa: Torre & Paiva 10732 (Ul), 
10951 (Ul, Z). 

VITEX RAPINI Beauvis, 

Quended synonymy: Vitex raplnll Beauvis, ex Moldenke, R^suml 
388, in syn. 1959; Guillaum., Mdm. Mus. Nat. Hist. Nat, Paris B. 
15: 315. 1967. 

Additional bibliography: Prain, Ind. Kew. Suppl. 3: 189. 1908 j 
Moldenke, R6sum6 206, 342, 388, & U78. 1959; Moldenke, Phytologia 
8: 78. 1961 J Guillaum,, Thome, & Virot, Univ. Iowa Stud. Nat, 
Hist, 20 (7;: U5. 1965; Guillaum,, M&n, Mus. Nat, Hist, Nat. Paris 
B.15: 315. 1967. 

Guillaumin, Thome, & Vitot (1965) cite Thorne 285U. from New 
Caledonia. Guillaumin (I967) states that the species grows in 
serpentine at 9OO meters altitude, and cites Baumann 823^ . 



3U PHYTOLOGIA Vol. 17, no. 1 

VITEX RAPINIOIDKS Gulllaum. 

Qnended synonymy: Vltex raplnoides Guillaum. ex A. \'l . liill, Ind. 

Kefw. Suppl. 9: 297. 1938. 

Additional L emended bibliography: A. V^ Hill, Ind. Kew, Suppl, 
9: 297. 1938; Uoldonko, Rfisum* 20^;, 388, £- U78, 1959; Moldenke, 
Phytologia 8: 78. I96I. 

Additional citations: UKUfffiSIA: lEV/ JIEbHIDES: Aneityum: J. P_^ 
Wilson 3.n. [Kajewski 992] (Bi— isotype) . Efate: Kajewoki m 
(Bi). Eromanga: Kajerwski 299 (Bi). 

VITEX REDNELLIANA Uoldenke 

Additional bibliography: E. J. Salisb., Ind. Kew. Suppl. 11: 
265. 1953; Moldenke, Phytologia 6: 83, 8U, 89, & 110—112. 1957; 
Moldenke, R6sum6 112 & U78. 1959. 

VITEX REHMANNI Gtlrke 

Qnended synonymy: Vitex rehmannii Gtlrke ex Moldenke, Alph. List 
Invalid Names 55, in syn. 19li2; J, Hutchinson, Botanist in South. 
Afr. 335. I9U6. 

Additional bibliography: Thiselt.-Dyer, Ind. Kew. Suppl. 2: I9U. 
I90I4; Watt & Breyer-ErandvrLjk, Med. &: Poison, PI. S. Afr., ed. 1, 
15U & 2la. 1932; J. Hutchinson, Botanist in South. Afr. 335. 1916; 
Moldenke, Phytologia 6: 2U. 1957; Moldenke, R^sum^ 15U, 388, 4 
U78. 1959; Moldenke, Phytologia 8: 78—79. 1961; Watt & Breyer- 
Brandwijk, Med. & Poison. PI. S. Afr., ed. 2, 1055 & lii5U. 1962; 
R. H. Canpton, Joum, S. Afr. Bot. Suppl. 6: 66. 1966. 

The corollas are described as "white" on Sidey I3IO. Hutchin- 
son (I9I46) cites his no. 211i8 , which, he says, had ♦•mauve" corol- 
las. Compton (1966) records the species from Swaziland. 

Additional citations: SOUTH AFRICA: Transvaal: Schlieben 7526 
(N); Sidgy 1310 (S) . 

VITEX REHMAIINI f . SUBTOMEIITOSA Moldenke 

Additional bibliography: Moldenke, Rlsrmi6 l5ii, 388, & I478. 
1959; Moldenke, Phytologia 8: 79. I96I. 

VITEX RESINIFERA Moldenke 

Bibliography: Moldenke. Phytologia 3: U^— liii7. 1951; Moldenke 
in Humbert, Fl. Madag. 17li: 72, 81—86, & 273, fig. H (U--6). 
1956; Moldenke, Phytologia 6: llli. 1957; Moldenke, R^sumfi 157 & 
li78. 1959; G. Taylor, Ind. Kew. Suppl. 12: l5l. 1959. 

Illustrations: Moldenke in Humbert, Fl. Madag, 17U: 85, fig. 
11 (U— 6). 1956. 

VITEX RIVULARIS Gtlrke 

Additional bibliography: Prain, Ind, Kew. Suppl, 3: I89. 1908; 
F. R. Irvine, PI. Gold Coast U38. 1930; Dalz., Useful PI. W. Trop. 
Afr. U58. 1937; Aubr^v., Fl. For. Cot, Iv., ed. 2, 3: 233, pl. 336, 
fig. 5—7. 19^9; F, R. Irvine, Woody PI, Ghana 76Ii. I96I; Moldenke, 
Phytologia 8: 79. I96I: Hviber in Hutchinson & Dalz., Fl, W. Trop, 
Afr,, ed, 2, 2: UU5 &. UU6, 1963; Moldenke, R5sm4 Suppl. 12: 7. 



1968 Moldenke, Notes on Vitex 35 

1965; lioldenke, Phjrtologla 1$: 2$\x & 2^6, 1967. 

Illustrations: Aubrlv,, Fl. For. Cot, Iv,, ed. 2, 3s pi. 336, 
fig. 5—7. 1959. 

Recent collectors and authors describe this plant as a forest 
tree, 30 — 75 feet tall, the stem with thin bark, often fluted, 
the bark -nhitish-green or broTm, fairly smooth, papery, longitud- 
inally furrowed, the wood soft or hard and white, the slash 
olive-brown, with brovmish longitudinal lines; leaves digitate; 
leaflets 5 — 7, elliptic-lamceolate, long-petiolulate, 15 cm. 
long, 5 cm, wide, entire, acuminate at the apex, pubescent be- 
neath, the secondaries 12 pairs; flowers small, numerous, white, 
lilac-tipped or tinged witBt purple, bluish in bud, borne in open, 
slender, irLch, long-pedunculate, wide-spreading cymes which are 
dichotomously branched; fruit edible, black, ellipsoid or obo- 
void, 1/2 inch long, borne in an enlarged cup-shaped frui ting- 
calyx. 

The species grows in deciduous forests, flowering in April and 
May, fruiting in Jxine, July, and October. The fruit is eaten by 
game. Additional vernacular names recorded for it are "ash", 
"akwakora gyahina", "(m)bli", "m'vassa", "nt^rowa", "^twe", and 
"otwe ntJroTra" . 

Huber (I963) states that the species occurs also in French 
Cameroun and the Congo. The Zenker s.n. [Bipindi], distributed 
as _V. rivularis , is actually V. longipetiolata Gtlrke. 

rluber (I963) cites the following collections: GHANA: F. R^ 
Irvine 951 ; Vigne FK.865 , 895 , ^ 1091 . LIBERIA.: Baldwin 6285 & 
6U9I. IVORY COAST: Chevalier 19097. SOUTHERN NIGERIA: Jones & 
Onochie FHI.I876O ; Kennedy 910 . BRITISH CAMEROONS: Mildbraed 
10535. 

Additional citations: CAMEROONS: Zenker 3761i (W— 55)4189). 
ANGOU: Cabinda: Monteiro & Murta 89 (Ul). 

VITEX ROBYNSI DeWild, 

This taxon is now regarded as conspecific with V, radula 
Mildbr, and should therefore be removed from my list of valid 
and accepted taxa, 

VITEX RUBRA Moldenke 

Bibliography: Moldenke, Phytologia 3: Uli7. 1951} Moldenke in 
Humbert, Fl. Madag. I7I4: 75, 115, 117—118, & 273, fig. 17 (6—8). 
1956; Moldenke, Phytologia 6: 116—117. 1957; Moldenke, R^sum^ 
157 & Ii78. 1959; G, Taylor, Ind. Kew. Suppl, 12: l5l. 1959. 

Illustrations: Moldenke in Humbert, Fl, Madag. 17U: 115, fig. 
17 (6—8), 1956. 

VITEX RUBRO-AURAHTIACA DeWild. 

Additional & emended bibliography: A, Iff, HUl, Ind, Kew, 
Suppl. 8: 2U9. 1933; Moldenke, R6sum6 lli3 <Sc li78. 1959; Moldenke, 
Phytologia 8: 79. 1961. 

Additional citations: CONGO LEOPOLDVILLE : Louis 518 (B), 5786 



36 PHYTOLOOIA Vol. 17, no. 1 

(B), 6171* (B). 

VITEX RUFESCENS A. L. Jusa. 

Additional bibliography: H.E,K., Nov. Gen. & Sp. PI., od. 
folio, 2: 200 (1817) and ed. quart., 2: 21*6. I8l6; Pers., Sp. 
PI. 3: 360. 1819; Steud., Nom. Bot., ed. 1, 888. I821j Barnhart, 
Pull. Torrey Bot. Club 29: 590. 1902: Jacks, in Hook, f . L 
Jacks., Ind. Ken., pr. 1, 2: 12lU (1895) and pr. 2. 2: 1211*. 
19U6; Hill & Salisb., Ind. Kenr. Suppl. 10: 2Ui. 19U7i MoLdenke, 
Phytologia 5: li30 (1956) and 6: 83. 1957; Moldenke, Rfisum^ 112, 
387, 389, & U78. 1959; Jacks, in Hook. f. 4 Jacks., Ind. Kew., 
pr. 3, 2: 1211i. I96O; k'oldenke, R^sumS Suppl. 12: 9. 1965; Mol- 
denke, Phytologia 15: 21*7. 1967. 

Tavares describes this species as a tree, about $ m. tall, 
with a tmmk diameter of 20 cm., known as "tamanqueiro", and 
fovind groning on the grounds of the Escola Agrononia do 'lordeste 
at Paraiba, /d.th the coiament "Cultavada? " . It was mis identified 
and specimens distributed as V. guerkeana Hiern. 

An isotype of V, perriana — Blanchet 3li3l; — in the herbarium 
of the Conservatoire et Jardin Botaniques at Geneva was photo- 
graphed there by Macbride as his type photograph nunber 30187, 
while the actual type, in the herbarium of the Musfium National 
d'Histoire Naturelle at Paris was photographed by him as his type 
photograph number 39501. 

It should be noted that the H.E,K, reference dates given above 
have been authenticated by consultation of the work by Barnhart 
(1902) on this subject. 

Additional citations: BRAZIL: Bahia: Blanchet 3i;3U [Macbride 
photos 30187 & 3950I] (W— photo, W— photo). Rio de Janeiro: H. 
F. Martins 209 [Herb. Cent. P«sq. Florest. 57U] (Z) . CULTIVATED: 
Brazil: Tavares 856 (W— 2U038IO) . 

VITEI RUFESCENS var, ABLUDENS (Moldenke) Moldenke 

Additional bibliography: Moldenke, Phytologia 6: 120 — 121, 
1957; Moldenke, Rfesum6 112, 388, & 1*78. 1959. 

VITEX SAMPSONI Hance 

Additional & emended bibliography: Jacks, in Hook, f, & 

Jacks., Ind. Kew., pr. 1, 2'- 1211*. 1895; Dunn & Tutcher, Kew 

Bull. Misc. Inf. Addit. Ser. 10: 201*. 1912; Hand.-Mazz,, Ann. 

Hort. Gothenb. 9: 68. 193U; Jacks, in Hook, f . & Jacks,, Ind. 

Kew., pr. 2, 2: 12ll*. I9U6; Moldenke, Risumfi 171 & 1*78. 1959; 

Jacks, in Hook. f. & Jacks., Ind, Kew., pr. 3, 2: 1211*. I96O; 

Moldenke, Phytologia 8: 79—80. I96I. 

The Levine collection, cited below, is marked "topotype". 
Additional citations: CHIM: Kwangtung: C O. Levine 3,n« 

[Herb. Canton Chr, Coll, 71*6] (¥—779018), 

VITEX SCABRA Wall, 

Additional & emended bibliography: Jacks, in Hook, f , & 
Jacks,, Ind, Kew., pr, 1, 2: 1211* (1895) and pr, 2, 2: lal*. 



1968 Moldenke, Notes on Vitex 37 

19li6; Moldenke, Phytologia 6: 122. 1957; Moldenke, R6svnn6 166 & 
1^78. 19^9} Jacks, in Hook. f. & Jacks., Ind. Kew., pr. 3, 2: 121it. 
I960. 

VTTEX SCANDENS Moldenke 

Bibliography: Moldenke, Phjrtologia U: 63~-6U (19^2) and 6: 122. 
1957; Moldenke, RSsumfi 202, 388, & U78. 1959; 0. Taylor, Ind. Ken. 
Suppl. 12: l5l. 1959; Moldenke, B&s\m6 Suppl. 12: 8. 1965. 

Clemens describes this plant as a scandent shrub, with flowers 
"dull brick purple with yellowish margin", growing at 2500 to 
1^500 feet altitude. 

Additional citations: MELANESIA: NEW GUINEA: Northeastern New 
Guinea: M. S. Clemens Ul775a (A) . 

VITEX SCHAUERIANA Moldenke 

Additional bibliography: Hill & Salisb.. Ind. Kew. Suppl. 10: 
2hh. 19U7; Moldenke. Phytologia 6: 123—12li. 1957; Moldenke, R5- 
sum^ 112, 386, & U7o. 1959. 

A cotype specimen of this species — Blanchet 2782 — in the 
herbarium of the Conservatoire et Jardin Botanicjues at Geneva, 
was photographed there by Macbride as his type photograph number 
30188, irtiile another of the same collection, deposited in the 
herbarium of the Naturhistorisches Museum at Vienna, is his type 
photograph nvanber 3U300. 

Additional citations: BRAZIL: Bahia: Blanchet 2782 [Macbride 
photos 30188 & 3li300] (W~photo of cotype) ; Fr6es 20182 (W— 
2li390U5) . 

VITEX SCHLIEBENI Moldenke 

Additional bibliography: Moldenke, R6sum6 Suppl. 1: 9. 1959; 
Moldenke, Biol. Abstr. 35: 1688. I96O; Moldenke, Phytologia 8: 
80. 1961 ; Hocking Excerpt. Bot. A.U: 592. 1962; G. Taylor, Ind. 
Kew. Suppl. 13: Uih. 1966. 

Additional citations: TANGANYIKA: Schlieben 6OO8 (N—isotype, 
W— 22lU711--isotype) . 

VITEX SCHOMBUEGKIANA Schau. 

Additional bibliography: Jacks, in Hook. f. & Jacks., Ind. 
Kew., pr. 1, 2: 1211; (1895), pr. 2, 2: 121it (19ii6), and pr. 3, 2: 
1211; . I960; Moldenke, Phytologia 8: 80. I96I. 

The type specimen of this species — J£i ^ Schcmbxirgk U21 — 
deposited in the herbarium of the Botanischer Garten und Museum at 
Berlin, was photographed there by Macbride as his type photograph 
number 17566, but is now destroyed. 

Additional citations: BRITISH GUIANA: M. R. Schomburgk U21 
[Macbride photos 17566] (W— photo of type) . 

VITEX SCHOMBURGKIANA var. GRANDIFLORA Moldenke 

Additional bibliography: Moldenke, Phytologia 6: 126. 1957; 
Moldenke, R6sum6 112 & 178. 1959. 



38 PIIYTOLOGIA Vol. 17, no. 1 

VITEX SEBESIAE H. J. Lam 

Additional bibliography: A. W. Hill, Ind. Kew, Suppl. 7: 252. 
1929; Moldenke, Phytologia 6: 126 — 128. 19^7; MoldonJ^e, R^sumft 
198 «c U78. 19!;9. 

VITEX SECUNDn-LORA H. Ilallier 

Additional bibliography: A. V/. Hill, Ind. Kew. Suppl. 6: 219. 
1926; Moldenke, Biol. Abstr. 32: 222 L 2353. 1?58; Moldenke, R6- 
sumfi 191 & U78. 1959; Moldenke, Phytologia 3: 80. I96I; Hocking, 
Excerpt. Bot. A. 5: U2. I962. 

VITEX SEINERI Gttrke 

Additional L emended bibliography: A. ^. HiU, Ind. lew, 
Suppl. 8: 21^9. 1933; Moldenke, Phytologia 6: 129. 1958; Uolaenke, 
R6suin6 lli8 L U78. 1959. 

VITEX SELLOWUNA Cham. 

Additional bibliography: Jacks, in Hook, f . 4: Jacks., Ind. 
Kew., pr. 1, 2: 1213 & 1211i (1895) and pr. 2. 2: 1ZL3 «c 121Ii. 
I9U6; Hill k Salisb., Ind. Kew. Suppl. 10: 2hk. 19ii7; Jacks, in 
Hook, f. «c Jacks., Ind. Kew., pr. 3, 2: 1213 & IZlU. I960; Mol- 
denke, Phytologia 8: 80. I96I. 

The type specimen of this species — Sellow lit37 — deposited 
in the herbarium of the Botanischer Garten und Museum at Berlin, 
was photographed there by Macbride as his type photograph number 
17567, but is now destroyed. 

Additional citations: BRAZIL: Rio Grande do Sul: Sellow ll;37 
[Macbride photos 17567] (W— photo of type). 

VITEX SERETI DeWild. 

Additional bibliography: Prain, Ind, Kew. Suppl. U, pr. 1, 21^8 
(1913) and pr. 2, 2U8. 1958; Moldenke, Phytologia 6: 132. 1958; 
Moldenke, R6sum6 lli3 & i;78. 1959. 

VITEX SIAMICA F, N, Will. 

Additional bibliography: Prain, Ind. Kew. Suppl. 3: 139. 1908; 
Fletcher, Kew Bull. Misc. Inf. 1938: U32 & U35. 1938; Anon., Kew 
Bull. Gen. Index 1929-1956, 293. 1959; Moldenke, Phytologia 8: 30, 
1961. 

Recent collectors have found this plant growing on limestone 
rock walls at sealevel, flowering in August, 

Additional citations: THAILAND: Larsen, Smitinand , & Warncke 
1238 (Ac), 

VITEX SIMPLIGIFOUJL Oliv. 

Additional synonymy: Vitex cordata AubrSv,, Fl. Forest. Sou- 
dano-Guin. 50lt. 1950. 

Additional & emended bibliography: Gtlrke in Engl., Pfl. Ost- 
Afr. C: 339. 1395; Jacks, in Hook, f , & Jacks,, Ind. Kew,, pr. 1, 
2: I21U. 1895; K. Schum. in Just, Bot. Jahresber, 28 (1): li97. 
1902; Thiselt.-Dyer, Ind, Kew, Suppl, 2: I9U. 1901;; J. H. Holland, 



1968 Moldenke, Notes on Vitex 39 

Kew Bull. Addit. Ser. 9 [Useful PI. Nigeria 3]: ^26. 1915; Lely, 
Useful Trees N. Nigeria ll6. 1925; Dalz., Useful PI. W. Trop. Afr. 
U57. 1937 j Aubrlv., Fl. Forest. Soudano-Guin. 50U. 1950; Moldenke, 
Phytologia 5: 305. 1955; Moldenke, R5sum6 133, 13U, 136~lIiO, lli3, 
381, 383, 389, & UTS. 1959; Jacks, in Hook. f. L Jacks., Ind. 
Kew., pr. 3, 2: 1211i. 1960; F. R. Irvine, ^Toody PI. Ghana 76U. 
I96I} Koldenke, Phjrtologia 8: 80. I96I; Jaeger & Winkoun, Bull. 
Inst. Franc. Afr. Noir 2li [ser. A, no. 1]: 79. 1962} Huber in 
Hutchinson 4 Dalz., Fl. W. Trop. Afr., ed. 2, 2: U;5 & hhl . 1963; 
Moldenke, Biol. Abstr. 1^8: 10099. 1967; Moldenke, Phytologia 15: 
100 & 232 (1967) and 16: U98. I968. 

Hiiber (1963) reduces V^ vogelii J. G. Baker to synonymy under 
7. simplicifolia . I regard it as a variety. 

Recent collectors and authors describe V. simplicifolia as a 
small tree or shrub, to 15 feet tall and with 1 foot girth, 
"often larger (?)", with dense pale indumentum; leaves 1- or 3- 
foliolate on the same tr«e, the leaflets 5 inches long, U 1/2 
inches wide, broadly elliptic, densely pubescent beneath when 
young, the secondaries 3 pairs, the petioles pubescent; flowers 
small; corolla greenish or mauve, tomentose, the lobes blue- 
purple 01* violet; fruit small, obovoid, violet-black, over 1/2 
inch long, 3-celled, on a hard saucer-shaped frui ting-calyx or 
"cupped like an acorn", with a thin edible pvilp, and a large 3- 
or U-seeded stone. 

The species is said to inhabit savannas, flowering frcm Janu- 
ary to June, fruiting in March and June. The twigs are used in 
Northern Nigeria to make "tooth sticks" or "chew sticks". In the 
Ivory Coast a lotion is made from the baric to use in the treat- 
ment of skin diseases and toothache. 

Additional vernacular names recorded for it are "abisa" and 
"nambara digali". The name, "bunmere", recorded previously for 
the fruit, is applied also to the fruit of Hannoa undulata . 

Huber (1963) cites the following collections: MALI: Aubr^ville 
1868 , Chevalier 2767, De Ganay 22. IVORI COAST: Aubr^ville 1^28 , 
I39U, 15U0, & 1967-1969 . DAHOMEY: Aubr^viUe U6d & 57d. NORTH- 
aiN NIGERIA: Barter I6U; , Dalziel 176 , Dent Young 206 , Lely 81;9 & 
P. 197 , Meikle 1070 , Trueblood FHI .1;319 . SOUTHERN NIGERIA: Barter 
1096. BRITISH CAMEROONS: Latilo & Daramola FHI.3Uii90 [this col- 
lection I regard as V, simplicifolia var. vogelii , vrtiich see] . 
He also comments "Also in Cameroun, Uganda and extending to Egypt 
and Sudan". Irvine (I96I) cites Brown 2236 , Kinloch 33U2, Kitson 
689 , and Vigne 3002 , 3777 , & 3786 from Ghana and says "Distribu- 
tion: Fr. Sudan to Cameroons and Sudan". 

VITEX SIMPLICIFOLIA var. VOGELII (J. G. Baker) Pieper 

Additional bibliography: K. Schinn. In Just, Bot. Jahresber. 28 

(1): U97. 1902; Thiselt.-Dyer, Ind. Kew. Suppl. 2: 19h» 190U; 

Moldenke, Phytologia 8: 80 — 81. I96I; Huber in Hutchinson & Dalz., 

Fl. W, Trop. Afr., ed. 2, 2: hhl . 1963. 

Huber (I963) reduces this taxon to synonymy under typical V. 



UO PHYTOLOGIA Vol. 17, no. 1 

almpllclfolla Oliv., but I am following Pieper in c^-vinc ^^ vari- 
etal status. The Latilo & Daramola 3ldi90 , which I havo previoua- 
ly cited as this variety, Huber cites under V, simplicif olia . 

VITEX SNETHLAGIANA Huber 

Bibliography: Moldeni:e, Known Geogr. Distrib, Verbenac., eci. 1, 
39 & lOU. 19li2; H. N. & A. L, Moldenke, Plant Life 2: 81. 19UBs 
Moldenke, Known Geogr, Distrib. Verbenac, ed. 2, 9$ ^ 202. 19u9l 
Moldenke, Phytologia 6: 136—137. 19^8; Moldenke, Rfisujnfi 112 & 
hlQ, 1959i G. Taylor, Ind. Kew, Suppl. 13: lUi. 1966. 

VITEX SPRUCEI Briq. 

Additional bibliography; Thiselt.-Dyer, Ind. Kew. Suppl. 2: 
19U. 190hi A. W. Hill, Ind. Kew. Suppl. 9: 298. 1938; Moldenke, 
Biol. Abstr. 33: 1215. 1959; Moldenke, Phytologia 8: 81. 1961; 
Hocking, Excerpt. Hot, A.5» Uk» 1962, 

The type specimen of this species — Spruce 2767 — deposited 
in the herbarium of the Conservatoire et Jardin Botaniques at 
Geneva, was photographed there by Macbride as his type photograjA 
n\Mber 2U705. 

Prance , Pena , Forero , Ramos , & Monteiro 3938 is said to have 
had its corollas "white with yellow center", and these collectors 
describe the plant as a tree, 22 m. tall, with a trunk diameter 
to U5 cm. 

The Murga Pires 781 , distributed as V. sprue ei , is not verben- 
aceous; it is probably something in the Bignoniaceae . 

Additional citations: BRAZIL: AmazBnas; Ducke ^l"(W — 1693056, 
W--187528U) ; Fr6e3 20510 (W— 2U39073); Prance , Pena , Forero , Ra- 
mos, & Monteiro 3938 (K, Rf); Spruce 2767 [Macbride photos 2ii765] 
(W — photo of type) . 

VITEX SPRUCEI var, LONGIDENTATA (Moldenke) Moldenke 

Additional bibliography: Moldenke, Phytologia 6: 139 — lUO. 

1958; Moldenke, R6sum6 112, 389, & U78. 1959. 

Additional citations: BRAZIL: AmazSnas: Fr6es 21398 (W— 

2li396l3) . 

VITEX SPRUCEI var. VAUPESENSIS Moldenke 

Additional bibliography: Moldenke, Biol. Abstr. 33: 1215. 
1959; Moldenke, Phytologia 8* 81. 1961; Hocking, Excerpt, Bot. 
A.5: hh, 1962. 

VITEX STAHELII Moldenke 

Additional bibliography: E. J. SsJj-sb., Ind. Kew. Suppl. 11: 
265. 1953; Moldenke, Phytologia 8: 81. 1961. 

Eerti describes this plant as "Arbol de 28 m. de altura total 
X 102 cm., yema terminal: complanada contomo m^s o menos c6nico. 
Ramitas terminales, verdosas con lenticelas alargadas y cremosas. 
Fruto: color morado negnizco. Semilla 1, envuelta en una pulpa 
cremosa, camosa" . It has been found in flower and fruit in May, 



1968 Moldenke, Notes on Vitex lO- 

Additional citations: VENEZUELA: Bolivar: E. L. Little 17659 
(Ve). Delta Amacuro: Berti ll;3 (N, S, Z), 163 (Ac, N) ; Wvirdack & 
Monachlno 3961i8 (N) . 

7ITEX STELLATA Moldenke 

Bibliography: Moldenke, Phytologia 3'- hhQ» 19^1; Moldenke in 
Humbert, Fl. Madag. 17U: 76, 12^, 126, & 273, fig. 19 (U--6). 
1956; Uoldanke, Phytologia 6: lli2~Ui3, 19^8; G. Taylor, Ind. 
Kew. Suppl. 12: 1^1 . 19^9; Moldenke, R^sumfi 1^7 & U78. 1959. 

Illustrations: Moldenke in Humbert, Fl. Madag. 17U: 125, fig. 
19 (U— 6). 1956. 

VITEX STRICKERI Vatke & Hildebr. 

Additional bibliography: Jacks, in Hook, f . & Jacks., Ind. 
Kew., pr. 1, 2: 12lU (1895), pr. 2, 2: 1211; (19U6) , and pr. 3, 2: 
121]i. I960; Moldenke, Phytologia 8: 81—32. 1961, 

Recent collectors describe this plant as a much-branched or 
scrambling shrub, to U or 5 feet tall, with rough bark, colorless 
sap, and panicles of aromatic flowers, the calyx brownish-green, 
filaments cream, and anthers brown, growing in groups in thickets 
on red-brown loam, the margins of thickets in Brachystegia wood- 
lands, or very local in Acalypha fruticosa - Acacia - Grot on - 
Haplocoelum - Grewia slmilis open to closed brushland on shallow 
black cotton soil ^vith lava rock pavements, to 2000 meters alti- 
tude, flowering in February. The corolla is said to have been 
"white" on Tanner 3J42O , "white tubular" on Greenway 9175 , and 
"cream" on Drummond & Hemsley I8IO . 

Additional citations: UGANDA: Meams 280 (W— 630295) . TAICAN- 
yiKA: Drummond & Hemsley I8IO (B); Tanner 2383 (B), 3U20 (S) . 
KENIA: Greenway 9175 (B) , 

VITEX STYLOSA Dop 

Additional & emended bibliography: A, W, Hill, Ind. Kew. Suppl. 
9: 298. 1938; Moldenke, Phytologia 6: 2h3—lUh» 1958; Moldenke, 
R63um6 177 & U78, 1959. 

VITEX SUMATRANA Miq. 

Additional bibliography: Jacks, in Hook, f . & Jacks., Ind, Kew,, 
pr. 1, 2: 121U (1895), pr. 2, 2: 1211; (19U6), and pr, 3, 2: 12lU. 
I96O; Moldenke, Phytologia 8: 82, I96I. 

VITEX SVrrNNERTONII S. Moore 

Additional bibliography: Prain, Ind. Kew. Suppl. 5, pr. 1, 273 
(1921) and pr. 2, 273. I960; Moldenke, Phytologia 8: 82, I96I. 

VITEX TANGENSIS GUrke 

Additional & emended bibliography: Gttrice in Engl., Pflanzerar. 
Ost-Afr, C: 339— 3l;0. 1895; K, Schum, in Just, Bot. Jahresber. 28 
(1): U97. 1902; This elt .-Dyer, Ind. Kew. Suppl. 2: 19li. 190U; Du- 
rand & Jacks,, Ind. Kew, Suppl. 1, pr, 1, U57 (1906), pr, 2, 1;57 



U2 P H y T L G I A Vol. 17, no. 1 

(19la), and pr. 3, 157. 1959; Uoldenke, Phytologia 8: 82. 1961; 
Cuf., Bull. Jard. Bot. Brux. 32: Suppl. 797-— 793. 1962; Uoldenke, 
Phytologia 1$: 315 (1967) and 16: U96. 1968, 

Recent collectors describe this plant as a shrub, 1.5 — h m. 
tall, several tines or' much branched from the base or near the 
base, many-stenmed, the bark pale gray-yellow, very finely retic- 
ulate. The corolla is described as "blue" on F. A. I'endonca 
2705 , "blue-lilac" on Dalslnhas 21; 2 , "violet" on Torre 958, "low- 
er lip violet" on Torre 2277, "cor de nalva" on Junod hlh , and 
"corolla-tube purple-mauve, large petal nauve, with yellow around 
the throat, throat purplish-mauve, the other petals white, fila- 
ments pale-mauve" on PolMll L Paulo 723. 

The species has been foxind growing in deciduous forests, in 
the substratum in dense forests, and in dune forests with Afzelia 
quanzensis , Dalium schlechterl , Garcinia livings tonei , Strychmos 
sp., etc. It is said to be common in the bush around cultivated 
land, with Adansonia , Allophylus , Carissa , Grewia , Hoslundia , 
Lannea , Sterculia , Stiychnos , Thespesia , etc., flowering in No- 
vember and December. 

Material has been misidentified and distributed in herbaria 
as V, amboniensis Gttrke. Torre 6323 is a mixture with V. oxycus - 
pis var. mossambicensls Moldenke. Torre 2277 is said to match 
well L. E. Godd 5U3U in the British Museimi and Kew hertarla, 
while F. A, Kendonca 2365 and Torre 3829 are said to match Volkens 
92, the type of the species, at the British Museum. Torre 3829 
is, however, described by Garcia as "intermediate" between 7. am- 
boniensis and V. tangensis. The A. Peter 39696, previously cited 
by me as deposited in ny personal herbarium, is now in the her- 
barium of the Texas Research Foundation at Renner, Texas. 

Additional citations: KENYA: Polhill L Paulo 723 (S) . PORTU- 
GUESE EAST AFRICA: Inhambane: Torre 3829 (Ul) . Lourengo llarques: 
Balsinhas 2U2 (Ul); Junod Ulli JmTi Torre 2067 (Ul, Z), 2277 
(Ul). Manica e Sofala: F. A. Mendonca 2365 (Ul), 2705 (Ul); 
Slmao 220 (Ul); Torre 6323 , in part (Rf, Ul, Ul) . llozambique: 
Torre 958 (Ul) . 

VITEX TELORAVINA J. G. Baker 

Einended synonymy: Vitex teleravlna J, G. Eaker apud Durand &: 
Jacks., Ind. Kew. Suppl. 1, pr. 1, ii57, sphalm. 1906. 

Additional &. emended bibliography: Durand L Jacks., Ind. Kew. 
Suppl. 1, pr. 1, 1'57 (1906) and pr. 2, it57. 19iil; Moldenke in 
Humbert, Fl. Madag. 17U: 77, 139, lU--lii2, & 273, fig. 22 (8 & 
9). 1956; Moldenke, Phytologia 6: 1U7— li;8. 1958; Durand & Ja;ks., 
Ind. Kew. Suppl. 1, pr. 3, U57. 1959; Moldenke, R^sumS 157, 389, 
& 178. 1959. 

Illustrations: Moldenke in Humbert, Fl. Madag. 17U: 139, fig. 
22 (8 & 9). 1956. 



1968 Moldenke, Notes on Vltex U3 

VITEX THOMASI DeWild. 

Additional & emended bibliography: A. W, Hill, Ind. Kew, 
Suppl. 8: 2ii9. 1933i Moldenke, Phytologia 6: lit8--ll(9. 1958; Mol- 
denke, R6sum6 ll;3 &: li78. 1959. 

VITEX THOMASI f . KASAIENSIS DeWild. 

Bibliography: DeVaid., Contrib. Etud. Fl. Katanga Suppl. 2: 
108—109. 1929; Moldenke, Phytologia 6: lli9— 150. 1958; Moldenke, 
R6sum6 ll;3 & UlQ. 1959. 

VITEX THONNERI DeV/ild. 

Additional & emended bibliography: A. W. Hill, Ind. Kew, Suppl. 
6: 219 (1926) and 8: 2i;9. 1933; Moldenke, Phytologia 6: 150. 1958; 
Moldenke, R^sum6 ll^O, lii3, & U78. 1959. 

VITEX THONNERI var. TIBATENSIS (Engl.) Pieper 

Additional bibliography: A. W. Hill, Ind. Kew. Suppl. 8: 2li9. 
1933; Moldenke, Phytologia 6: 151. 1958; Moldenke, R6sum6 139, 
389, & U78. 1959. 

VITEX THORELII Dop 

Additional & emended bibliography: A.W. Hill, Ind. Kew, Suppl. 
9: 298. 1938; Moldenke, Phytologia 6: 151—152. 1958; Moldenke, 
R^sumS 177 & U78. 1959. 

VITEX THYRSIFLORA J. G. Baker 

Additional bibliography: Durand & Jacks., Ind. Kew. Suppl. 1, 
pr. 1, U57. 1906; Prain, Ind. Kew. Suppl. 3: I89. 1908; I. Bailey, 
Ecology 1: 17U~189. 1920; Bequaert, Bull. Am. Mus. Nat. Hist. U5: 
333—383. 1922; A, V^, Hill, Ind, Kew. Suppl. 6: 219 (1926) and 9: 
297. 1938; Durand & Jacks., Ind. Kew. Suppl. 1, pr. 2, ii57. 19Ul; 
Uphof, Bot. Rev. 8: 569—571. 19U2; Durand & Jacks., Ind. Kew. 
Suppl. 1, pr. 3, U57. 1959; Moldenke, Phytologia 8: 82. I96I; 
Huber in Hutchinson & Dalz., Fl. \i, Trop. Afr., ed. 2, 2: hh$ & 
UU6. 1963; Moldenke, Phytologia 15: 312. I967. 

Recent collectors and authors describe this plant as an vmder- 
shrub, shrub, or small tree, 2m. tall, or a sarmentose shrub to 
6m. tall, with glabrous branches, 5-foliolate leaves, and small 
whdte flowers in terminal panicles, growing in forests or open 
Brachystegia forests, the herbaceous layer being dominated by Di~ 
gitaria and Panicum , at 280 meters altitude, flowering from Janu- 
ary to July and in September, fruiting from July to October, and 
csdled "namep^prlr" . The corolla is described as "yellow" on 
Torre 1268 . 

Bailey (1920) found this species inhabited by the ant, Vitici- 
cola tessmanni . The plant has lateral cavities or pits excavated 
in the woody parts of the stele of stout dry stems and branches. 
Furthermore, there are in stout stems exit-holes resembling those 
of the lateral pits subtended by them. This may be due to an in- 
herent tendency to form hollow stems and branches. It is not 
known whether the ants accelerate formation of the cavities 



Ui PHYTOLOGIA Vol. 17, no. 1 

throughout the center as has been demonstrated by Fiebrlg with 
Cecropla . The pseudo-gall-like structures made i/ Viticicola are 
histologically very complex. The insects enter throughi thie cir- 
cular apertures in the swollen internodes. According to Dequaert 
(19?2) this species of host has heteroplasias similar to thiose of 
Plectronia laurcntii. 

Dan Janzen, in a memorandum to my son, Andrew R. Uoldenke, re- 
fers to the original description of V. staudtii Gtlrfce as stating 
that the leaflets are glabix)U3 on the under surface but densely 
covered with minute, golden-yellow glands. He continues "These 
glands are critical (if they are indeed glands) to understanding 
the mymecophytic relationship that Viticicola has with Vitex 
staudtii ." He speaks of discussions of this plant and its myrme- 
cophily by Bequaert and by Bailey in VHieeler's "Ants of the Bel- 
gian Congo" . 

Material of V_. th^^Tsiflora has been misidentified and distrib- 
uted in herbaria as V. radula Mildbr. 

Huber (I963) cites the following collections: GUINEA: Baldwin 
9669 ; A. Chevalier 13199 & 13267 ; Jacques-Felix 8$2 . SIERKA LE- 
ONE: Delghton 31h7 i N. W. Thomas 1692 & 1953. LIBEPJA: J. T. 
Baldwin 6172, 9$10 , & 99hS i Harley s.n. [Ganta] ; Konneb 175. 
IVORY COAST: A. Chevalier 17055 , 193UO , & I9805. TOGO: Baimann 
56U . NORTHERN NIGERIA: Killick 67. SO'JTHERN NIGERIA: Harrison 
5; Olorunfemi FHI. 38057 ; Rowland s.n. [W. Lagos]; Symington FHI . 
5052 ; Talbot s.n. [Oban]. BRITISH CAMEROONS: Maitland 565 & 
1577; Olorunfemi FHI.306O8 ; Ujor FHI. 29283 . He ccmiments "Extends 
to Congo", 

Additional citations: SIERRA LEONE: N, W. Thomag 1692 (S) . 
PORTUGUESE EAST AFRICA: Mozambique: Torre 1268 (Ul); Torre & Pai- 
va 986U (Ul) . 

VITEX THYRSIFLORA var. UXIFLORA Pieper 

Additional bibliography: A. W. Hill, Ind. Kew. Suppl. 7: 252. 
1929; Moldenke Phytologia 6: 15>-15U. 1958; Moldenke, R5sum6 
139, 386, & U78. 1959. 

VITEX TOMENTULOSA Moldenke 

Additional bibliography: A. W. Hill, Ind. Kew. Suppl. 9: 298. 
1938; Anon., U. S. Dept. Agr. Bot. Sub j . Index 15: lli362. 1959; 
Moldenke, Phytologia 8: 83. I96I. 

VITEX TRICHANTHA J. G. Baker 

Additional & emended bibliography: Jacks, in Hook, f, & Jacks., 
Ind. Kew., pr. 1, 2: 1211; (1895) and pr. 2, 2: I21I4. 19U6; Mol- 
denke in Humbert, Fl. Madag. I7U: 75, 120—122, & 273, fig. 18 
(U— 6). 1956; Moldenke, Phytologia 6: 155—156. 1958; Jacks, in 
Hook, f . & Jacks., Ind. Kew., pr. 3, 2: 1211i. I96O. 

Qnended illustrations: Moldenke in Humbert, Fl. Madag. I7U: 
121, fig. 18 (U~6). 1956. 



1968 Moldenke, Notes on Vitex kS 

VITEX TRIFLORA Vahl 

Additional & emended synonymy: Vitex sericea Poepp. ex Btting- 
sh., Blatt-Skel. Dikot. 79, pl. 32, fig. 6. 1861 [not Y. sericea 
Poepp, ex Moldenke, 1936]. Pyrostoma tematum G. F, W, Mey. apud 
Jacks, in Hook. f. & Jacks., Ind. Kew., pr, 1, 2: 667. 189^. 
Vitex triflora tenuifoliai Huber ex Stapf, Ind. Lond. 6: U79. 1931. 
Vitex trifolia Vahl ex Moldenke, Suppl. List Invalid Names 11, in 
syn. I9UI [not V. trifolia Graham, I966, nor Hemsl., 19li9, nor 
L., 1753, nor L. f ., 1895, nor Moon, 1895, nor Sess^ & Moc, I9U0, 
nor "sensu Matsumiira & Hayata" , I963] . 

Additional & emended bibliography: H.B,K., Nov, Gen. & Sp. PI., 
ed. folio, 2: 200 (I8I7) and ed. quart., 2: 2U6. l8l8j Pers., Sp. 
PI. 3: 360. 1819} Steud., Nom. Bot., ed. 1, 888, 1821; Jacks, in 
Hook, f . & Jacks., Ind. Kew., pr. 1, 1: Ui;7 (1893) and 2: 667 & 
1211i. 1895; Bamhart, Bull. Torrey Bot. Club 29: 590. 1902; Le 
Cointe, Amaz. Bras, in Arv. & Plant. Uteis, ed. 1. U30. I93I4; 
Jacks, in Hook. f. & Jacks., Ind. Kew., pr. 2, 1: iili7 (19l*6) and 
2: 667 & 121ii. 19li6; Le Cointe, Amaz. Bras. Ill Arv. & Plant Ut- 
eis, ed. 2, U57. 19U7; Hill & Salisb., Ind. Kew. Suppl. 10: 2hh. 
19U7; Jacks, in Hook. f. & Jacks., Ind. Kew., pr. 3, 1: UIi7 
(i960) and 2: 667 & 1211i. 1960; Moldenke, Phytologia 8: 83. 1961; 
Soukup, Biota 5: 137. 1961^; Moldenke, R6sum6 Suppl. 15: 25. 1967; 
Moldenke, Phytologia 15 : 229, 2ii2, & 267 (1967) and 16: h9^* 
1968. 

It should be noted here that the V. trifolia of Linnaeus the 
elder is a valid species, with the homonym ascribed to Linnaeus 
the younger as a synonym, while the V. trifolia accredited to 
Graham is a synonym of V. negundo L., that ascribed to Hemsley 
and to "sensu Matsumura & Hayata" is V. trifolia var, simplicl - 
folia Cham., that ascribed to Moon is V. altissima L. f ., and 
that accredited to Sess6 & Mocifio is V, mollis H.B.K, 

LeCointe (19U7) records the vernactilar variant "tanand da 
mata" and conments "Nas capoeiras e mata secundlida. — E' a es- 
p6cie mais vulgar de AmazSnia.. ..0 fruto 6 emeniigogo e diuret- 
ico; as f8lhas enqDregam-se contra as cistites e uretrites; a 
raiz 6 tSnica e febrifuga." 

The Huber (1909) reference in the bibliography of this spe- 
cies is dated "1907-8" by Stapf (1931), but I9O9 seems to be the 
actual date vrtien the pages in question appeared. It should be 
noted that the H.B.K. reference dates given above have been au- 
thenticated by consultation of the work by Barnhart (1902) on 
this subject. 

The type specimen — Herb. Vahl s.n. — deposited in the her- 
barium of the Universitetets Botaniske Museum at Copenhagen, was 
photographed there by Macbride and is his type photograph nxnnber 
22779. 

The corollas on Mur<;a P^res & Cavalcante 52602 are described 
as having been "purple", the flowers slightly fragrant, and the 
plant itself "rar«". 

The Archer 80U7 and Barbosa de Silva 155, distributed as the 



Il6 PHYTOLOGIA Vol. 17, no. 1 

typical form of V. tri flora , are actually var. coilacea Huber, 

Additional citations: VENEZUELA: Amazonas: Ll^ Wiiliama 1$688 
(Ve— 8096, W— 2li2882]i) . BRAZIL: AmapA: Irwin , Lfur^a Pires, ^ 
Westra U83II (N); Mur<;a Fires U8$60 (Ui, N) ; l^rga. Pires ;^ Caval - 
cante 52602 (N, Rf)j Mart; a Pires , Kodrigues, «f Irvine ^OUSC (IJ) . 
Amaz3nas: Krukoff U70U (W— 166271?), 6869 (W— 1660920) . Pari: 
Fr6e3 20381 (vr— 2U390U2); Killip ^ Smith 30598 (V—lUShlBh) ; Ifur- 
ga Pires $1907 (N) . LOCALITY OF COLLECTIOli Ul.TJETEIda NED : Kerb. 
Vahl s.n. [ex India; Herb. Willdencnr 11 701 j Uacbride photos 
22779] (W— photo of type). 

VITEX TRIFLORA var. ANGUSTILOBA Huber 

Synonymy: Vitex tid flora angustiloba Huber apud Stapf , Ind. 

Lond. 6: U79. 1931. 

Additional bibliography: Moldanke, Phytologia 6: 161 — 162. 
1958; Moldenke, R6sum6 112 & Ii78. 1959; Moldenke, R^sum^ Suppl. 
15: 25. 1967. 

The Huber (I909) reference in the bibliograph of this variety 
is dated "1907-8" by Stapf (1931), but the pages involved seem 
to have appear first in 1909. 

VITEX TRIFLORA var. CORIAGEA Huber 

Synonymy: Vitex triflora coriacea Huber apud Stapf, Ind. Lond. 

6: U79. 1931. 

Additional bibliography: Moldenke, Phytologia 6: 162. 1958; 
Uoldenke, Rlsum^ 112 & 1;78. 1959; Lloldenke, R^sumS Suppl. 15: 
25. 1967. 

Recent collectors describe this plant as a shrub, 2 feet tall, 
or a large tree, knomi as "piquia-rana" , flowering in Novenber 
and December. The corollas on Archer 8OU7 are said to have been 
"lavender" . 

The Huber (I9O9) reference cited in the bibliography of this 
variety is dated "1907-8" by Stapf (1931), but the pages involved 
appear not to have been issued until 1909. 

Material has been inacciirately identified and distributed in 
herbaria as typical V, triflora Vahl. 

Additional citations: BRAZIL: Pari: Archer 80li7 (N); Barbosa 
de Silva 155 (N) . 

VITEX TRIFLORA var. FLORIBUNDA Huber 

Additional synonymy: Vitex triflora floribunda Huber apud 
Stapf, Ind. Lond. 6: U79. 1931. 

Additional bibliography: Moldenke, Phytologia 6: 162 — I63. 
1958; Moldenke, R^sumS 112, 387, & U78. 1959; Moldenke, RSsumS 
Suppl. 15: 25. 1967. 

As mentioned above under the other varieties of this species, 
the Huber (1909) reference in the bibliography is cited as 
"I907-8" by Stapf (1931), but it seems that the pages involved 
here did not actually appear in print until I909, 



1968 Moldenke, Notes on Vitex i;7 

AddlUonal citations: BRAZIL: Par^: Ducke 971 (W— 1332289) . 

^/ITEX TRIFLORA var. KRAATZII Ruber 

Additional synonymy: Vitex triflora kraatzii Huber apud Stapf , 
Ind. Lond. 6: U79. 1931. 

Additional bibliography: Moldenke, Phytologia 8: 83. 1961; 
Moldenke, R6sum5 Suppl. 1$: 25 . 1967. 

The original publication of this variety by Huber (1909) is 
inaccurately cited by Stapf (1931) as "1907-3". 

T/ITEX TRIFLORA var. QUINQUEFOLIOLATA Moldenke 

Additional bibliography: Moldenke, Phytologia 8: 83. 1961. 
The Ecuadorean collection cited below consists only of leaves 

and fruit and it is therefore placed here only tentatively. 
Additional citations: ECUADOR: Guayas: Gilmartin ^k& (W — 

2U28I4I2) . 

VITEX TRIFOLIA L., Sp. PI., ed. 1, 638 [as " trifoliis "] . 1753 
[not V. trifolia Grahaa, I966, nor Hemsl., 19U9, nor Moon, 
1395, nor Sess^ & Moc, 19liO, nor Vahl, 191^1, nor "sensu 
Matsumura & Hayata", I963] . 

Additional &. emended synonymy: Vitex triflora odorata , syl- 
vestris J. Buna., Thes. Zeyl. 209—210, pi. 109. 1737. Vitex 
incisa Wall, apud Watt, Diet. Econ. Prod. India 6 (U): 251, in 
syn. 1893 [not V, incisa Bimge, 1927, nor Lam., 1788, nor 
Thunb., 19ii7] . Vitex agnus castus var. Kurz ex Watt, Diet. E- 
con. Prod. India 6 (U): 251, in syn. 1893. Vitex trifolia L. f. 
ex K. Schum., Notizbl. Bot, Gart. Berl. App. 1: 55, sphalm. 
1895. Vitex trifoliolata L. apud J. Matsumura, Ind. Pi, Jap. 2 
(2): 53U— 535. 1912. Vitex trifoliolata var. trifoliolata 
Schau. apud J, Matsumura, Ind. PI. Jap. 2 (2): 53U — 535. 1912. 
Vitex trifolia Q, trif oliata Cham, apud Hara, Ertum. Sperm. Jap. 
1: 191, in syn. I9U8 . Vitex trifolia Oj trifoliolata Schau. 
apud Kara, Enum. Sperm. Jap. 1: 191, in syn. 19li8. Vitex tri- 
folia trifoliolata "Schau. ex Blanco" apud Stapf, Ind. Lond. 6: 
U79. 1931- Viiex trifolia L. ex Hosokawa, Journ, Soc, Trop. 
Agr. Taiwan 6: 206, sphalm. 19 3U. 

Additional tc emended bibliography: J. Buim., Thes. Zeyl. 209 — 
210 Sc 229, pi. 109. 1737} J. F. Gmel. in L., Syst. Nat., ed. 13, 
pr. 1, 2: 962 (1739) and pr. 2, 2: 962. 1796; Horsf., Verh. Bat. 
Gen. 8: lOU. I8I6; Pers., Sp. PI. 3: 36I. 1819; Steud., Norn, 
Bot., ed. 1, 838. 1821; Ro:!d)., Fl. Ind., ed. 2 [Carey], 3- 69. 
1832; Schnitzl., Icon. Fam. Nat. Reg. Veg. 137. 1856; Mason, 
Burmah & its People, ed, 2, I4I3, U79, & 792, i860; Miq,, Cat. 
Mus, Bot, Lugd.-Bat, 70, 1870; Beddome, Forester's Man. Bot, S. 
Ind. 172. 1873; Gamble, Man. Ind, Timb,, ed, 1, 296. I88I; 
Watt, Econ. Prod. India 5: 29li— 295. 1883; Vidal, Phan. Cuming. 
Philip. I3U. 1885; Warb. in Engl,, Bot, Jahrb, 13: i;28--i;29. 
I89I; v;att, Diet. Econ. ?rod. India 6 (U): 251. 1893; W. A, Tal- 



U8 PHYTOLOGIA Vol. 17, no. 1 

bot, Syst. List Trees Shrubs Bomb. I6l & 229. 189U: Jacks, in 
Hook, f . ^ Jacks., Ind. Kew., pr. 1, 2: 1213 ^ 12]X. 13?^; K. 
Schum., Notizbl. Dot. Gart. Berl. App. 1: 55 (1695) and 1: 2<jC. 
1896; Anon., Notizbl. bot. Gart. berl. App. 1: 3^6. 1897; K. 
Schum., Notizbl. Bot. Gart. Berl. App. 2: 1hk—lhS» I898; Anon., 
Notizbl. Bot. Gart. Berl. App, 2: Ul9. 1899; Gamble, l!an. Ind. 
Timb., ed. 2, 539- 1902; Prain, beng, PI., eri. 1, 2: 832--333. 
1903; C. B. Clarke in J. Schmidt, Bot. Tidsskr. 26: 173. 190U; E. 
D. Merr., Philip. Joum. Sci. Bot. 1, Suppl. 1: 121. 1906; Kaora- 
kami. List PI. Formos. 85. 1910; Duthie, Fl. Upper Gang. Plain 2: 
22U. I9II; Craib, Kew Bull. Uisc. Inf. 9: UI43. 1911; Craib, Con- 
trib. Fl. Siam Dicot. 16U— 165. 1912; Dunn ?c Tutcher, Ketw bull. 
Misc. Inf. Addit. Bar. 10: 20li. 1912; J. Matsumura, Ind. PI. Jap. 
2 (2): 53U--535. 1912; E. D. Merr., Interpret. Rumph. Herb. Am- 
boin. U53, 52ii, & 59U. 1917; H. J. Lam in Lam F.c Bakh., Bull. 
Jard. Bot. Buitenz., ser. 3, 3: 53. 1921; Haines, Bot, Bihar & 
Orissa U: 711 & 712. 1922; Nakai, Trees & Shrubs Indig. Jap., ed, 
1, 1: 350, fig. 190. 1922; H. N. Ridl., Joum. Malay Br. Roy. 
Asiat. Soc. 1: [Mai. For. Trees] 83. 1923; H, J, Lam in Engl., 
Bot, Jahrb. 59: 27, 28, & 92—93. 192U; C. J. F. Skottsberg, 
Medd. GOteb. Bot. Tradg. 2 [Haw. Vase. PI.]: 259. 1925; Gamble, 
Fl. Presid. Madras 2: 1101 & 1102. 192U; Mezger, Ann. Mus. Col. 
Marseille, s6r. it, h: pi. 60. 1926; A. W. Hill, Ind. Kew. Suppl. 
7: 252. 1929; C. A. Gardn., Enum. PI. Austr. Occid. 3: 112. 
1931; A, ¥, Hill. Ind. Kew. Suppl. 8: 2li9. 1933; Kanehira, Fl. 
Iticrones, 3U3 ^ U57. 1933; Tu, Chinese Bot. Diet., abrdg. ed., 
1337. 1933; Hosokawa, Joum. Soc. Trop. Agr. Taiwan 6: 206. 
I93I4; Terazaki, [Illustr. Fl. Jap.] fig. 2U99. 1938; Fletcher, 
Kew Bull. Misc. Inf. 1938: Ii31--li33. 1938; Comer, Card, Bull, 
Straits Settl. 10: 256—260, 1939; Jacks, in Hook. f. k Jacks,, 
Ind, Kew., pr, 2, 2: 1213 & 121U. I9U6; Selling, Bishop Mus, 
Spec, Publ, 38: 275 & Un, 19li7; L. H. Bailey, Man, Cult. PI., 
ed. 2, 81i3, 8U4, & inli, 19ii9; W. J, Bean in Chittenden, Roy. 
Hort, Soc. Diet, Gard. U: 2250, 1951; Hocking, Diet, Tenns Phai^ 
macog, 166 & 21^3. 1955; Kuck & Tongg, Mod. Trop, Gard, 77 & 236, 
1955; Darlington & Wylie, Chromosome Atl., pr. 1, 323, 1955; 
MoMenke in Humbert, Fl, Madag. I7I;: 71, 72, 79—33, & 273, Tig, 
10 (5 & 6). 1956; Anon., Biol, Abstr, 30: 1;370, 1958; H. St. John, 
Nomencl. PI. 7U. 1958; Moldenke, Biol. Abstr. 32: 2353. 1958; 
Anon., Kew Bull, Gen, Index 1929-1956, 293. 1959; Nath, Bot. Surv, 
South, Shan States 30l;--305. I960; Jacks, in Hook, f . 4 Jacks., 
Ind. Kew., pr, 3, 2: 1213 & 12lU, I960; Darlington & T/ylie, Chro- 
mosome Atl,, pr. 2, 323. I96I; Moldenke, Phytologia 8: 83 — 3U. 
I96I; Cave, Ind, PI. Chromosome Numb. 2: 137. 1961; Deb, Bull. 
Bot. Surv. India 3: 315. I96I; Hocking, Excerpt. Bot. A. 5: U2, 
1962; Moldenke, Biol. Abstr. 37: 1062, 1962; F. R. Fosberg, Bish- 
op Mus. Occas. Papers 23 (2): U. — U2, 1962; Thothathri, Bull. 
Bot. Surv, India li: 291. 1962; Hatusima, Mem, South. Indust. Sci. 
Inst. Kagoshima Univ. 3: 31. 1962; Van Steenis-Kruseman, Fl. 
Males. Bull. 3: 695 & LI. 1962: Li, Wood. Fl. Taiwan 973. 1963; 
Hocking, Excerpt. Bot. A.6: 53U. 1963; Prain, Bang. PI,, ed, 2, 
621 & 1012. 1963; Sharma & Mukhopadhyay, Joum. Genet. 58: 359, 



1968 Moldenke, Notes on Vitex U9 

366, 376, 379, & 539. 1963; E. E, Lord, Shrubs & Trees Austral. 
Gard., rev. ed., 232. 196Uj Cave, Ind. Pi. Chromosome Numb. 2: 
331. 1961ii Merminger, Seaside Pi. 32. 15U, & 155, pl. 223. 196U; 
Duffy, Joum. Appl. Ecol. 1^ 227—228, 231, 23U, 2li2, 2U3, & 2li8. 
I96U} Straatmans, Micronesica 1: 115. 1961i; Backer & Bakh., Fl. 
Java 2: 60U & 605. 1965; J. S, Beard, Descrip. Cat, W. Austr. PI. 
93. 1965; Moldenke, RfisumS Suppl. 12: 8. 1965; HStnsel, Leuckert, 
Rimpler, & Schaaf, Phytochem. h: 19 & 21. 1965; Quisumbing, Govt. 
Sarawak Sympos, Ecol. Res. Humid Trop. Veg. 35 & 36. 1965; Bose, 
Handb. Shrubs 96 & 97. 1965; Malick, Bull. Bot. Surv. India 8: 
55. 1966; Gaussen & al., Trav. Sect. Sclent. & Tech. Inst. Frang. 
Pond. Hors ser. 7: 71 & lOU. 1966; T. C. Whitmore, Giiide Forests 
Brit. Solomon Isls. 206. I966; Lourteig, Taxon 15: 28. I966; 
Moldenke, R6sum6 Suppl. 15: 15 & 25. 1967; Sauer, Plants & Man 
Seychelles 102. 1967; Moldenke, Phytologia 15: 78 & 267 (1967), 
15:' hl2 (I968), and 16: Ii95. 1968. 

Additional & emended illustrations: Terazakl, [Illustr. Fl. 
J^.], fig. 2li99. 1938; Moldenke in Humbert, Fl. Madag. I7U: 79. 
fig. 10 (5 & 6). 1956; Menninger, Seaside PI. 15U, pl. 223. 196U. 

Backer & Bakhuizen van den Brink (1965) describe this plant 
as being a very aromatic shrub, the stem erect, not rooting from 
the nodes, the leaflets 1 — 3, those of the 2- or 3-foliolate 
leaves either all sessile or the median (largest) leaflet on a 
petiolule of less than 0.5 cm, in length, ovate-elliptic to ob- 
long-obovate, the largest leaflet of the 2- or 3-i'oliolate 
leaves U — 9.5 cm. long and 1.7—3.7 cm, -wide, the unifoliolate 
leaves 2 — 6.5 cm. long, 1.3 — 3.5 cm. wide, all very densely- 
covered with white or gray hairs beneath; panicles narrow, 3*5 — 
2U cm. long; cymes 2 — 6.5 cm. long (including the 2 — 25 mm. 
long peduncle), 3 — l5~f lowered, rather dense to rather lax; 
calyx 3 — U.5 nmi. long; corolla-tube 7 — 8 mm. long; median seg- 
ment of the lower lip U — 6 mm. in diameter. They say that the 
species is found in teak forests, brushwood, secondary forests, 
and "periodically very much desiccating localities", and is also 
cultivated as a hedge-plant in Java. The make the further com- 
ment that "Some specimens closely approach the next species [ V. 
paniculata Lam.]". 

The corolla is described as having been "purple" on S. Olsen 
879 . Bose (1965) reports the plant as "very hardy, leaves 
simple or 3-foliolate", best propagated by the so-called "gootie" 
method. The plant has been collected in fruit in January as 
well as d\iring the months previously recorded by me. Cave (I96I, 
I96U) reports the diploid chromosome number for this species as 
26 and 3U. 

Deb (1961) says of this plant: "1ft. glabrous above, tomentose 
beneath, panicles white tomentose, corolla tomentose, lavender 
blue. Very common in valley, gregarious, in damp or moist waste 
land, along drains and roads or liver banks" and cites his no. 
128 . 

It should perhaps be noted here that the V. trifolia accred- 
ited to Graham is a synonym of V. negundo L., that ascribed to 



50 PHYTOLOGIA Vol. 17, no. 1 

Hemsley and to "sensu Matstunura & Hayata" is V_, trlfolia var. 
almpllclfolla Cham., that accredited to Moon Is V_. altlBsima L.f ., 
that of Sess^ ft. Moclflo is V, mollis H.b.K., while that ascribed 
to Vahl is V. trlflora Vahl. 

Accordinc to Lourteig (1966) the name, V_. trifolia L., is 
based on and typified by I\ Henaaim 70 . liocking (1955) informs 
us that the leaves of this plant have a volatile oil containing 
cineol and methyl alcohol, and that this oil is used luedicinally. 
Additional vei-nexsiilar names recorded for the plant are "Cayenne 
popper", "hamago", "kyaung banm ye-kyi-yo-ban" , "lagunding dagat", 
"mitsuba-hamagS", "pani-sanbhalu" , "shiru-fflki", and "tachi- 
hamago". 

Lord (1961i) recommends the species for planting in coastal 
climates in Australia, Malick (I966) reports it not so cocunon in 
West Bengal, citing Biswas 35. T, C. "iWhitmore (I966) cites Wa- 
terhovLse 60 from the Solomon Islands. Vidal (1885) cites Cvuning 
lit93 from the Philippines. 

Duffy (I96I1) states that " Vitex trifolia " [surely one of the 
varieties, not the true species 1] was introduced in 1858 on As- 
cension Island, having been received in a consigrment of 228 
species of plants from the Capetown Botanic Garden, and is now 
widespread on the island. He also avers that beetles are a fonn 
of insect life scarce on Ascension Island, but are found on this 
"Vitex trifolia " , as well as on Opuntia sind Acacia, there. 

Straatmans (I96U) informs us that V. trifolia is among the 
tropical seashore buoyant-seed plants in the coastal conxiunity on 
Eua island, but it is probable that he is here actually referring 
to var, bicolor (Willd.) Moldenke. 

The Lam (192li) reference in the bibliography of this species 
is often cited as "1925", but the latter is merely the title- 
page date for the volume; the pages cited appeared in I92I4.. The 
Blanco (I878) reference is dated "I878-8O" by Stapf (1931), but 
the plate which concerns us here seems to have been issued in 
1878. The "Basu, Ind. Med. Pi. pi. 2ii99" references given by me 
in the bibliography published in 1958 should be deleted; they 
are the result of errors in transcription for Terazaki, [Illustr. 
Fl. Jap.] fig. 2i;99 (1938). Prain (1903) writes the Watt refei^ 
cnces given in the bibliography above as "E. D. 5^ 181", but 
this is actually a paragraph reference, not a page referencel 

The following incoc^jlete bibliographic references occur in 
the literature of V, trifolia , but have not as yet been located 
by me in any library consulted: Aplin, Rep. on the Shan States, 
Settl. Rep. Chanda app, 6; Baden Powell, Fb. Pr. 36U; Boorsma, 
Plantenstoffen ii: HI; Cooke, Oils k Oil-seeds 81; Fleming, lied. 
PI. & Drugs [Asiatic Reser. 11] 181;; Gazetteer Mysore & Coorg 1: 
6ii; Koord., Natuurk Tijdschr. v. B. 1, li8: 89 and 20: 223; Ridl., 
Mai. Geneesmiddeln 28; Pharm. Ind, 163; Tijdschr. v. Land- en 
Tuihbotrw en Boschcultuur $: 55ii; Waitz, Practische Waamemingen 
10. 

The D. Anderson 211^3, Elmer 15236, Haenke s.n. [Mariana, 1792], 



1968 Moldenke, Notes on Vltex $1 

Kajewski 2U17 , and H. E. Parks 208^7 , distributed as typical V. 
trifolia , are all var. bicolor (Willd.) Moldenke; A, A, Hellor 
2731 , Taam 1702 , M. lu. To?maend s.n. [Oct, 20, 19iiO] , and C. 
Wright s.n. [Hong Kong] & s.n. [Bonin Islands] are var, simplici - 
folia Cham.} E. H. Bryan 131$ , Chapin 8^3 , E. I. Dawson 1982$ , F. 
R. Fosberg II98I & 36709 , K. P. Fosberg 15» £1 ^ Gillespie J438O , 
S^ K. Lau 270, £. W. Moore 696, Native collector DI.ll;9 [Herb. 
Roy. Forest Dept. 3$67], Quayle 1281 , £.£.£. Rock 232$ , 2969 , 
7838 , & s.n. [S. Kona, April 28, 1958] , H. Saint John 1U2$2 & 
16573, Schiffner 2li5U , A. C_. Smith U559 & 6078 , A. M. Stokes 1, 
Toroes 910, and Waterhouse 60 [Herb, Mus, Yale Sch. Forest. 
2266U] are all var. subtrisecta (Kuntze) Moldenke; and H^ Saint 
John 16705 is the type collection of var, subtrisecta f . albi- 
flora Moldenke. 

Additional citations: WESTERN PACIFIC ISLANDS: PHILIPPINE IS- 
UNDS: Mindoro: H. H. Bartlett 13707 (Mi) . Papahag: S. 01s en 
879 (Cp). IIDX)NESIA: GREATER SUNDA ISLANDS: Sumatra: Yates IU80 
(Mi), 19la (Mi). MELANESIA: BISMARK ARCHIPELAGO: New Britain: 
Pissing 2722 (Cp, Z). 

VITEX TRIFOLIA var. BICOLOR (Willd.) Moldenke 

Additional synonymy: Vitex negundo var. bicolor Lam., in herb. 

Additional k emended bibliography: Steud., Nam, Bot., ed. 1, 
888. 1821; Bocq., Adansonia 3: [Rev. Verbenac] 253. 1863; Watt, 
Diet. Econ. Prod. India 6 (U): 2I4.8. 1893} Jacks, in Hook, f, & 
Jacks., Ind. Kew,, pr, 1, 2: 1213. 1895; H. J. Lam in Engl,, Bot. 
Jahrb, 59: 27—28 & 93. 1921;; H, J, Lam in Bakh, & Lam, Nov, 
Guinea Hi, Bot. 1: I69. 192li; A. W. Hill, Ind. Kew. Suppl. 8: 
2U9. 1933; Corner, Card. Bull. Straits Settl. 10: 257. 1939; 
Jacks, in Hook, f, t Jacks., Ind, Kew,, pr. 2, 2: 1213. 19i;6; 
Hill & Salisb., Ind. Kew. Suppl, 10: 2hh* 191+7; Moldenke in Hum- 
bert, Fl. Madag. 17U: 72, 83, & 272—273. 1956; Yuncker, PI. 
Tonga 232. 1959; Jacks, in Hook. f. & Jacks., Ind, Kew., pr. 3, 
2: 1213. I960; Moldenke, Phytologia 8: 8U— 86. I96I; Backer & 
Bakh., Fl. Java 2: 605. 1965; B. C, Stone, Micronesica 2: 132, 
1966; Moldenke, Phytologia 1$: 306. I967. 

Backer & Bakhuizen van den Brink (1965) adopt the name, V. 
paniculata Lam., for this taxon, but admit that it is very fre- 
quently confused with V. negundo L. and is sometimes "difficult 
to be distinguished" from V. trifolia "with which it seems to 
hybridize." Corner (1939) says "This variety is so curiously 
intermediate between V. negundo and V, trifolia , that one cannot 
doubt that it covers their hybrids." Lam (192li) regarded 
Volkens U25 , from Yap, as a hybrid between what he called V, ne- 
gundo var. bicolor and V^ trifolia var. trif oliolata . In regard 
to the theory that this taxon is a natural hybrid between V, ne- 
gundo and V. trifolia , it is worth pointing out that it has been 



52 P H Y T L G I A Vol. 17, no. 1 

collected — often abundantly — on at least seventy-five ialanda 
in the Pacific Ocean area on which V, negundo does not occur, or, 
at least, has never been found and is very unlikely to occtir. I 
have no doubt that these two species do hybridize [see ander V. 
negundo in these notes], but this taxon does not represent thds 
hybrid. Nor do I feel that it is worthy of specific rank. As 
Backer and Bakhulzen van den Brink themselves admit, there are 
many specimens intermediate between it and the typical V. trifo - 
lia. 

Recent collectors and writers describe this plant as a shrub, 
6 — 12 feet tall, or a tree, 8 — 10 m. tall, with 3 — ^j leaflets per 
leaf, the petiole 2 — 6 cm. long, the middle leaflet on a petio- 
lule 0.5 — 2 cm. long, ovato-oblong or oblong-lanceolate, 3.5 — 10 
cm. long, 1.5 — 3*5 cm, wide, very acutely aciminate at the apex, 
the 2 adjacent leaflets (in the 5-xOliolate leaves) smaller or 
shorter-petioluled, the outermost leaflets (in 5--follolate 
leaves) smallest, sessile or subsessile; panicles pyra mid al -ovoid, 
lax, 6 — 20 cm. long, the cymes distinctly forked, 2 — 10 cm. long 
(inclusive of the pedxincle which is 5— UO mm. long), many- flow- 
ered, lax; calyx 1,5 — 3 nm. long; corolla-tube U — 5 dhh. long, the 
median lobe of the lower lip 3 — u mm. long and 2.5 — 3 nun. wide. 
The corolla is described as having been "blue" on Jaiiowsky 5l8 
and "pale-lilac" on Purseglove P.5Q15 * 

The plant has been collected on coral limestone, in thickets, 
above beaches, on sandy beaches and adjacent localities, especial- 
ly on the older parts of the beach-wall, rarely more inland. 
Yimcker (1959) says that it is occasional throughout Tonga and 
notes for its general distribution "Frcm eastern Africa and India 
through Malaysia to Polynesia. Presianably the V. trifolia of 
Honsley's and Btirkill's list." The leaves are used as a medicine 
in the treatment of fever in Samoa. Additional vernacular names 
recorded for it are "agulundi" and "garaulega" . Stone (1966) re- 
cords the plant from Nuktioro in the Caroline Islands, where it is 
known as "kSsik" « 

It should be pointed out here that the Lam (192ii) reference in 
the bibliography of this variety is often cited as "1925", but the 
latter date is merely the title-page date for the volume; the 
pages involved here appeared in the year 192U. In this work Lam 
cites Janowsky 5l8 frcxn Dutch New Guinea, Hollrung li86 , Lewandow - 
sky lt8 , Nyman 210, and Schlechter 11^253 fron Northeastern New Gui- 
nea, Dahl llt9 and Lauterbach 166 from New Britain, Kraemer s .n. , 
Ledermann llj.22 , and Raymxmdus 178 from the Palau Islands, Krae- 
mer s.n. and Ledermann 13531 from the Caroline Islands, and Haenke 
s.n. and HOfer 25 from the Mariana Islands. The Feudlleteau de 
Brxiyn )|T)| which he also cites is actually f , albiflora (Kuntze) 
Moldenke. He notes that Lewandowsky Ii8 shows one 1-foliolate 
leaf. 

The £. A. Price s.n. [May 10, 19U31, distributed as var. bi- 
color, is actually var. variegata Moldenke. 



1968 Moldenke, Notes on Vitex 53 

Additional citations: TAMGANYIKA: Tanner 2960 (S) . ZANZIBAR: 
H. G. Faulkner 2389 (S) . WESTERN PACIFIC ISLANDS: PHILIPPINE IS- 
LANDS: Cagayan: Kondo & Edafio s.n. [Philip. Nat. Herb. 39032] 
(Bi). Luzon: Elmer 1^36 (Bi) , Mindanao: Elmer 11999 (Bi). Min- 
doro: G. T_, Velasquez 11 (Bi) . Naranjo ; Kondo ^ Edafio s.n. 
[Philip. Nat. Herb. 38739] (Bi). PoliUo: R. C^ McGregor s.n. 
[Herb. Philip. Bur. Sci. 10270] (Bi). MARIANA, ISLANDS: Guam: H. 
M. Kayo s.n. [Oct. 2k, 19h7] (Bi) ; P. Nelson 522 (Bi), 53| (Bi, 
Bi) . Saipan: W. H. Lange 1(7 (Bi) . Tinian: R, S. CoTfan s.n. [A- 
pril 3, 19li5] (Bi); Hosokatra 7700 (Bi)j Kanehira 55 (Bi); Kondo 
1 (Bi), 58 (Bi) . Island undetermined: Haenke s.n. [Mariana, 1792] 
(Bi). INDONESIA: GREATER SUNDA ISIANDS: Sarawak: Purseglove P. 
5015 (N). Sumatra: Ltttjehams U655 (Bi, Bi) . MICRONESIA: CARO- 
LINE ISLANDS: Arekalong: Takamatsu 1697 (Bi). Dublon: Takamatsu 
I3U (Bi) . Ifaluk: Abbott & Bates 78 (Bi) . Kusaei: Takamatsu U87 
(Bi). Lele: Glassman 2716 (Bi) . Lxikunor: D. Anderson 2lU3 (Bi). 
Ponape: Takamatsu 780 (Bi). MELANESIA: NEVY GUINEA: Dutch New 
Guinea: Aet & Idjan 3U8 (A) . SOLOMON ISLANDS: Florida: Seale s_j_ 
n. [May 23, 1903] (Bi) . Guadalcanal: Kajewski 2la7 (Bi) . NEW 
HEBRIDES: Aneityum: Kajewski 8OI (Bi) . YASAWA FIJI ISUNDS: Fu- 
langa: A. C, Smith 1200 (Bi) . Kansavu: A. £. Smith 31It (Bi) . 
Koro: A. C. Smith 1075 (Bi) . Ovalau: J. W. Gillespie U503 (Bi, 
Bi). Taveuni: J. W. Gillespie 1|687 [wood no. 211^5] (Bi). Vanua 
Levu: A. C. Smith ^22 (Bi) . Viti Levu: E. H. Bryan 208 (Bi); 
Mac Daniels IOO8 (Bi); Meebold 161;92 (Bi), 21385 (Bi); H. E. Parks 
20800 (Bi), 20857 (Bi); Tothill & Tothill 66O (Bi) . LAU FIJI IS- 
LANDS: Thithia: E, H. Bryan 556 (Bi) . TOICAN ISLANDS: Eua: H. E. 
Parks 16178 (Bi) . Nomuka: Yuncker 15901 (Bi) . Tonga: McKem 27 
(Bi). Tongatabu: Yuncker 15011 (Bi) . POLYIESIA: VIESTERN SAMOA.: 
Savail: E. Christophersen 936 (Bi), 28U9 (Bi) . Upolu: A. £. 
Eames 36 (Bi). EASTERN SAMDA: Ofu: Yuncker 9566 (Bi) . Safotu: 
Vaupel 389 (Bi). Taut D. W. Garber 611 (Bi); Yuncker 9IOU (Bi) . 
Tutuila: W. A. Setchell 531 (Bi) . NIUE: Yuncker lOOiq (Bi) . 
COOK ISUNDS: Rarotonga: Parks & Parks 22573 (Bi)VGi P. Wilder 
1000 (Bi). CULTIVATED: Samoan Islands: D. W, - Garber 99^ (Bi); 
G. P. Wilder IjS [2li8] (Bi). 

VITEX TRIFOLIA var. BICOLOR f . ALBIFLORA (Kuntze) Moldanke 

Synonymy: Vitex agnus-castus ^ negundodes f . albiflora Kuntze, 
Rev. Gen. PI. 2: 510. I89I, Vitex agnus-castus var. negundodes 
f . albiflora Kuntze ex Moldenke, Rfistund 38O, in syn. 1959. Vitex 
trifolla var. bicolor f . albiflora Moldenke, Phytologia 8: 86. 
1961. Vitex agnus-castus var. negundoides f . albiflora Kuntze, 
in herb. 



$U PHYTOLOGIA Vol. 17, no. 1 

Bibliography: Kuntzo, Rev. Gen. PI. 2: $10. 1891; H. J. Laa in 
Bakh. 'a Lam, liov. Guin. lii, Bot. 1: 169. 1921^', Uoldenke, BAa\m6 
30O. 19^9; Itoldenke, Phytologia 8: 86. l?6l. 

Colleciora describe this plant as a tree, ^ m. tall, the trink 
9 cm. in diameter, the leaves white beneath, and the corolla 
white. 

The type of the form, as originally described by me, is H. ij. 
Parks 16178 , from Bua Island in the Tongan group. However, 
Kuntze apparently described the taxon earlier, based on a col- 
lection made by himself in Dakkan, Bombay, India. Since he al- 
so gave the taxon form rank, it is obvious that his description 
is the valid one and mine, being so much later, is illegitimate. 
Ilis collection, therefore, becomes the true type of the taxon. 

The Feuilleteau de Bruyn Ulit , cited by Lam (192U), apparently 
belongs to this form since its corollas as described as having 
been white. It was collected on Schouten Island, New Guinea, 
but I have not as yet been able to examine it, nor Kuntze 's type. 

VIT5X TRIFOLIA var. PURPUREA Lord, Shrubs Sc Trees Austral. 

Gard., rev. ed., 232 [as " trifolia ' purpurea '"] . I96I1; Uol- 
denke, R6sunl Suppl. 1$: 1$. 196?. 
The original description by Lord (1961i) of this variety is 
" Vitex trifolia ' purpurea ' with soft clean leaves, purple be- 
neath". It is apparently cultivated in Australian gardens and I 
know nothing else about it. 

VITEX TRIFOLIA var. SILIPLIGIFOLIA Chan. 

Additional L emended synonymy: Vitex trifolia var. unifollata 
Miq., Cat. Mus. Bot. Lugd.-Bat. 70. I87O. Vitex trifolia var. 
unifoliata Schau. ex Kawakani, List PI. Fomos. 35. 1910, Vitex 
rotundifolia L. ex S. Sasaki, List PI. Formos. 3S3 & 35U. 1928. 
Vitex trifolia unifoliolata Schau. ex Stapf, Ind. Lond. 6: U79. 
1931. Vitex trifolia unifoliolata "Schai. in DC." apud Worsdell, 
Ind. Lond. Suppl. 2: 501. 19iil. Vitex trifolia ovata Kak. ex 
fforsdell, Ind. Lond. Suppl. 2: 501. 19U.. Vitex agnus-castus Tl 
ovata (Thunb.) Kuntze ex Kara, Enum. Sperm. Jap. 1: I90, in syn. 
I9U8. Vitex trifolia Hemsl. apud Rehd., Eibliog. Cult. Trees 
585, in syn. 19U9 [not V. trifolia Graham, 1966, nor. L., 1753, 
nor L. f ., 1895, nor Moon, 1895, nor Sess^ ^ Moc, I9U0, nor 
Vahl, I9UI] . Vitex trifolia subsp. literal is Van Steenis, Blumea 
8: 516. 1957. Vitex trifolia var. unifoliata DC. ex Uoldenke, 
Phytologia 6: I8U, in 3301. 1958. Vitex trifolia var. unifoliola- 
ta DC. ex Itoldenke, Phytologia 6: I8U, in syn. 1958. Vitex ro- 
tundifolia var. rotundifolia Mizushima ex Uoldenke, Phytologia 
8: 86, in syn. I96I. Vitex trifolia var. ovata Schau. ex Uolden- 
ke, Phytologia 3* 86, in syn. I96I. Vitex trifolia "sensu Uat- 

sum. & Hayata" apud Li, Wood. Fl. Taiwan 83U, in syn. I963. Vi- 
tex trifolia var. unifolia Judd, in heA. Vitex trifolia ^ 



1968 Moldenke, Notes on Vitex 55 

vmifoliata Schau., in herb, Vitex tri folia var. ovovata Mak., in 
herb. 

Additional & emended bibliography: J, F. Gmel, in L,, Syst. 
Nat., ad. 13, pr. 1, 2: 962 (1789) and pr. 2, 2- 962. 1796; Pers., 
Sp. PI. 3: 3S9, I8l9j Steud., Norn. Bot., ed. 1, 888. 1821} Hook. 
L Am., Bot. Beechey Voy. 206, pi. U7. I836; Miq., Cat. Mus. Bot. 
Lugd.-Bat. 70. I87O; Jacks, in Hook. f. & Jacks., Ind. Kew., pr. 
1, 2: 1211i. 1895; C. B. Clarke in J. Schmidt, Bot. Tidsskr. 26: 
173. I90U} E. D. Merr., Philip. Journ. Sci. Bot. 1, Suppl. 1: 121, 
1906; Matsumura & Hayata, Journ, Coll. Sci. Univ. Tokyo 22: 301, 
1906 J Kawakami, List PI. Fomos. 85. 1910 j J. Matsumura, Ind. PI, 
Jap. 2 (2): 53U— 535. 1912; H. J. Lam in Lam & Bakh., Bull, Jaad. 
Bot. Buitenz., ser. 3, 3: 53. 1921; Nakai, Trees & Shrubs Indig. 
Jap., ed. 1, 1: 350, fig. 190. 1922; H. J. Lam in Engl., Bot. 
Jahrb. 59: 27. 192li; H. J. Lam in Bakh. & Lan, Nov. Guin. lU, Bot. 
1: 169. I92U; C, J. F. Skottsberg, Medd. GOteb. Bot. TrSdg. 2 
[Haw. Vase. PI. 1]: 259. 1925; S. Sasalci, List PI. Fomos. 3S3 & 
35U. 1928; Tu, Chinese Bot. Diet., abrdg. ed., 1337. 1933; Hoso- 
kawa, Journ. Soc. Trop. Agr. Taiwan 6: 206. 1931;; Fletcher, Kew 
Bull. Misc. Inf, I938: U31~U33. 1938; J. Matsumura, [Bot, & 
Zool.] 10: 288, fig, 125. 19U2; Jacks, in Hook, f, & Jacks,, 
Ind. Kew., pr, 2, 2: 1211;, 19U6; Selling, Bishop Mus. Spec. Publ. 
38: 275 & 101. I9li7; Li & Keng, Taiwania 1 (2—1;): 127. 1950; 
Van Steenis, Bluraea 8: 516, 1957; Anon,, Biol. Abstr, 30: U370, 
1958; Cave, Ind, PI. Chrcmosome Numb. 1: U6, 1958; Moldenke, Bi- 
ol. Abstr. 32: 2353. 1958; Anon., Kew Bull. Gen. Index 1929-1956, 
293. 1959; Jacks, in Hook, f, & Jacks,, Ind, Kew,, pr. 3, 2: 
1211; . I96O; Kitamura & Okamoto, Col. Illustr. Trees St Shrubs 
Japan 221, pi. 65. I960; Moldenke, Phytologia 8: 86—83, I96I; 
Hocking, Excerpt. Bot. A. 5: 1;2. 1962; F. R. Fosberg, Bishop Mus. 
Occas. Papers 23 (2): la— 1;2, 1962; Nobuhara, Okada, & Fujihira, 
Jap. Journ. Ecol. 12: 101—103, 105, & 107. 1962; Liu, Illustr. 
Nat. & Introd. Lign. PI. Taiwan 2: 1231, pi. 1039. 1962; M. J. 
Van Steenis-Kruseman, Fl. Males, Bull, 3: 695 & LI. 1962; Hatu- 
sima, Mem. South. Indu»t, Sci, Inst, Kagoshima Univ. 3 (1): 31. 
1962; Li, Wood. Fl. Taiwan 832, 83I;, & 973. 1963; Chuang, Chao, 
Hu, 5c Kwan, Taiwania 1 (8): 5U, 58, & 63, pi. 3, fig. UO. 1963; 
Taniguti, Amat. Herb. 2h (3): 9. 1963; Cave, Ind. PI. Chromosome 
Numb. 2: 331. 196U; Neal, In Card. Hawaii, ed. 2, 728, fig. 277. 
1965; Backer k Bakh., Fl. Java 2: 60i;. 1965; Ohwi, Fl. Jap. 765. 
1965; Hatusima, Mem. Fac. Agr. Kagoshima Univ. 5 (3): 1;7 — US. 
1966; Nobuhara, Journ. Jap. Bot. I9: 326—328, 330, 332—331;, 
336—338, 3U— 3l;5, & 3U8. 1967; Moldenke, I^sum^ Suppl. 15: 25. 
1967: Moldenke, Phytologia 15: 267 (1967), 15: 1;72 (I968), and 
16: I;95. 1968. 

Additional L emended illiistrations: Hook, & Am,, Bot, Beech. 
Voy. pi. 1;7. 18 36; Nakai, Trees & Shrubs Indig. Jap., ed. 1, fig. 
190. 1922; J. Matsumura, [Bot, & Zool,] 10: 288, fig, 125. 19i;2; 
Kitamura & Okamoto, Col. Illustr. Trees & Shrubs Japsin pi. 65 
[in color]. I96O; Chuang, Chao, Hu, & Kwan, Taiwania 1 (8): 63, 
pi. 3, fig. 1;0. 1963; Neal, In Card. Hawaii, ed. 2, 728, fig. 277. 
1965. 



$6 P H Y T L a I A Vol. 17, no. 1 

Recent collectors and writers describe thds plant as a procuja- 
bent or ascending, creeping shrub, 6 — 30 cm. tall, or a woody 
trailing vine, the whole "plant with stinky odor", the main sten 
1 — 2 m, long, often entirely buried in the sand from which only 
the flowering branchleta emerge, densely gray-white puberulent 
throughout; stems creeping, copiously rooting at the nodes, emit- 
ting many, er«ct, short, flowering branchlets; branches U-angledj 
leaves 1-foliolate; petioles 1.5 — 3.5 nnn. long ["cm" by error in 
Backer & Bakh. (1965)]; leaflet-blades herbaceous, broadly ovate 
or broadly elliptic to oval-elliptic-obovate, 1.5—5 cm. long, 
1.3 — 3.5 cm. wide, obtuse to rounded at the apex, entire (or a few 
2- or 3-partite), abruptly acute at the base, green and thinly 
puberulent above, densely grayish-puberulent beneath or densely 
white-tonentose especially beneath; panicles terminal, narrow, 1— 
9 cm. long, densely flowered, with very short branches; peduncles 
1 — ii cm. long; cymes 1 — U-f lowered, the lower ones often in the 
upper axLls of the leaves; corolla blue, light-blue, or bluish- 
violet to purple-blue, purple, deep-purple, lavender, or red, a- 
bout 13 mm. long, from the insertion of the stamens inside up to 
half the length of the lower lip densely white-hairy, silky- 
pubescent on the outer surface, the tube about 7 jam, long, the 
median segment of the lower lip about 5 nnn. long; calyx greenish, 
silky-pubescent; style about 15 mm. long; bases of the filaments 
villous; fruit drupaceous, globose, dry, black, 5 — 7 mm, wide, 
the lower half enclosed by the persistent fnii ting-calyx; pyrenes 
corky. 

The corolla is described as having been "bluish-i^iite" on F_. 
R, Fosberg 8S81 , "purple" on Hxirusawa 202 , "blue" on H. L. Porter 
3 and E. H. Wilson 10978 , "deep-purple" on Ichikawa 2006^1 , "red" 
on Tsang s.n. [Kerb. Lingnan Univ. 166U9] , "lavender" on R. C. 
Ching 1967 , "light-blue" on McClure s.n. [Kerb. Lingnan Univ. 13095], 
and "purple-blue" on Liang 62926. Cave (1958) reports the haploid 
chromosome number as 16. 

It should be noted here that the V^ trifolia of Linnaeus the 
elder is a valid species, with the homonym ascribed to Linnaeus 
the younger as a synonym, while the V, trifolia of Graham is 7. 
negundo L., that accredited to lioon is V, altissina L. f ., that of 
Sess6 & Mociflo is V^ mollis H,B,K., and that ascribed to Vahl is 
V. triflora Vahl. 

Vitex trifolia var. simplicifolia has been collected on sandy 
beaches, in sandy places by the sea, on loany seashores, and along 
rocky roadsides, blooming from July to September. Fosberg (1962) 
and Comer (1939) feel that the plant should be called V, ovata 
Thunb., and in this they are followed by Backer Sc Bakhuizen van 
den Brink (1965) who note "Ridley. .. .states that he saw specimens, 
transplanted into the interior, develop into V. trifolia . If 
this statement proves correct, V. ovata has to be considered an 

edaphic form of V. trifolia I never saw any transitional form, 

nor were such forms ever observed by Comer." 



o 



PHYTOLdM&^SARY 

Designed to expedite botanical publication 



:i;C3 



Vol. 17 August, 1968 No. 2 



BOTANICAL GARDEN 



CONTENTS 



BOIVIN, B., Flora of the Prairie Provinces (part) 57 

DEGENER, O. & I., Retiezf 113 

MOLDENKE, H. N., f<otes on new and noteworthy plants. L. . . . . 113 
MOLDENKE, H. N., Additional notes on the genius Vitex. IX . . . .114 



Published by Harold N. Moldenke and Alma L. Moidenke 

303 Parkside Road 
Plainfield, New Jersey 07060 
U.S.A. 

Price of this number, $1; per volume, S6.75, in advance, 
or S7 at close of volume 



r 



PROVENCHERIA 
3 
Memolres de I'Herbier Louis-Marie 
Facultg d 'Agriculture, Universite Laval 



FLORA 
OF THE 

PRAIRIE PROVINCES 



A HANDBOOK 
TO THE FLORA OF THE PROVINCES OF 
MANITOBA, SASKATCHEWAN AND ALBERTA 

by 

BERNARD BOIVIN 

Herbier Louis-Marie, Universite Laval 
and Department of Agriculture, Ottawa 



Part II 

Digitatae, Dimerae, Liberae 
(Continued) 



58 PHYTOLOGIl Vol. 17, no. 2 

2. SCLERANTHU:: L. KNAWPX 

Sepals fused; the tube becoming thick and hard arxl enclos- 
ing the utricule. Petals lacking. 

1. S. ANNUUS L. -- Knawel, German Knotgrass (Q navelle^ 
Herbe aux alouettes) — Leaves opposite and connate in the man- 
ner of a Caryophyll. Puberulent annual with numerous stems. 
Flowers green. Calyx lobes membrancws-margined, slightly longer 
than the tube. Early to mid summer. Uncommon weed of roadsides 
arid cultivation.— NS-0, S-BC, (US), Eur. 

A Manitoba report by Montgomery I96U is not substantiated 
by any specimen at OA.G or elsewhere (Montgomery in litt.). 

Order hh. GHENOPCDIALES 
Like the Polygonales , seems to be derived from the Garyo- 
phyllales j with the fruit reduced to a 1-seeded utricule or 
achene. But the flowers typically $-merous and the embryo, vi- 
sible through the seed coat, is annular or spirally curled. 

a. Flowers bractless, or exceptionally subtended 

by herbaceous bracts 78. Ghencpodiaceae 

aa. Each flower subtended by scarious bracts .. 

79. Amaranthaceae p. 129 

78. GHENCPCDIACEAE (GOOBEFOOT FAMILY) 

Herbs often thickish or fleshy. Hairs often short and 
thick, ± subglobular. A family usually readily recognized by 
the curled eirbryo and the usually semi-fleshy and alternate 
leaves , 

a. Fleshy herb with vestigial leaves 12, Salicomia 

aa. Leaves well developed, 
b. Shrubby. 

c . Very spiny 13 . Sarcobatus 

cc. Not spiny. 

d. Leaves flat 5. Atriplex 

dd. Strongly revolute 7 . Eurotia 

bb. Annual herbs. 

e. Fruit hidden between a pair of bracts. 

f. Bracts free at least above the 

middle 5. Atriplex 

ff . Bracts fused to the tip and enclo- 
sing the fruit h. Spinacia 

ee. Fruit not hidden. 

g. Fruit flanked by a pair of fused 
bracts; pistillate flower without 

perianth 6, Suckleya 

gg. Pistillate flower and fruit bract- 
less, or the bracts neither fused 
nor hiding the fruit, 
h. Galjrx imich reduced and not sur- 
rounding the fruit. 
SCLERANTHUS 118 



1968 Boivin, Flora of Prairie Provinces 59 

i. Main leaves hastate to 

rhomboid-lanceolate 3« Monolepis 

ii. Leaves * linear 11. Corispermum 

hh. Fruit surrounded by the 

marcescent calyx Group A 

Group A 
Annual herbs. Fruit surrounded by the marcescent calyx. 
Bracts lacking or small. 

a. Flowers unisexual, the staminate ones borne 
in a conspicuously differentiated terminal 

spike 8. Axyris 

aa. Flowers all perfect or scxne of them pistillate, 
b. Upper leaves and bracts stiff and ending 

in a sharp and spiny point 1$ . Salsola 

bb. Foliage not spines cent. 

c. Fruit surrounded by a continuous 

horiz ontal wing 2. Cycloloma 

cc. Not winged or with a discontinuous 
series of winged lobes. 
d. Foliage glabrous or glandular or 
mealy. 

e. Calyx thin 1, Chenopodium 

ee. Calyx fleshy. 

f . Flowers in axillary glccie- 

rules of 3 lU . Suaeda 

ff . Fmiits in large strawberry- 

like glcmerules 1. Chenopodium 

dd. Foliage pubescent, the leaves and 
bracts long ciliate. 
g. Inflorescence densely pubescent, 

including the calyx 10. Bassia 

gg. Calyx glabrous, or the lobes 

scmetimes ciliate 9. Kochia 

1. CHENOPODIUM L. GOOSEFOOT, PIGWEED 
The basic and unspecialized type of the family. Flowers 
bractless, perfect, - $-merous, with a persistent calyx enve- 
loping the fruit. 

a. Fruit a large strawoerry-like glomerule . . . . 1. C. capitatum 
aa . Fruit not or very little fleshy and the 
inflorescence less congested. 
b. Leaves narrowly lanceolate to linear, 
entire or nearly so. 
c. Grayish -mealy, especially on the 

undersurface of the leaves 5, C. leptophyllum 

cc. Pale green and nearly glabrous.... UT C . subgja'brum 
bb. Leaves oblong-lanceolate to deltoid, ~ 
mostly coarsely toothed. 

119 CHENOPODIUM 



60 PHYTOLOOIA Vol. 17, no. 2 

d . Leaves deltoid, nearly as broad as 

lonp and *■ truncate at base 6. C. rramontil 

dd. Leaves ovate to oblong-lanceolate, 
roxinded to cuneate at base, 
e. Plants t^labrous and green. 

f. Tjcaves entire or essentially 

so 9. C. polyapermum 

ff. Leaves lobed. ~ 

g. Seeds mostly vertical, 

Q.b-1.0 mm wide 3. C. rubrmn 

e,p . Seeds horizontal, 

l.U-3.0 mm wide 10. C . hybridum 

ee. More or less mealy-puberulent, 
especially in the inflorescence 
and undersurface of the leaves, 
the latter paler to whitish below, 
h. Seede mostly borne vertically, 

1mm wide or less 2. C. glaucum 

hh. Seeds nearly all horizontal, 
1 ran wide or more. 
i. Early flowering, the main 

leaves typically ovate 8. C. album 

ii. Late flowering, the main 

leaves oblong and subentire .. 
7. C. strictum 

1. ^. y^gitatug^ (L.) Asch. (Blitum capitatum L.) — Straw- 
berry-31ite, Indian Paint (Blette) — Calyx becoming fleshy and 
bright red at mat'ority. Leaves triangular-hastate, coarsely 
dentate. Fruiting calyces aggregated in strawberry -like fruits, 
these partly axillary, partly in terminal leafless racemes. 
Early summer. Infrequent but conspicuous in disturbed or shal- 
low soils. —Mack^ka, NS, NB-3C, US, (Eur). 

An Alberta report of C. Bonus Henricus L. by Groh 1$50 was 
based on H. Groh , Edson, 1'535 (DAO), a sheet since reidentified 
to C. capita turn . 

G. f olioium (Moench) Asch. was reported by Wahl 195ii.> pa- 
ge 9,'"as was C. vlrgatum (L.) Ambrosi by Aellen 1929, page Uii. 
Both from Alberta and both based on a sheet, A.H. Brinionan 28^8 , 
Battle, woods, Aug. 28, 1927 (Aellen; DAO, photo), revised 5y 
Aellen to C. capitatum more than a quarter of a century ago. 
We concur. 

2. C. Rlaucum L. var. pulchrum Aellen (var. salinum 
(Standley) Boivin; C. s a lin'om StandYey ) — Leaves tending to be 
the smallest, vmitis^h-mealy below, nearly glabrous above. Erect 
to creeping and very branchy. Leaves broadly lanceolate and 
coarsely few -toothed. Fruit peltate or mostly erect, i.e. la- 
terally compressed, and aoout 1 ram wide. Mostly after mid sum- 
mer. Mostly exsiccated saline shores, often weedy. — (K)-Mack, 
Aka, Q-BC, US. 

In our variety the glcmerules are gathered on ultimate 
branchlets bearing reduced leaves almost to the tip, tepals are 
CHENOPODIUM 120 



1968 Boivin, Flora of Prairi* Provinces 61 

mostly obovate and the inflorescence is often farinose -puberulent. 
The eurasian var. glaucum is none too readily recognized by its 
flowering branchlets almost devoid of reduced leaves, except to- 
wards the base, its mostly elliptic or oblong tepals and its gla- 
brous inflorescence. In Eastern Canada both varieties will be 
met with as infrequent weeds. 

5* /£.• ru br um L. (var. hunile (Hooker) Watson; C. chenopo - 
dioides (J?. )^Aeltehj C. humile Hooker) — Fat Hen, French Spi- 
nach — Stamens only 1-2 and the fruiting calyx reddish and slight- 
ly fleshy. Plant erect to depressed, glabrous or nearly so. 
Foliage thickish. Leaves - rhombic-triangular, lobed, the lobes 
inclined forward. Glooierules less than 5 nun wide, rather nume- 
rous. Fruit erect, 1 mm wide or less. Mid summer. Saline sho- 
res, rarely weedy. ~ sMack, Y, (NF-SPM), NS, NB-BC, US, Eur. 

The basis for an Alberta report of C. ambrosioides L. by 
Groh 19$0 seems to be the collection G.H. Turner U3> Fort Sas- 
katchewan, garden, Aug. U, 1537 (DAO), since revised to C. rubrum 
by Dr. H.G. Wahl in 1953- 

Scmietimes divided in two or, more rarely, in three species. 
Plants of more open habitats, and especially of pioneering ha- 
bitats, are more or less depressed (C. humile ) ; obviously an 
ecological form. More luxuriant specimens have rarely been se- 
gregated as C. chenopodioides . 

U. C^. subglabrmn (Watson) Nelson (C. leptophyllum Nutt. 
var. subglabrum Watson) — Similar to the following, but barely 
pubemilent and thus pale green in colour. Main stem leaves usu- 
ally quite glabrous, becoming slightly mealy in the inflorescen- 
ce. The latter broad and diffuse, with scattered flowers. First 
half of summer. Rare or inconspicuous pioneer on wind eroded 
sand. — (swO)-Man-S, US. 

A species of eroded dunes, it is almost skeletic and thus 
easily overlooked. It may be much more common than herbarium 
sheets indicate. Thus far we have only one Manitoba record: 
Boivin & Laishley l3l3o , entre Oak Lake et Routledge, k k milles 
au nord du lac de Ch&nes, dune active, U juillet 19^9 (DAG). 

5. C. leptophyllum Nutt. (var. oblongifolium Watson; C. 
dessicatum Nelson; C. pratericola Rydb., ssp. dessicatum (Nel- 
son) Aellen) — Grayish -mealy, and usually virgate, annual herb 
of li^t soils. Leaves narrow and entire or with a pair of weak 
lobes, grayish -mealy at least below. Fruits mostly horizontal, 
about 1 mm wide. Around mid summer. Steppes, especially on 
light or wind -eroded soils. — Y, NS, swQ-BG, US, Eur, (GA). 

6. C. FrgJJl^itii Watson (C. atrovirens Rydb.) — Mostly 
occuring as a native annual in dry woods. A rather facile and 
stiffly erect nerb. Leaves usually thin and wilting very quick- 
ly. Spikes of glOTverules very remotely moniliform. Fruit ho- 
rizontal, 1.2-1.5 mm wide. Early to mid summer. Dry woods; 
often under shrubs, especially Rrunus , sometimes on shores. — 
s**Ian-BG, US, (GA). 

An extension of range to Yukon by Hulten 1950 was based on 
Anderson & Brown 103U7» near Garcross, alkali flat, 3^ July, 
i9U& 1.CAN ; DAG, photo), since revised to G. rubrum . 

121 GHENCPCDIUM 



62 PHYTOLOGIA Vol. 17, no. 2 

Thicker-leaved plants are sometimes identified C. atrovi - 
rena . 

7. C. STRICTUM Roth (var. Klaucophyllum (Aellen) Wahl; C. 
flaucophyllum Aellen) — Resembling the following, out flowering 
later and cocrlmonly larger. Lower leavea ovate and shallowly 
serrate but the middle and upper oblorp and - entire. Calyx 
lobes * elliptic. FVuit 1 mm wide or slightly larger. Late 
summer and early fall. Waste places and disturbed soils > es- 
pecially in towns. — sQ-sS, BC, US. 

Introduced plants in North America are usually distingoished 
as var. Rlaucophyllum but the souridness of tne distinction is 
questionable. The eurasian material at hand does not coriform in 
its reputed differences witn tne neogean phase. Seems likely tnat 
here many varietal identifications were made oil the basis of 
geography rather than morjihology. 

Most floras do not distinguish this species but it appears 
to be rather widely distributed in botn Canada and tne U.S.A. 
In tne field C . strictum is rather readily spotted by its prefer- 
ence for town¥ and waste -lots, its late flowering, its brancning 
and its leaf dimorphism. C. strictum does not attract atten- 
tion and does not begin to riower~~until late Aup^ast or early 
September, at a time when C. album is already heavily loaded with 
ripe fruits and shedding tnemi CT strictum is also heavily 
branched down to the base, the many Icwer branches are closely 
set togetner and often nearly as long as tne stem. The stem 
leaves are rather similar to those of C. album but the branch 
leaves are mostly entire and oblong-elliptic. Because of its 
heavy branching and size, usually a good meter tall, C. strictum 



does not lend itself to making pood specimens and tne average 
herbarium sheet is likely to be a mere snipping or a selected 
small (hence often depauperate) individual. But the later flo- 
wering time, the narrower shape of the calyx lobes and the smal- 
ler fruits snould provide good diagnostic features. 

8. G. album L. (f. lanceolatum (Muhl.) Aellen; C. Berlan - 
dieri Moq., var. farinos ijm (Ludwig) Aellen; C. Boscianum Moq.; 
C. dacoticum Standley; C. lanceolatum yuhl.;~C. paganum Reich.; 
C. Zschackei Murray) — ~Pigweed , Lamb's Quarters (Chou gras, 
Poulette grasse) — The common middling type. Annual erect 
herb, * mealy, especially on the lower leaf surfaces. Main lea- 
ves more or less ovate and coarsely toothed. Calyx lobes del- 
toid. Seed ± 1.5 mm wide, borne horizontally. Mostly mid sum- 
mer. Common weed of disturbed soils and humanized places, see- 
mingly native on shallow soils over rocky outcrops. — (G), 
Mack^ka, L-NF-(Sm), NS-BC, US, (CA), Eur. 

Plants with larger leaves and fruits have been distinguished 
as C. Bushianum Aellen or C. paganum . The merit of the distinc- 
tion is not clear to us. 

Native plants are reputedly distinguishable (as C. Boscianum 
or C. Berlandieri) by their ovaiy wall free from the achene or 
by being more preeminently keeled on the sepals, characters 
which have also been detected in a number of european specimens 
at hand. We are not yet satisfied that seemingly native plants 

CHENCPCDIUM 122 



1968 Boivin, Flora of Prairie Provinces 63 

can be convicingly discriminated on these or any other charac- 
ters. 

9. C. POLYSPf-RMUM L. (var. acutifolivun (Sm.) Gaudin) — 
Allseed (Limoine, Poiree sauvage) — Leaves glabrous, thin and 
entire, the main ones ovate to lanceolate. Seed maturing pur- 
ple-red, then black, about 1 mm across, horizontal. Second 
half of summer. Rare town weed: Wallwort. — NB-0, S, US, Eur. 

10. C^. ]iY^33^ L. var. gi gant os pernrum (Aellen) Rouleau — 
Sowbane (Pied d^oieT" — Large, uffn^'^'ovate^eaves with i 3 pairs 
of large teeth or lobes . Flowers mostly in terminal panicles . 
Fruit greenish. Mid to late summer. Infrequent in dry woods 
and casually weedy. — Y, NB-BC, US. 

American plants are supposed to have larger seeds, but our 
specimens do not conform to this pattern. However, our Canadian 
specimens do have black, shiny and essentially smooth seeds, 
while our European ones (var. hybridum ) have seeds that are dull 
and finely but clearly rugose -reticulate . 

A Saskatchewan report of C_. Bonus - Henricus L. by Groh 1950 
was based on two sheets of which the first, Shevkenek 127 , 
Qu 'Appelle Valley, 1938 (DAO) is now filed under C. hybridum 
var. gigantospermum , while the other, Garmichael 37, Regina, 
19U1 (DAO) has since been revised to Atriplex hoi^nsis . 

2. CYCLOLOMA Moq. WINGED PIGV/EED 

Calyx developing a peripheral wing at maturity. Otherwise 
as in Chenopodium. 



1. C^. ati^^ljcifolium (Sprengel) Coulter — Tumbleweed — 
Resembling CnenopodTunr "but 1 i gh tly lanate and not mealy. Lea- 
ves - oblanceolate, coarsely toothed. Flowers in moniliform 
spikes. Fruit about 3 mm across including the wing. Seed con- 
cave above, convex below. Mid to late summer. Disturbed 
sands: Agassiz Delta, Grande-ClairiSre. — swQ-sMan, US. 

We have been unable to substantiate a report from Baildon, 
Sask., by Russell 19Ui, 19$U, Groh 1950 and Breitung 1959, re- 
peated by Boivin 1966. 

3. MONOLEPIS Schrader 
Calyx reduced to a single sepal which thus takes on the 
appearance of a small bract. 

1. y^. Nuttalliana (R. & S.) Greene — Povertyweed — Ra- 
ther res emblrrig Chenopod ium glaucum in general habit and leaf 
shape but the inflorescence much more leafy. Leaves not white 
below, merely slightly mealy. Fruit apiculate. Early summer. 
Native on saline shores, but mainly found as a weed of distur- 
bed soils. — Mack-^ka, Q-(0)-Man-BC, US, (CA, SA). 

U. SPINACIA L. SPINACH 

Resembling Atriplex but the pistillate bracteoles fused 
all around and forming an accessory envelope around the seed. 
Flowers dioecious . 

123 SPINACIA 



6U PHYTOLOGIA Vol. 17, no. 2 

1. S. OLfJUCEA L. — Spinach (Epinard, Spina g^e ) — f-'njit 
with 2-U lons^ spiny lobes. Loaves nastate to trianfnjlar, ra- 
ther large. Stamlnate flowers in spikes of glomerules. Pistil- 
late flowers in axillary glomerules. Early sujnmer. Sometijiee 
cultivated, rarely occurinp as a waste grourid or roadside weed. 
— Mack, (Aka),Alta, (USi Eur. 

5. ATRIPLEX L. CRACHE 

Flowers dimorphic, the pistillate ones reduced to a naked 
ovary between 2 bracteoles. Staminate flowers as in Chenopo - 
dlum. Pistillate bracts fused at base only. 

a. Shrubby 1. A. Nut* alii 1 

aa . Annual herbs. 

b. Pistillate bracteoles orbicular and 

entire 2. A. hortensis 

bb. Bracts variously shaped and cut. 

c. The whole plant, and especially the 
leaves , more or less silvery, being 
densely covered by a scaly or mealy 
puberulence. 
d. Pistillate bracteoles coarsely 

toothed to summit Ii. A. argentea 

dd. Entire above the middle $. A. ?c»/ellii 

cc. Leaves glabrous or ligitly mealy. 

6. Terminal spikes entirely staminate, 
the pistillate flowers borne only in 
inconspicuous axillary clusters. .. .6. A. dioica 
ee. Terminal spikes at least partly ~ 
pistillate, except in entirely 
staminate plants 3 . A . patula 

1. A. Nuttallii Watson var. Nuttallii (A. canescens AA., 
var. aptera AA.) — Salt-Sage, Moundscale — Semi -shrubby, pro- 
ducing numerous erect herbaceous shoots frcm a woody base. Fo- 
liage densely mealy-puberulent and grayish-silvery. Herbaceous 
shoots simple, but with numerous axillary tufts of small leaves. 
Dioecious. Staminate flowers in yellow, moniliform, flexuous, 
and bractless spikes of glomerules. Pistillate flowers in a 
leafy terminal spike of glomerules. Mid summer. Eroded hills 
and badlands, sometimes in steppes on saline soils. — s\*lan- 
Alta, US. 

Leaves mostly 0.5-1.0 cm wide and rather elliptic-lanceo- 
late to oblong-lanceolate. Otner varieties occur further soutn, 
including a var. falcata M.E. Jones with narrower and rather 
linear leaves. 

All previous reports of A. canescens (Pursh) Nutt. and of 
its var. aptera (Nelson) G.L.~Hitchc, from our area were based 
on specimens of A. Nuttallii . This remark includes the Moodie 
collection from Rosedale (GH; DAO, pnoto). 

2. A. HCRTENSIS L. (A. nitens Schrank) — Orach, French 
Spinach (Bonne -dame, Arrocne) — Fruit larger, suborbicular, 

ATRIPLEX 12U 



1968 Boivin, Flora of Prairie Provinces 6$ 

entire, flat, ± 1 cm across. Tall, conspicuous, virgate herb. 
Leaves triangular, rather large, the lower remotely dentate, the 
upper entire, whitish -mealy below. Mid summer. Sometimes culti- 
vated and readily spreading to waste places and railway yards. — 
swMack, (Aka), Q-BC, US, Eur — Cv. ATR03ANGUINEA ~ Stem leaves 
and fruits more or less tinged in bright red: Hoosier. — S. 

3. A. patula L. var. patula (var. hasta ta (L.) Gray; A. 
hastata L.) — Spearscale (, Belie dame. Bonne dame) — Resembling 
a Ghenopodium, but with about 3 main pairs of stem leaves being 
opposite. Diffusely branched. Leaves deltoid to lanceolate, 
± dentate, the 2 lower teeth much larger. Flowers in terminal 
spikes which are bractless at least above tne middle. Mid sum- 
mer and early fall. Native in saline places and a frequent 
weed of towns and disturbed soils. — (seK)-Mack, (Aka, NF)-SPM, 
(NS4^B)-Q-BC, US, Eur —Var. oblanceolata (Vict.& Rouss.) Boi- 
vin (A . glabriuscula AA . ) — Terminal spikes conspicuously 
bracted, the bracts mostly entire and lanceolate or oblanceola- 
te. Sea shores. — (G, K, L)^^F, l.S, NB-CKO-nMan, US) —Var. 
LITTORaLIS (L.) Gray — As var. patula, but the leaves narrower, 
i linear, and entire. A coastal variation rarely appearing in- 
land as a weed. — (K), NS41an, BC, (US), Eur. 

As per a tradition now over 200 years old, the larger- 
leaved (i.e. deltoid-hastate) extreme is often segregated as A. 
hastata . It is not clear to us how this distinction facilita" 
tes in any way the intellectual apprehension of this polymorphic 
species . 

U. A. fTgente^ Nutt. — Saltbush, Silverscale — A whitish 
silvery annual wi th i deltoid leaves. Very leafy. Glomerules 
axillary, not forming distinct spikes. First half of summer. 
Open saline soils. — swMan(Melita)-swS-BC, US. 

5. A. Powellii V/atson — Like the preceeding but smaller. 
Bracteoles entire, at least in the upper half. Upper leaves 
more reduced. Mid to late summer. Badlands: Steveville, Ro- 
sedale, — sAlta, wUS . 

6. A. dio ica (Nutt.) Macbr. (Endolepis Sucklej-i Torrey) — 
Rillscale — Staminate glomerules pinkish and forming lightly 
bracted terminal spikes. Pistillate glomerules inconspicuous 
in the lower axils. Leaves lanceolax-e, subacuminate, somewhat 
glaucous, glabrous or nearly so. Early to mid summer. Saline 
flats. — swS-Alta, US. 

6. SUGKLEYA Gray 
Pistillate flowers as in Atriplex but the bracteoles fu- 
sed laterally to the ovary instead of hiding it. 

1. S. Suckleyana (Torrey) Rydb. — Leaves flabellate and 
flabellately dentafeV Somewhat mealy. Diffusely branched and 
resembling Amar anthus albus in habit. Fruit ovate -rhomboid, 
often with a pair of lobes on the angles, bifid at apex. Summer. 
Saline shores, sometimes weedy, but rather rare. — S-seAlta, 
(US). 

125 SUGKLEYA 



66 PHYTOLOGIA Vol. 17, no. 2 

7. EIJROTIA Adanaon 
Pistillate flowers and bracteoles much as in Suckleya . 
Bracteoles with a conspicuous t'oTt of long hair. 

1. JJ. lanat^a (Purah) Moq. — Winter -Fat , White Sage — 
Densely stellate -pubescent throughout. Semi-shruoby in tne man- 
ner of Atriplex Nu ttallii . Dioecious. Leaves linear, revolute. 
Inflorescence - lonR-pilose. Early summer. Dry hilla. — 
swMan (Virden)-Alta, US. 

a. AXYRIS L. 
Staninate flox-zers in a terminal, naked spike of glomerulea. 
Pistillate flowers solitary, axillary. Otherwise resembling 

Chenopodium . 

1. A. AMARANTHOIDES L. — Russian Pigweed — Teminal spi- 
ke conspicuously differenciated, yellowish, and elongate. Other 
spikes much smaller and terminating the branches. Lightly to 
densely stellate -puberulent throughout. Leaves lanceolate. 
Calyx membranous. Mid summer. Frequent weed in disturbed soils, 
invading native habitats in shaded places. — swMack, (NS)-FEI, 
Q^C, US, Eur. 

At times seemingly native, but the earliest Canadian col- 
lection goes back only to 1886. 

9. KXHIA Roth 

As Chenopodium , but the mature calyx developing a peripher- 
al wing or ridge, yet this character not obvious in our only 
species. Not mealy-pubescent. 

1. K. SCOPARIA (L.) Roth (K. trichophila Hort.) ~ Summer- 
Cypress, Burning Bush ( Petit s soTdats , Petits~Pins ) — Very 
branchy and very leafy annual. Densely puberulent witn tufts 
of long hairs in the inflorescence. Leaves linear. Bracts 
very long-ciliate. Calyx glabrous. The whole plant often turn- 
ing red in the fall. Late summer. Cultivated ornamental, fre- 
quent weed of streets, roadsides and waste places. — NS, sQ- 
3C, (US), Eur. 

The weed is perhaps distinct from the cultivated ornamen- 
tal , but we know not how to differentiate them clearly. 

10. 3AS3IA All. 

As Kochia, but the mature calyx developing $ spirally coil- 
ed horns"! However most herbarium specimens are collected too 
early when this character is not yet readily observed. 

1. B. HYSSOPIFOLIA (Pallas) Ktze.-- Rather siJiilar to 
Kochia and easily confused with it, but not so branchy and the 
calyx as densely pilose as any other part of the inflorescence. 
Bracts lacking the long, spreading cilia of Kochia . After 
mid summer. Infrequent weed of railways and roadsides in alka- 
line areas. — swS-3C, US, CA, (Eur). 

EUROTIA 126 



1968 Boivln, Flora of Prairie Provinces 6? 

11. GORISPERMUM L. 
Flower much reduced, with only l-(2) stamens and the calyx 
reduced to 1 sepal. 

1. C. hvssqpifolium L. var. hyssopi folium (C. marginale 
Rydb.; C. simplicissimum Lunell) — Bud-Seed, Tick-Seed — Flo- 
wers not in glcxnerules, but solitary in the a.d.1 of large bracts. 
Very branchy and glabrous to stellate -pubescent, not mealy. In- 
florescence a terminal spike, rather dense and the bracts hiding 
the fruits. Seed discoid, with a peripheral wing 0.3-0.6 mm wi- 
de. Mid summer. Loose sands. — Mack-(Y^ka), Q-Alta, US, (CA), 
Eur — Var. rubricaule Hooker (C . nitidum Kit.) — Spikes not so 
dense. Bra ctssmaTler , 1-3 mm wide, mostly narrower than tne 
fruits. ~ wO-S-(Alta-BC), US, Eur ~ Var. emarginatum, (Rydb.) 
Boivin (C. orientale Lam. var. emarginatum ^Rydo . ) ' Ma cbr . ; C. 
villo3um~ Rydb . ) — Seed fairly large, 3-U mm long, and merely 
sharp-margined , without a marginal wing. — swQ-Alta-(BC, US, 
Eur). 

Within our range our three varieties present themselves 
like mere extremes of variations, but in Eurasia their ranges 
appear to be highly individualized. 

12. SALICORNIA L. GUSSWORT, SAMHiIRE 
Fleshy plants with vestigial leaves. Flowers in 3's and 
more or less embedded in a depression of the next internode 
above. Calyx fleshy. Stamens only l-(2). 

1. S. eurgpaea^ L. var. prona (Lunell) Boivin (3. rubra 
Nelson) — Sand^ire , Gla s swortt cbra il , Passe-pierre) — Small 
herb reduced to its fleshy stem and branches, often turning red 
in late summer. Annual. Intemodes swollen into joints. Each 
joint with a membranous -margined collar at the upper end. Flo- 
wers inconspicuous, in terminal spikes of opposite glomerules. 
Mid summer. Saline shores. — sMack-Y-(Aka), Man-BC, US- 
All the inland material belongs to our variety in which 
the stem internodes pass abruptly into the much shorter inflo- 
rescence intemodes, the latter usually 1.5-2,5 mm long. Upper- 
most stem internode generally more than twice longer than the 
lowermost inflorescence internode. In the East Coast and Old 
World var. europea the spike is less strongly contrasted and 
its intemodes are mostly (2)-U-(5) mm long; the uppermost stem 
internode usually less than twice as long as the adjacent spike 
internode. 

13. SARCOBATUS Nees GRAESE'iOOD 

Staminate flowers in catkins which show a marked similar- 
ity to the spikes of Equisetum, each flower being reduced to 3 
stamens and a stipitate, peltate scale. Pistillate flower so- 
litary, axillary. Fruit with a broad horizontal and circular 
wing. 

127 SALICORNIA 



68 PHYTOLOOIA Vol. 17, no. 2 

1. S. vermiculatus (Hooker) Torrey — Oreasewood, Pulpy 
Thorn — very spiny ahrub f^rowiriK in large colonies. Young 
branches pale to whitish. Leaves fleany, linear, alternate aoo- 
ve to opposite or verticillate below. Early summer. Hitfily al- 
kaline flats at the oottcm of the major coulees. — svS-seAlta- 
seBC, US. 

lU. SUAEDA Forsk. S£A .BLUE 

Flowers in axillary glomsrules of 3. Calyx flesny. Otner- 
wise resembling Chenopodium . 

1. S. maritima (L.) Dum. var. maritima — Seablite (Blan- 
chette, ^languety^- Annual herb withT^strong tendency to turn 
dirty black during the second naif of summer. Very oranchy. 
Leaves linear, fleshy. Bracts much as the leaves, 1.0-1.5 nm 
wide, oblong to linear, of uniform width, but snorter than tne 
leaves. Mid summer to early fall. Seashores. — (Mack-Y)^Aka , 
NS-Q, nMan, wBC, US, Eur — Var. americana (Pers.) Boivin (S. 
depressa (Pursh) Watson; S. erecta (Watsonj Nelson) — Bracts 
more sharply dif f erenciated from the leaves . Lower leaves * 1mm 
wide, linear of uniform width. Bracts mucn snorter, l,$-3.0 mm 
wide, at the base, ovate to narrowly triangular -lanceolate, gra- 
dually narrowed from the base. Alkaline shores, sonetimes 
weedy. — seK-Y, (NF), NS^C, US. 

The more southern S. intermedia Watson nas reported frcm 
Alberta by Hitchcock 19'5U, but this may have been only a lapsus 
calami as we nave been unable to substantiate this report. 
There was no justifying sheet at WTU in 1967 and there was no 
specimen under that name in any of the herbaria visited. A sys- 
tematic review of all tne Saskatchewan and Alberta sneets of 
Suaeda at DAO in 1967 failed to turn up any S. intermedia mas- 
querading under another name. 

1$. SALSOLA L. SALTWCRT 

Flowers as in Chenopodium , but with 2 bracts. Fruit deve- 
loping a circular horizontal wing as in Cycloloma and Sarcoba - 
tus . 

1. S. KALI L. var. TENUIFOLIA Tausch (3. pestifer Nelson) 
— Russian Thistle ( Chardon de Russie] — An- 

nual herb~at first soft and fleshy, soon hardening into a bun- 
dle of norribly spinescent foliage. Very branchy. First lea- 
ves filiform, and soft, the later ones and the bracts shorter 
and ending into a whitish, stiff and very sharp point. Flower 
axillary, solitary, subtended by 3 bracts, i.e., the foliage 
bract and the 2 floral bracts. Mid summer to frost. Very com- 
mon weed of bare or disturbed soils, seemingly native on eroded 
dunes. — NS-BC, US, Eur. 

Typical var. Kali is native along the East Coast and in 
the Old World. Its leaves are shorter, the main ones not over 
3 cm and usually not over 2 cm; they are also as thick, stiff, 
and spinescent as the shorter and later leaves. 

SUAEDA 128 



1968 Boivin, Flora of Prairie Provinces 69 

79. AI11PANTHACSAE (AMARANTH FAMILY) 
Each flower subtended by a scarious bract and 2 scarious 
bracteoles. Otherwise similar to the Chenopodia ceae . 

1. amaPu\:;thus l. amaranth 

The basic genus of the family, with alternate leaves and 
the calyx present. 

a. Spiny in the leaf axils 6. A. spinosus 

aa. Not spiny. 

b. Flowers in small axillary inflorescences. 

c. Seed about 1.$ mm wide U« A. blitoidea 

cc. Smaller, slightly less than 1 mm wide. 

d. Bracts and bracteoles 2-3 mm long... 3. A. albus 
dd. Shorter, less than 2 mm long .. 

5» A. calif omicus 

bb. Terminal inflorescences present, larger 
and conspicuous. 
e. Spike -like inflorescences lax and moni- 
liform, at least in the lower half .. 

7. A. tuberculatus 

ee. Spike or panicle dense throughout or 
essentially so. 
f . Bracts 2-3 mm long, only slightly 

longer than the calyx 1. A. hybridus 

ff . Bracts 3-8 mm long, much exceeding 

the calyx 2 . A . retroflexus 

1. A. HYdRIDUS L. var. HYBRIDUS (A. cruentus AA.j A. du- 
bius Mart.) — Pilewort, Pigweed (BrMe de Malabar) — Glcoieru- 
les in numerous, narrow, elongate spikes, usually less than 1cm 
wide. Flowers and bracts small, otherwise similar to the fol- 
lowing. Inflorescence green. Late summer. Sometimes cultiva- 
ted and casually escaped: Winnipeg, — Q-Man, (US, CA), SA, 
Eur, (Afr, Oc) —Var. CRUENTUS (L.) Moq. (var. h^ijochondriacus 
(L.) Baileyj A. paniculatus L. ) — Prince's Feather, Love-Lies - 
Bleeding ( Cannes , CordeliSre) — Inflorescence red. Fort Sas- 
katchewan "^^^"Q^, cAlta-(BC, US). 

Our only sheet of var. cruentus was reported as var. hypo - 
chondriacus by Groh 19U9. 

T. aT retroflexus L. var. RETROFLEXUS — Red Root, Pig- 
weed (Herbe grasse) — The taproot commonly reddish . A stiffly 
erect annual with large oval leaves and a dense greenish pani- 
cle. Villous, especially above. Calyx lobes obtusn or rounded, 
commonly erose, often mucronate. Mid summer. Common weed of 
open soils and cultivation. — Mack, (Aka, NS-NB)-Q'-0-(Man)-S- 
BC, US, (CA), Eur, (Afr) —Var. FSEUDORETROFLEXUS (Thell.) Boi- 
vin (var. Powellii (Watson) Boivinj A. Powellii Watson) — Ca- 
lyx lobes acute to acuminate. Not so densely villous, someti- 
mes nearly glabrous. Inflorescences tending to be less thick 
and not quite so dense. Native further south, but only a rare 
weed with us: Melfort, Lethbridge. — PEI, 0, cS-BC, US, (CA, 
SA, Eur). 129 AMARiu\'IHUo 



70 PHYTOLOGIA Vol. 17, no. 2 

Var. peeudoretroflex\je (Thell.) stat. n., A. chloroetachya 
W. var. pgeudoretrof lexus~ f hell . , ViertelJ. NatT Ges . ZQrich 52: 

hh3, 190T. ^ ^ 

3. A. albugo L. var. albug — Tumbleweed (Fleur de jalousie) 
— A bushy tumCleweed resemCTinp the following, but t^ie leaves 
gradually decreasing Jn size from ti-ie base up. Branchy with a 
well defined main axis whicn is more or less erect. Glabrous 
or sparsely puberulent. Mid sunmer to early fall. Sandy soils, 
Bometijnes weedy. — NS-BC, US, Eur. 

The more southern var. pubescens (Uline & Bray) Fern, is 
viscid -puberulent . 

U. A. BLITOIDES Watson (A. graecizans AA.) — Matweed — 
A carpet weed with the leaves conspicuously dimegueth. Stem 
usually indistinct, but the many branches more or less spread 
out flat on the ground. Leaves obovate, usually retuse, tnose 
of the main branches all about the same size, coranonly 2-5 cm 
long including the petiole, those of tne secondary branches only 
half as large. Summer. Common weed, tolerates trampinp, prefers 
bare soils. — (Aka), swQ-BC, US. 

5. A. CALIFORNICUS (Moq.) Watson — Similar to tne proceed- 
ing, but generally smaller. Leaves only half as large. Seeds 
small, like those of A. albus . Mid to late summer. Rare road- 
side weed: Cypress Hills, Calgary, Herronton, Manyberries. — 
swS-sAlta, wUS. 

6. A. SPINOSUS L. — Careless Weed (Eplnard rouge, Epinard 
epineux) — Most leaf axils bearing a pair of snarp spines about 
1 cm long. Erect annual. Leaves ovate. Spikes thin and elon- 
gate. Mid to late summer. Rare and evanescent weed, collected 
once at Fort Garry. — swO-^an, US, Eur. 

7. A. tiiberculat,\j^s (Moq.) Sauer — (A. tairiariscinus Nutt.; 
Acnida tamarisciiia^ Nutt . ) Wood) — Dioecious"! Erect annual. 
Leaves narrowly ovate to lanceolate. Glojnerules in numerous, 
very thin, elongate and moniliform spikes. Mid summer. Sandy 
shores: Souris River. — swQ-O-(sMan), US. 

Order 15. PRIMUTJiT.ES 
Calyx and corolla fused. Stamens opposite the petals. Flo- 
wer repTolar. In nearly all other groups the stamens are either 
more numerous than the corolla lobes or alternate with them. 

a. Style 1 80. I>rimulaceae 

aa . Styles 5 81 • Plumbaginaceae 

80. FRIMULACEAE (PRD-IROSE FAMILY) 
Herbs with opposite or verticillate leaves and a dry fruit. 

a. Leaves all basal except sometimes for an invo- 
lucre subtending tne inflorescence. 

b. Flower solitary 2. Douglasia 

bb. Flowers in an umbel. 

c. Corolla lobes elongate, sharply 

ref lexed U • Dodecatheon 

AMARANTHUS 130 



1968 Boivin, Flora of Prairie Provinces 71 

cc. Lobes ascending to spreading. 

d. Calpc shorter than the tube of 

the corolla 1. Priinula 

dd. Calyx as long or longer 3« Androsace 

aa. Stem leafy. 

e. Upper leaves alternate 9. Centunculua 

ee. All leaves opposite or verticillate . 

f. Flowers nearly sessile in the axils 7. Glaiix 

ff. Flowers pedicellate. 

g. Leaves borne in a single 

verticil 6. Trier.talis 

gg. Leaves boime at more than one 
node. 

h. Corolla yellow 5. Lysimachia 

hh. Brick-red 8. AnagalHi 

1. PRIMULA L. PRIMROSE, CCfWSLIP 
Flowers ^-merous in an umbell. Leaves all basal. Stamens 
borne on the upper third of the cylindrical corolla tube. Co- 
rolla lobes bilobed. 

a. Yellowish or whitish farinose on the calyces 

and lower leaf surfaces 2 . P. incana 

aa. Green or only slightly farinose. 

b. Leaves entire U. ?• egaliksensis 

bb. Leaves dentate or crenate; flowers larger. 
c. Pedicels many times longer than the 

bracts 1. P. mistassinica 

CO. Not more than twice as long at flower- 
ing time 3« ?• strict a 

1, ^ J2i,Sj;£x§i2i££ M^* var. mis t a^ s ini c a (P. MacCalliana 
Wieg.) — Bird's Eye, Primrose — Small and usually less than 
12 cm high. Leaves denticulate, mostly obovate. Bracts 2-6ram 
long, flat at base. Pedicels up to 3 cm long. Flowers white 
to mauve, commonly 1 cm across. Late spring and early summer. 
Bogs, shores and wet rocks. ~ K^^Iack-(Y-Aka, L)-i«fF-(SFM), NS, 
N3-BC, US, (eEur). 

The leaves are green in our variety, but yellowish farino- 
se below in var. intercedens (Fern.) Boivin, a plant similarly 
small, yellowish farinose on the calices, ma gni lacustrine in 
its distribution, 

P. borealis Duby, a minor segregate of P. mistassinica , 
was reported from as far north as Banks Island by Hulten 1$'U8, 
Anderson 19U9 and Simmons, "A Survey of the Phyto geography of 
the Arctic Archipelago, Lunds Un. Arskr. 1^: I-I63. 1913," but 
this has never been confirmed and may have been based on a spe- 
cimen of P. stricta, the only Primula species otherwise known 
to occur In the Franklin District. Hence the restricted range 
accepted above. 

2. P. in cana. M.E. Jones (P. farinosa AA.) ~ Larger and 
the calyces "and^Tower leaf surfaces densely farinose. Mostly 

131 PRIMULA 



72 PHYTOLOGIA Vol. 1?, no. 2 

2-U dm htpti- Leaves dentate, oblanceolate. Early summer. Mar- 
shy places. — (Mack-Aka, nwQ), Man-Alta-(BC ), wU^. 

3* -£• SiS^i2^ Horn. — Somewtiat coarser than P. mistassi - 
nica, but thtTTIowera smaller. Mostly 1-3 dm hi^h. Leaves obo- 
vate to lanceolate. Bracts saccate at base. Flowers somewnat 
less than 1 cm across. Early swnmer. Wet places in arctic and 
subarctic habitats. — (G-F)-K-Mack-(Y-Aka, L), Q-Msn, (Alta-BC, 
wUS), Eur. 

li. p. egaliksensis Wormsk. — Resembles P. mistassinica , 
but the leavesentTre''and broadly obovate to spatulate. Flowers 
less than 1 cm across. Early summer. Arctic shores and mar- 
shes. — (G), sK-(Mack-Y)-Aka, (L)-NF, Q-nMan, (Alta)-3C. 

2. DOUGIJISIA Lindley 
Flowers as in Primula, but the corolla lobes are entire. 

1. D. montana Gray — Cushion-forming perennial with the 
general pTesentation of Silene acaulis . Leaves thick, ciliate. 
Peduncle stellate -pubescent. Flower pink to white. Early sum- 
mer. High alpine on rocky ridges and scree slopes: Waterton. 
— swAlta, wUS. 

Reported by Hitchcock 1959 as "Waterton Lakes, 3.G.," an 
obvious lapsus calami for "Waterton Lakes, Alta" . The B.C. re- 
port by Taylor 1966 may be based on the above lapsus, as there 
was no corresponding B.C. specimen at UBC in 1966. 

Douglasia nivalis Lindley is known to occur only in the 
mountains of the state of Wasaington except that the type col- 
lec"oion is supposed to come from the Canadian Rockies, hence 
the frequent reports fran Alberta and B.C. Lindley describes 
the type locality as follows in Edin. Bot. Reg. 22: 1886. I836: 
"Upon his journey across the rocky mountains in April 1827, in 
latitude ^CN., longitude llQ"\i., at an estimated elevation of 
12,000 feet above the level of the sea, the attention of Mr. 
Douglas was attracted by a brilliant purple patch amidst the 
surrounding snow. . ." 

Part of the journal kept by Douglas was published in the 
Comp. Bot. Mag. vol. 2 of I836. We learn frcm it that in the 
spring of 1827 Douglas went up tiie Columbia to the junction of 
Canot-Toume river. On April 28 he left the Columbia to strike 
east. On May 1st he climbs Mount Brown (alt. 91$6 ft.) to which 
he assigns an altitude of 16 - I7OOO ft. By May 3rd he has 
crossed the height of land and he is now going down the Athabas- 
ka . There is no suggestion of Douglasia among the plants men- 
tioned in his journal for these few days. 

Considering that Douglasia nivalis has never been collect- 
ed again in the Rockies either of Canada or of tbe U.S.A., and 
despite the circumstancially detailed report by Lindley, we are 
of the opinion that as long as Lindley 's report remains uncon- 
firmed, we must assume an error of locality and date and that 
the type of Douglasia must have been collected witnin the state 
of Washington where Douglas was collecting in I826 and where 
the plant has been collected repeatedly since. 
DOUGLASIA 132 



1968 Boivin, Flora of Prairie Provinces 73 

3. ANDROSACE L. 
Rather similar to Primula, but the corolla tube shorter, 
constricted at the mouth and more or less dilated by the ovary. 

a. Perennial with the flowers much longer than 

the calyx 3« A. Chamaejasme 

aa. Annual with small flowers. 

b. Involucral bracts sessile, lanceolate to 

linear 1. A. geptentrionalis 

bb. Bracts subpetiolate, spatulate or 

obovate 2. A. occidentalis 

1. A. segtentiu^nalis^ L. (var. diffusa (Small) Knuth, var. 
puberulenCa 'J^^dbTjKnutnpVar . subumbellata Nelson; A. puber - 
ulenta Rydb . ) — Like the following, but the bracts narrower 
and broadest at the base. Late spring and early summer. Dry 
places. — (GrAka), NF, Q-(0)-Man-BC, (wUS), Eur. 

2. A. occidentalis Pursh — Inconspicuous annual consist- 
ing mainly of a very leafy rosette and thin and wiry stems and 
pedicels. Stems usually many. Involucral bracts broadest abo- 
ve the middle. Pedicels rather long and uneven. Corolla 
shorter than the calyx. Second half of spring. Li^t and loose 
soils, sometimes weedy. — (wO)-Man-BC, US. 

3. A. Chamae^a^sme Host — Flowers white with a yellow eye. 
Stolonif erous ^erenrnal with solitary scapes. Villous. Pedi- 
cels rather short, not much longer than the bracts. Late spring 
to mid summer. Rocky slopes, montane or alpine. — swF, Mack- 
Aka, swAlta-(eBG, nwUS, Eur). 

U. DODEGATHEON L. AMERICAN COVISLIP 
Flower very showy and rather unusual, resembling an arrow- 
head, with the conspicuous stamens in the point and the long 
reflexed petals as the ears. 

a. Foliage glandular-pubescent 1. D. conjugens 

aa . Folia ge glabrous 2 . D. pulchellum 

1. D. c on jugens Greene var. Beamishii Boivin (var. visci - 
dum AA.; D. cylindrocarpum AA.; D'. pubescens Rydb.) — Flower 
showy, with a rather unusual arrangement of successive colour 
rings. The corolla lobes are bluish -purple; while the tube is 
whitish; the connectives form a yellowish ring and the anthers 
are bluish black below, paler to whitish above. Leaves oblan- 
ceolate. Corolla lobes 10-2^ mm long. Fruit 13-22 mm long, 
circumcissile near the top. Spring and early summer. Montane 
prairies: Cypress Hills and Rockies. — swS-seBC, nwUS — F. 
lacteuin Boivin — Flowers white . — swAlta . 

Var. Beamishii nom. n,, D. pubescens Rydb., Mem. N.Y. Bot. 
Gard.^: 306. 1900. Var. Beamishii is glandular -pubescent, but 
otherwise not different from the more western and glabrous ty- 
pical variety. Miss K.I. Beamish is a student of Dodecatheon 
and herbarium curator at the University of Britisn Columbia. 

133 ANDROSACE 



7U PHYTOLOOIA Vol. 17, no. 7 

Our variety has also been called var. vi,£cidufn but it has Deer, 
pointed out tnat tne type of the lattor nair.e is apparently tne 
hybrid D. conjuyens X Cusic kii. See Bull. Torr. 3ot. Club _b2: 

361, 19??. 

F. lacteujn f.n. floribus alois. Type: D.K. N orris 19, 
Pasque Mtn., UO miles almost due r.orth of Coleman; open prassy- 
rocky slope; flowers wnite, rare, alt. 7500', July 8, 19$6 
(HAO). 

2. D. ouj^neUu": (Raf.) Merr. var. ^^c he llurn (D. Mae^j: a 
AA.; D. rjaucTftorurTTSurand) Gre'^ne; D. radicat'un Greene; t). sa- 
linum~Nelson) ~ SKootinp Star — Similar, 'but glaorous and tne 
flower generally smaller. Corolla lobes ^~lh "w^ long. Fruit 
8-lU mm lon^, openinp by longitudinal slits. Mid spring to 
early summer. Wet places on saline soils. — Mack^Aka, sMan- 
BC, US, (CA). 

Many authors have expressed doubts as to the exact identity 
of D. pauciflorum and D. radicatum . Fortunately, as pointed 
outljy Merrill, Journ. Arn. Arb. 2^: 212. 19Uo, an earlier name 
is available: Eximia pulchella Raf., Aut. 3ot. 16$. 18U0, Tnis 
is based on an excellent illustration and description oy Hooker, 
Curt. Bot. Magr. §j^' 3622. I837 so that the interpretation of 
Rafinesque's name presents no difficulty. Four otner varieties 
occur to the west and soutn of us. Tnese and the typical phase 
are as follows. 

Var. pulchellum — Normally l-3-(U) dm high. Herbage gla- 
brous. Leaves oblanceolate and gradually attenuate at base. 
Filaments yellow. 

Var. Watsonii (Tiri) stat. n., D. Watsonii Tid., Proc. Biol. 
Soc. Wash. 36: I83. 1923 — Smaller than the first and gene- 
rally 2-10 cm hign. Known in Canada only on Mt. Arrowsnith in 
Vancouver Island. A map of the full range of this and otaer va- 
rieties is given by Thompson 19^3. 

Var. album (Suksd.) stat. n., D. Cusickii Greene var. album 
Suksd., V/erdenda 1: 30. 1927; D. Cusickii Greene, Erytnea 3: 
37. 139$ — Like'^he first but the nerbage glandular -pub eruient, 
especially the inflorescence. Known from south-central B.C. and 
the northwestern U.S. 

Var. alaskanum (Hulten) stat. n., D. macrocarpum (Gray) 
Knuth var. alaskanum Hulten, Fl. Aka, Yuk. 3^: 1289. 19li8 — 
Leaves broadest towards the base, ovate to ovate-lanceolate, 
abruptly rounded to a petiole clearly set off from tne limb. 
Occurs along the coast from southern Alaska to northwestern 
Oregon. 

Var. monanthum (Greene) stat. n., D. pauciflorum (Durand) 
Greene var. monanthuro , Pittonia ^: 73- 18 90. Differs from var. 
radicatum by its purple filaments. Tnis would seem to be widely 
distributed in Canada according to a map by Thompson 1953, page 
117, but on closer inspection it appears that the symbols for 
D. radicatum ssp. radicatum and ssp. monanthum have been inter - 
changed and that the latter entity does not occur in Canada. 

DODECATHEON 134 



1968 Boivin, Flora of Prairie Provinces 75 

5. LYSIMACHIA L. LOOSESTRIFE 

A middling type with stamens opposite the petals. Flowers 
yellow. Herbs with opposite or verticillate leaves. 

a. Not flowering, but bulbiferous in the 

axils !• L. terrestris 

aa. Floriferous. 

b. Flowers in racemes. 

c. Raceme open, terminal 1. L. terrestris 

cc. Raceme dense, axillary 2. L. thyrsiflora 

bb. Flowers axillary or in terminal cymules. 

d. Leaves narrowly linear and ses- 
sile 5. L. quadriflora 

dd. Broader and petiolate. 

e. Leaves ciliate, - ovate 3» L» ciliata 

ee. Not ciliate and narrower U. L. hybrid a 

1. L. j;^;:restris (L.) BoP. — Sterile and usually simple 
stems with" reddish axillary bulblets . Much less common than 
the flowering type, not yet collected from Manitoba. — L-(NF, 
NS-PEl)-NB-0, US — F. flS£i£§E2 Boivin ~ Swamp-Candles, Bog- 
Loosestrife — Sepals, petals and fruit with dark purple lines 
or dots. With one or more terminal racemes of long-pedicelled 
flowers. Slimmer. Lake shores. — L-SPM, NS-seMan, US. 

Both forms apoear to have essentially the same distribu- 
tion, but the typical bulbiferous phase was not represented frcm 
Manitoba among the many specimens examined from loans and during 
inventories or revisions. Because this sterile phase is much 
less conspicuous, its lack of representation from our area may 
be due only to lack of collecting. 

2. ^. JhjSlsi^lora ^« ( Na umber gia thyrsiflora (L.) Reich.) 
— Tufted Loosestrife(Corneille en bouquet) — Leaves, stem 
and flowers abundantly and finely purple -dotted. No terminal 
raceme, but the simple stem bearing 2-8 axillary racemes on long 
peduncles. Pedicels shorter than the flowers. Early summer. 
Freshwater shores. — Mack-(Y)-Aka, NS-BC, US, Eur. 

3. L. ciliata L. ( Steironema cilia turn (L.) Raf.) — A com- 
mon and conspicuous yellow-flowered herb with a variable floral 
arrangement, but usually with some flowers solitary in the 
axil of opposite leaves while others are in terminal cymules of 
U-6 flowers subtended by a verticil of U leaves. Long stoloni- 
ferous and without basal rosettes. Leaves mostly 3-$ era wide. 
Peduncle (2)-U-(6) cm long. Mid summer. Light woods and wetter 
prairie spots. — NS-BG, US. 

Gleason 1952 would extend the range to Yukon, but we found 
no corresponding specimen at NY in 196$. 

U. Ji- hybrida Mx. ( Steironema hybridum (Mx.) Raf.; S. lan - 
ceolatum (WalterTTiray var. hybridum (Mx.) Gray) — Readily con- 
fuied with the preceeding, but the leaves not ciliate and nar- 
rower. Not stoloniferous, but producing basal rosettes. Leaves 
0.5-2,0 cm wide, ± lanceolate, usually verticillate on the last 

135 LYSIMACHIA 



76 P H Y T L G I A Vol. 17, no. 2 

2-3 nodes. Flowers all or mootly verticillate. Mid summer. Wet 
meadows. — swQ-wAlta, U3. 

5. L. quadriflora Sims (L. lonp;lfolia Purah; Steironeg.a 
quadrif lormn^ 't^ims^ Hitchc . ) — Leaves linear and sessile. Tuft- 
ed with rosettes, the basal leaves mucn smaller and obovate to 
elliptic. Leaf and flower arrangement mucn as in tne last two. 
Mid summer. Chernozem prairies, rare: Kleefeld. — swO-aeMan, 
US. 

6. TRnMALIS L. CHICKW2ED WlUTKR(SiEEti 
Flower usually T-merous. 

a. Leaves rhomboid -lanceolate, acute to aub- 

acuminate at tip 1. T. borealis 

aa . Leaves oblanceolate to obovate, obtusish 

to rounded at tip 2. T. europaea 

1. T. bgrealis Raf. (T . americana Pursh) — Star - Flower — 
Leaves alt o? mostly in a single terminal verticil. Other lea- 
ves, if any, very much reduced and alternate. Larger leaves 
usually over 5 cm long. Flowers white, terminal, usually two. 
Early summer. Frequent in forests. — (seK), L-SP^I, NS-neBC, 
neUS . 

2, T. eurogaea L. (var. arctica Fischer) — Similar, but 
the leaveT broadest near the tip and usually less than 5 cm 
long. Stem leaves usually present and not so mucn reduced, 
nearly as large as the smaller ones of the terminal verticil. 
— Mack-(Y)^ka, nwAlta-BC, (nwUS), Eur. 

Quite variable as to leaf size and there is a strong ten- 
dency to smaller leaves (var. arctica ) in America. But tnis is 
only a matter of frequency as the range of variation appears to 
be essentially the same on both sides of the Pacific. It seems 
difficult to implement here a distinction that would not be 
either artificial or based primarily on the locus of collection. 

A report of T. lat ifolia Hooker from Alberta by Hitchcock 
19^9 and Boivin 1966 may have been due to a lapsus calami as 
there was no corresponding specimen at VITU in 1967. 

7. GUIIX L. SE.^ MIUC'.'CRT 

Corolla lacking, the calyx somewhat petaloid . 

1. G^. ma rit.ijng L. var. angustifolia Boivin — Black Salt- 
wort (Herbe au lait) — LeavesveryTinery punctate in sli^tly 
darker green. Small perennial herb with milky juice. Somewhat 
fleshy. Leaves mostly around 1 cm long, lanceolate, entire. 
Calyx, marcescent, the lobes pinkish with white margins. Early 
summer. Wettish alkaline soils. — sMack-sY, sMan-sBC, US. 

It is primarily by its narrower leaves that our inland va- 
riety is distinguished from either the east coast (var. obtusi - 
folia Fern.) or the west coast (var. macrophylla Boivin) vica- 
rianEs . 

trientahs 136 



1968 Boivin, Flora of Prairie Provinces 77 

8. ANAGALLIS L. PIMPERNEL 

Sepals free. 

1. A. ARVENSIS L. — Pimpernel, Scarlet Pimpernel (Mouron, 
Mouron rouge) — Flower brick-red. Rather similar to Stellaria 
media in general presentation. Foliage obscurely punctate in 
purple . Leaves ovate, sessile. Peduncle becoming sharply re- 
curved in fruit. Summer. Rare garden weed: Lacombe. — (G, 
NF)^PM, NS-(PEI-NB)-Q-0, Alta-3C, US, Eur. 

9. CEiNTUNCULUS L. CHAF!^//5ED 

Flowers insignificant, U-merous. Leaves mostly alternate. 

1, C. mininnjs L. — Chaffweed — Capsule whitish with a 
brown equatorial line. Small annual with obovate leaves, the 
lowermost opposite. (Mid summer?). Marshy places in the prai- 
rie. Rare or inconspicuous. — NS, S-BC, US, (CA), Eur, 

We have cnecked specimens (DAO) from Mortlach, Long Lake, 
Cory and Einpress. We also know of a report from Reed Lake (CAN). 

81. PLUMBAGINACEAE (LEADWORT FAMILY) 
Plants with the stamens opposite the petals and otherwise 
generally similar to the Primula ce a e but the styles ^ and the 
leaves (and brancning) alternate or basal. 

a. Flowers in a branched inflorescence 1. Limoni um 

aa. In a dense head 2. Statice 

1. LIMONIUM L. SEA -LAVENDER 

Petals free or nearly so. Each flower tightly wrapped in 
(2)-3 scarious bracts. Cal;/x petaloid . 

1. L. VULGARE Miller — Sea -Lavender (Saladelle) — Flo- 
wers in a corymb of secund spikes. Leaves all basal, broadly 
oblanceolate, fairly large. Branching somewhat dichotomous, the 
branches trigonous and winged. Calyx white with $ thick and 
green nerves. Corolla pink. Mid summer. Cultivated and rare- 
ly spreading around old cemeteries: Big Muddy. — sO, scS, Eur. 

Both collections examined (REG, TRT) belonged to the white- 
flowered cv. Album. 

2. APJ-ERIA W. 
Scapose herbs with the flowers in a globose head. 

1. k^. maritima (Miller) W. var. interior (Raup) Lawr. 
( Statice interior Raup) — Thrift, Sea^5inir~(Gazon d 'Espagne, 
Herbe S sept t&tes) — Head subtended by numerous membranous 
bracts, the lowest one being reflexed and tubular. Rosette 
leaves numerous, marcescent and narrowly linear. Head inter- 
spersed by numerous bracts. Early summer. Dunes of lake Atha- 
baska. — (Mack), nwS. 

137 LIMONIUM 



78 PHYTOLOOIA Vol. 17, no. 2 

A variable type to be organized into (reopraphical varietiea 
only with some difficulty. Our present understanding of tne Ca- 
nadian variations may be sxanmarized in tne following key: 

a. Calyx Rlabrous var. interior 

aa. Pubescent at least alon^ the nerves. 

b. Outer involucral bracts triangular- 
lanceolate, - acute at tip, and as 
long or longer than the inner ones — 

Vancouver var. californica (Boies.) Lawr. 

bb. Broader, rounded at tip and shorter. 

c. Outer involucral bracts less tnan 
half as long as tne inner — Arctic 

regions — var. sibirica (Turcz.) Lawr. 

cc. Not quite so short, hence less strongly 
imbricated, 
d. Less than 2 dm high; cai.yx puoes- 

cent on both the nerves and tne 

internerves . — Arctlr -alpine .. 

var. labradorica (Vallr. )Lawr . 

dd. Usually taller; calyx pubescent 

on the main nerves, glabrous on 

tne internerves. — West Coast .. 

var. purpurea (Kert. & Koch) Lawr. 

Order 116. LYTHRALES 
Ovary inferior, but the petals free or lacking. Petals 
borne on the summit of a calyx tube. 

a. Flower vrithout perianth, reduced to a single 
stamen or ovary or both. 

b. Fruit an achene; leaves verticillate. . . Hippuris , p. IhO 
bb. Fruit a diachene; leaves opposite .. 

8$ . Callitrichaceae , p . I)i6 

aa . Flower normal or much less reduced . 

c. Petals more tnan U, usually 6... 82. Lythraceae , p. 138 
cc. Petals (3)-U, rarely lacking. 

d. Fruit an achene. Aquatics .. 
83. Halorrhagidaceae , p. 13*? 

dd. Fruit a capsule. Terrestrial 

plants 8U. Onagraceae , p. lIiO 

82. LYTHRACEAE (LOOSESTRIFE FAMILY) 
Like the Onagraceae, but the floral parts usually more nu- 
merous and the hypantnium (or calyx tube) free frorr. the ovary. 

1. LYTHRUy; L. 
Petals usually 6, free and borne on the summit of the 
elongate hypanthium. 

1. L. SALICARIA L. (var. gracilior Turcz., var. tomento - 
sum (Miller) DC.) — Purple Losestrife ( Salicaire , Roupie de 
ARMERIA 138 



1968 Boivin, Flora of Prairie Provinces 79 

coq d'Inde) — Showy species of snores and ditches with a termi- 
nal inflorescence of magenta flowers. Coarse perennial with op- 
posite lanceolate leaves. Inflorescence a raceme of opposite 
glomerules. Mainly late summer. Sometimes cultivated and 
spreading readily to freshwater habitats. — NF, NS-sMan, swAlta- 
BC, US, Eur. 

83. HALORRHAGIDACEAE (WATKR MITJ'OIL FAMILY) 
Aquatic plants with a rather small or somewhat reduced flo- 
wer, similar to the Onagraceae , but the fruit indehiscent. 

a. Leaves finely divided 1. Myriophyllu m 

aa . Leaves entire 2 . Hippuris 

1. hTYRIOPHYLLUM L. WATER MILFOIL 

Submerged aquatics with verticillate pectinate leaves. 

a. Flowers and bracts all or mostly alternate .. 

1. M. alterniflorum 

aa. Verticillate and the leaves longer. ~ 
b. Inflorescence bracts closely pectinate to 
entire, many times shorter than the leaves .. 

2. M. spicatum 

bb. Bracts remotely lobed and at least half as 

long as the leaves 3« M. pinnatum 

1. JM. alternjjClorum DC . — Leaves smaller than in the fol- 
lowing, (5)-8-10i(12) mm long. Fruit deeply U-lobed, the lobes 
rounded and smooth on the back. Second half of summer. Shallow 
waters, becoming sterile in deeper waters. — G, (Mack, Aka), 
NF-SPM, NS, NB-nMan-nS, US, Eur, (Afr). 

We know of only 3 collections (CAN; DAG, photo) from our 
area: Cochrane river. Reindeer Lake and lake Axis. The last 
is not typical, the leaves being part alternate like the inflo- 
rescence bracts. 

2. M^. s£icatiOT L. (M. exalbescens Fern.j M. verticillatum 
L., var. pectinatum WallrT) — Water -milfoil (Volant d'eau) — 

A common submerged aquatic with verticillate and pectinately 
divided leaves. Leaves (l)-2-(3) cm long. Flowers inconspi- 
cuous, verticillate in a monilifonn and emerged spike. Fruit 
shallowly U-lobed, the lobes rounded and sometimes smooth or 
more commonly somewhat verrucose. Mid to late summer. Common 
submerged herb in shallow to deeper water. — G-(F)-K-Aka, (L- 
SPM), NS-BC, US, (SA), Eur, Afr. 

We are not convinced that the neogean plants are separable 
from the paleogean ones except on a statistical basis . 

3. M. pinnatujTi (Walter) BSP. —Usually with some of the 

leaves or'^f lowers alternate, the otners verticillate. Leaves 

1-2 cm long, the lobes few and rather short, passing gradually 

into the not very reduced bracts. Fruit deeply Ii-lobed, the 

lobes squarish, with 2 tuberculate ridges on the back and 3 

concave sides. Late summer. Submerged in sloughs, rare: 

139 MIRIOPHYLLUH 



80 P H Y T L G I JL Vol. 17, no. 2 

Wordsworth, Mortlach. — aS, US, (CA). 

We have checked only the Wordsworth collection. 

2. HIPPURIS L. MARE'S TAIL 

Palustrine and simple herbs with verticillate and entire 
leaves. Fl'jwers insignificant. Perianth lacking, the ovary 
enclosed by the overgrown hypanthivun. Stamen only 1 or none. 

a. Leaves verticillate in li's 2. H. tetraphylla 

aa . More numerous and narrower T . H. vulgaris 

1. Jj^. VTilraris L. — Bottle-Brush, Mare's Tail (Queue de 
cheval, Fessed^eauT — Common herb of shallow waters with sTm- 
ple stems and verticillate leaves. Stem fleshy. Leaves In 
6's - lO's, entire, 1-3 cm long, acute or acutish, 1-3 mm wide. 
Early summer. Forming large colonies on muddy shores and shal- 
low waters. — G-Aka, L-SFM, NS^G, US, (SA), Eur, (Afr). 

In so far as our two species are shore plants, emerged and 
submerged forms are part of the normal variation of each spe- 
cies and we have made no attempt at distinguishing them, even 
if the submerged forms can be strikingly different. They have 
already received names: f. fluviatilis (Coss. & Germ.) Glueck 
for the first, f. lacunarum Dut. & Lep. for the second. 

2. ji. tetra£h^2i^ L. f . ~ Leaves 0.5-1.0-(1.5) cm long, 
broader, thickish and verticillate in U's-(6'e), oblong-lanceo- 
late and obtuse or rounded at tip. Second half of summer. Ma- 
ritime shores. — (F-K)-Mack-(Y)-Aka, (L), Q-nMan, (BC), Eur. 

8U. ONACiiACEAE (EVENING-PRIMROSE FAMILY) 
Flower U-merous, of free parts, but the ovary inferior, 
being enclosed in a long-tubular hypanthium. 

A Manitoba report of Isnardia palustris L. (» Ludwigia pa- 
lustris (L.) Ell.) is undoubtedly incorrect as pointed out by 
Scoggan ly57 and the Saskatchewan reports by Hooker 1832 and 
Macoun I883 are probably equally unjustified. 

a. Fruit catchy, covered with hooked hairs 6. Circaea 

aa. Not catchy. 

b. Fruit short, indehiscent 5. Gaura 

bb. Elongate, a dehiscent capsule. 

c . Seeds with a pappus 1. Epilobium 

cc. No pappus. 

d. Capsule bilocular, opening by 

2 valves U. Gayophytum 

dd . U-locular and opening by U valves . 
e. Petals entire to merely emar- 

ginate 3« Oenothera 

ee. Petals conspicuously bilobed .. 

2. Boisduvalia 

1. EPILOBIUM L. WILLCW-HERB 

Seed with a pappus of capillary bristles. Otherwise as 
in Oenothera . 

HIPRJRIS IhO 



1968 Boivin, Flora of Prairie Provinces 81 

a. Petals large, at least 1 cm long. 

b. Flowers numerous , subtended by small 

bracts 1. E. angustifolium 

bb. Flowers few in a leafy inflorescence...?. E. latifolium 
aa. Petals smaller. 

c. Leaves linear. 

d. Annual; fruit 2-3 cm long 3« E. paniculattim 

dd. Perennial with longer fruits U. E. palustre 

cc. Leaves lanceolate to ovate. 

e. Low plant with usually ovate to 

elliptic leaves 6. E- alpinum 

ee. Taller, the leaves mostly lanceo- 
late 5. E. ciliatum 

1. E. angustifolium L. (var. intermedium AA., var. macro - 
phyllum (fiaus^rrfFernT^var . platyphyllum (Daniels ) FernT^ 
Chamaenerion spicatujn (Lam.) S.F. Gray) — Fireweed , Pink Tops 
( Lilas de montagne , Bouquets rouges) — Showy virgate herb with 
one large terminal raceme of spreading magenta flowers. Stolo- 
niferous, commonly 1 m high. Leaves ± lanceolate, thin, paler 
and somewhat rugose below. Bracts mostly about as long as the 
pedicels. Flower buds reflexed; flowers spreading; fruits 
slightly ascending. Mid to late summer. Open places, often 
very abundant after a f ire.--G-(F)-K^ka, L-SPM, NS-BC, US, Eur 
— F. albiflorum (Dum.) Hauskn. — Flowers white, including the 
sepals. — Mack-Aka, L-NF, NS-^, US, Eur — F. spectabile 
(Simmons) Fern. — Petals white, but the sepals purple. — Aka, 
NS, Q, Man-S-(Alta), Eur. 

2. E. latifolium L. — River -Beauty — Similar to the abo- 
ve but smaller and somevihat fleshy. Only 1-U dm high. Leaves 
rhomboid to lanceolate, rather thickish, the lateral nerves in- 
conspicuous. Bracts large and leaf -like, mostly at least as 
long as the buds. Flowers (and buds) 2-3-(12), erect. Fruit 
erect. Mid summer. Arctic and alpine habitats, especially wet 
gravels. ~ G-Aka, L-NF, Q-(nO)-nMan, swAlta-BG, US, Eur. 

3. E^. parnxulatmn^ Nutt . (f. adenocladon Hausskn.; var, 
subulatum (HaussknTTFern. ; E. adenocladon (Hausskn.) Rydb.) — 
The bark usually exfoliating~on the lower part of the stem. An- 
nual, usually diffusely branched. Leaves linear, conduplicate, 
falcate. Fruit attenuate at both ends, mostly falcate. Mid 
summer. Shores of sloughs and disturbed soils. — swQ-CB, US. 

U. E^. pal^^st^ L. var. palustre (var. grammadophyllum 
Hausskn.; var. monticola AA., var. oliganthum (Mx.) Fern.; E. 
davuricurn Fischer; E. densum Raf.; E. leptophyllum Raf,; E.~ 
lineare ~AA . ; E. molTe Torrey; E. oliganthum Mx.; E. strictum 
Muhl.; E. wyomingense Nelson )~ — Resembling the next, but the 
leaves narrowly linear and the flowers usually white. Glabrous 
to grayish pubescent. Leaves less than 5 mm wide. Perennial 
by thin, fragile stolons. Fruit 3-7 cm long. Mid summer. 
Swampy ground. ~ (G-F)-K-Mack-(Y^ka ), L-SPM, NS-BC, US, Eur. 

Somewhat variable and subjected to much splitting. We 
have accepted the consolidation proposed by Hitchcock 1961 as 

Uil EPILOBIUM 



82 PHYTOLOQIA Vol. 17, no. 2 

it aeema realistic. The next two apeciet aro ijiao tne reeult of 
similar consolidation procedures. 

On the east coast tnere is a var. sabulonerise (Fern.) Boivin 
with larger flowers, the petals 8-10 mm lon(^. 

5' j^« Siii£k>© Raf« var. siii^iijii (^* adenocaulon Haueskn., 
var. perplexans Trel.; E. americanum Hausskn.; E. Drummondii 
Hausskn.; E. glandulosum Lehjn., var. adenocaulon (Kausekn.) Ferxi., 
var. cardiophyllum Fern., var. Macounii (Trel.)~C.L. Hitchc, 
var. occidentale (TreljFem., var. tenue (Trel.) C.L. Hitcnc; 
E. leptocarpum Hausekn., var. Macounii Trel. ; E. saximontanum 
ffausskn.; E. scalare Fem.j E. Steckerianum Fern.j E. Wateonii 
Barbey) — A common middling type, 2-8 dm hi^. Perermial by 
fragile stolons. Leaves 0.^-2.0 cm wide, lanceolate, denticula- 
te. Fruits and flowers erect, the latter usually pinkisn or 
mauve. First half of summer. Wet ground. — (Mack)-Y-(Aka) , 
L4JF-(Sm, NS-PEI)-NB-BC, US, (Eur). 

The absence of pappus cnaracterizes an eastern endemic, 
var. econosum (Fassett) Boivin, known only from the estuary of 
the Saint Lawrence. 

Earlier reports by Hooker I832 and Hacoun I883 of E. colo- 
r a turn Muhl. were based on specimens wnich, according to""Macoun 
l89U, were mostly revised by Trelease to E. adenocaulon. Con- 
sidering the absence of E. coloratum from~We£tern Canada, a si- 
multaneous report by Macoun lb9U of the hybrid E. coloratum X 
adenocaulon from Little Slave Lake cannot be rated as anything 
but highly improbable . 

6. JE. alpinum L. (var. albiflorum (Suksd.) C.L. Hitchc, 
var. clavaturr XTreT. ) C.L. Hitchc, var. gracillimom (Trel.) C. 
L. Hitchc, var. lactiflorum (Hausskn.) C.L. Hitchc, var. nu- 
tans (Horn.) Hooker; E. anagallidifolium Lam.j E. glaberrimum 
Barbey var. f astigiatum (Nutt.) Trel.; E. Hornenan HIi Rchb.; E. 
lactiflorum Hausskn.; E. platyphyllum Rydb.) — LiTce'tne pre-" 
ceeding but smaller anH perennial by rooting decumbent bases or 
superficial stolons. Only l-2-(U) dm high. Leaves ovate to 
narrowly oblong, rather few and commonly only 3-U pairs to a 
stem. Flowers few, usually pinkish or nauve. Mainly mid sum- 
mer. Cold mountain springs. — (G-F)-K-(Mack-Y)-Aka, L-(NF, 
NS), Q, Alta-BC, US, (Eur). 

Re E. minutum Lindley reported for northern Alberta by 
Macoun IB83, see comment about Rosa nutkana p. 65, part I. 

2. BOISDUVALIA Spach 
Petals bilobed, othei'wise as in Oenothera. 

1. B^. Si£^®iiS. (Nutt.) Walpers — Inconspicuous annual. 
1-2 dm high, usuali^^ decumbent and ± branched from the base. 
Herbage more or less hirsute. Leaves narrowly lanceolate below 
to broadly lanceolate above. Fruit often curved, somewhat 
shorter than its leaf -like bract. Mid summer. Bare alkaline 
clays, rare. — swS-BC, US, (SA). 

A collection of 3. densiflora (Lindley) V/atson labelled 
M.O. Malte, Alberta, Lethbridge, Aug. 27, 1911 (CAN; DAO, photo) 

EPILOBIUM lli2 



1968 Boivin, Flora of Prairie Provinces 83 

was mentioned by P. Raven in Brittonia 12' 250. 1965 and was the 
basis for the Alberta entry in Boivin 1966. The accuracy of the 
locality on the label was questioned by Raven and his doubts 
proved to be fully justified. We did not locate Malte's field 
records for that year, but a checking of other herbarium sheets 
at DAO showed that in late August 1911 Malte was collecting in 
British Columbia, not in Alberta. A similar check by Miss H. 
Harkness at the National Musei:im neatly confirmed and completed 
our sampling. The consolidated samplings provide us with the 
following spot-check on Malte's 1911 itinerary: 

Aug. 7-8, 1911 ~ Fernie, B.C. 

Aug. 11 Nelson, B.C. 

Aug. 15 Salmon Arm, B.C. 

Aug. l6 Kamloops, B.C. 

Aug. 20-21 Vancouver, B.C. 

Aug. 24 Victoria, Cedsr Hill, B.C. 

Aug. 27 New Westminster, B.C. 

Aug. 31 SujTimerland , B.C. 

Sept. 3 Banff, Alta. 

Sept. 5-6 Calgary, Alta. 

In all likelihood the collection labelled Lethbridge came 
from the vicinity of Victoria, B.C., the only area where B. den - 
siflora is known to occur in Canada. 

3. OENOTHERA L. EVENING-PRIMROSE 

A basic type, 4-merous and the perianth of free parts, but 
the ovary inferior. 

A very heterogeneous genus comprising 15 subgenera many of 
vrhich are rated as distinct genera by various authors. We have 
found the treatment by P. A. Munz, N. Am. Fl. TI , 5: 79-177.1965 
to be the most practical solution, while being intellectually 
as satisfactory as any other arrangement known to us . 

a. Stemless or the stem rather short, overtopped by 
the basal leaves . 

b. Flowers very large, white 8. 0. caespitosa 

bb Smaller and yellow. 

c. Petals 1-2 cm long 7. 0. flava 

cc. Shorter, 6-10 mm long 9. 0. brevi flora 

aa. Stem much taller than the rosette leaves. 

d. Petals white, fading purplish 2. 0. Nuttallii 

dd. Petals yellow. 

e. Petals 1-3 ram long 6. 0. andina 

ee. Petals 5 ram long or more. 

f . Ovary and capsule roiinded on the 
angles . 

g. A low shrub 3. 0. serrulata 

gg. Biennial herb 1. 0. biennis 

ff . Ovary and fruit winged or crested on 
the angles . 

143 OENOTHERA 



8U P H y T L G I A Vol. 17, no. 2 

h. Petals 5-9 nrni lonf 5. 0. perennis 

hh. Larper, 10-25 nun 4. 0. fruticosa 

1. jg^. Jsigncig L» var. ,bi^nnjs — Lvenini^-Primrose, Candle- 
stick (Herbeauxanes, Mache roupe) — Larpe yellow flowers in 
the shape of a maltese cross, biennial herb, p:reen, more or 
less pubescent. Leaves lanceolate, entire to remotely denticu- 
late. Flower borne at the end of a loni^ thin tube, termed hy- 
panthiuin, longer than the ovary and enclosinp it. Mid to late 
summer. Pioneer in open soils, — (NF, NS-Nt)-Q-(0-Man)-S-&C, 
US, (Eur) — ^ • JEJiJiiSS^^ ( L • ) Eoivin (0, muricata L. ; 0. parvi - 
flora L. ) — Pubescence partly of stiff hairs with a red and 
inflated base. — (NF, NS-NB)-Q-0-(Man-BC, US) — Var. cajTescens 
T. 8r G. (var. hirsutissima Gray; 0. striposa (Pydb.) Mack. * 
Bush) — More pubescent, prayish or whitish hairy, especially in 
the inflorescence. Muricate hairs none or few. — (NS-O)-Man- 
Alta-(BC), US, (CA). 

In the east it has been minutisected into umpteen micro- 
specie-^ as the result of genetic studies. Fortunately our local 
populations have remained completely outside these developments 
towards the miniaturization of the species concept. 

2. jQ^. Nuttallii Sweet (0. pallida AA.; Anogra Nuttallii 
(Sweet) N els on)--^ Stem bone-white. Tufted perennial. Leaves 
linear. Flowers large and showy, opening white in late after- 
noon, fading pink, drying reddish blue. Mid summer. Scattered 
tufts on sandy soils. — 0-BC, US. 

3. ^. serrulata Nutt. (Meriolix serrulata (Nutt.) Walp. — 
Shrubby in the lower half. Leaves lanceolate to linear, cons- 
picuously serrate, tending to be conduplicate and falcate, Fmit 
"lirH^r. Summer, Prairie on sandy or gravelly soils. — (C)- 
Man-Alta, US. 

4. 0. Ft-.UTICOSA L. (var. line aris (Mx.) Watson) — Sun- 
drops — Le^.'>''eE alternate, becoming congested in the inflores- 
cence. Tufted perennial. Fruit ellipsoid, stipit^te. Early 
siommer. Rare weed of gravelly soils: Bird's Hill, — (sMan), 
eUS, 

5. 0. perennis L. (0. pumila L.) — Sundrops — Fruit cons- 
picuously st"iprtateV Generally similar to the preceeding,but 
the flovxers smaller and the inflorescence racemose. Early sum- 
mer. Prairies on gravelly soils, rare: Teulon. — NF-(SPK), 
NS-0-(Man, swBC, eUS). 

6. 0. andina Nutt. var. andina — Small annual with minute 
flowers. >lround 1 dm high and very branchy. Fruit largest at 
the base and gradually, tapered. Early sujnmer. Light soils, r^- 
re: Pend-d' Oreille. — sAlta-(sBC), wUS. 

In var. Hilgardii (Greene) M\mz from the state of Washington 
the petals are about twice longer. 

7. 0_. fl^vj^ (Nelson) Garrett ( lavauxia flava Nelson) — 
Similar to tliefoll owing but generally smaller and the flower 
yellow when fresh. Pubescence somewh-t shorter and less dense. 
Petals 1-2 cm long, fading purplish. Anthers 4-8 mm long. Hy- 

OENOTHEEA IW 



1968 Boivin, Flora of Prairie Provinces 85 

panthium and sepals finely glandular. Capsule slightly hirsute 
and finely glandular, the pnfTes not verrucose and not particu- 
larly sinuous. Early summer. Steppes and eroded hillsides, — 
sS-sAlta, (US, CA). 

8. 0. caespitosa Nutt. var. caespitosa (var. montana 
(Nutt.) Eharand; Pachylophus caes pitosus ^Nutt . ) Raim.; P. rnon- 
tanus (Nutt.) Nelson) — Showy perennial with huge white flowers 
fading pink or red. Stemless with rosette leaves resembling 
those of a Taraxacum . Petals 2,5-4.5 cm long. Anthers 8-13 ™" 
long. Hypanthium and sepals strigose. Capsule strigose or gla- 
brescent, strongly sinuose- verrucose on the angles. Early sum- 
mer. Bare clays and badlands, local. — sS-sAlta, wUS — Var. 
£sanraiophila (Nels. & Macbr.) Munz — Stem present, about 1 dm 
long. More restricted: Cardston. — swAlta, nwUS. 

Var. m.ontana is apparently only a less common glabrous ex- 
treme, sporadic in the range of the typical pubescent phase. 

9. 0. .ijreviflQra^ T . & G. (C. breyi folia sphalm. ; Taraxja 
breviflorT ( tTT'gT) Nutt . ) — Like the previous 2 but the lea- 
ves more deeply divided, ]yrately pinnatipartite, and the flo- 
wers smaller. Puberulent throughout, including the sepals, hy- 
panthium and capsule, the latter merely rounded on the angles. 
Petals yellow, 6-10 mm long, fading reddish. Anthers less than 

1 mm long. Early summer. Saline clay flats, rare. — swS-sAlta- 
sBC, US. 

4. GAYOPHYTUM Jussieu 
Capsule bilocular and opening by 2 valves. Otherwise as 
in Oenothera , 

1, G. humile Juss . (G. racemosum T. &. G.) — Capsule deeply 
sulcate on both faces. Inconspicuous and small annual, somewhat 
puberulent. Leaves linear. Capsules linear. Mid summer. Dis- 
turbed sandy ground, rare: Mt. Glendovm. — swAlta, wUS, (SA). 

Closely related to, and none to clearly distinct from, the 
more western G, ramosissimum Nutt. 

5. GAURA L. BUTTEKFLY WEED 

Fruit short and indehiscent. Otherwise as in Oenothera . 

1, &. coccinea (Nutt.) Pursh var. coccinea — Fruit rhom- 
boid. Tufted perennial with decumbent stems and terminal race- 
mes. Herbage pubescent and tending to be grayish, especially in 
the inflorescence. Flowers pinkish in bud, darkening and fading 
deep scarlet. Early to mid summer. Common on hillsides, dry 
prairies, roadsides, etc, — O-Alta-(BC), US — Var, glabra 
(Lehm.) T. & G. (G. glabra Lehm.) — Glabrous or nearly so. Less 
frequent and of more restricted distribution. — S-Alta, US. 

6. CIRCAEA L. ENCHANTER'S NIGHTSHADE 
Floral partsin 2's. Fruit catchy by hooked hairs. 

a. Fruit broadly oblanceolate 1. C. alpina 

li*5 GAURA ~ 



86 P H Y T L G I A Vol. 17, no. 2 

aa. Broadly obovoid 2. C, quadrisulcata 

1. _g^ alpina L. (C. paci fica Asch. ft Kapnus) — A delica- 
te forest species with small catchy fruits in terminal racemes. 
1-4 dm hiph. Leaves broad, ovate, remotely denticulate, haceme 
minutely and obscurely bracteolate, the bractlets mostly 0.1-0,3 
mm lonp. Flowers small, white. Petals ± 1 mm lonp. Fruit ± 

1 mm wide, not ridred. Early to mid summer. Common in damp fo- 
rests. — (Nack), Aka, L-SPM, NS-BC, US, Eur ~ Var. P«cificn_ 
(Asch. & Mapnus) M.E. Jones — Raceme bractless excepTsometirnes 
the lov;ermost l-(3) flowers. Rockies. — swAlta -EC, wfJS. 

Specimens of var. paci fica will commonly exhibit a number 
of other characters such as being taller and hevinp leaves not 
cordate at base and less saliently toothed. Distinctions based 
on these additional characters have proved rather unsatisfactory 
as a certain proportion (about one in ten) of more eastern spe- 
cimens will also exhibit these same features in a sporadic 
fashion. We have therefore shifted the emphasis entirely to 
the presence or absence of bractlets in the inflorescence, a 
character more clearly restricted in its geography. 

2. jC^. 9ijadrisulcata (Max.) Franch. & Sav. var. canadensis 
(L.) Hara — Rachis of the raceme purplish at the base of each 
pedicel. Like the preceeding, but larger throughout. 3-8 dm 
high. Petals 1 2 mm long. Fruit 2_3 mm wide, with 6-10 longi- 
tudinal ridges. Summer. Alluvial woods on the Coteau de Prai- 
rie. — (NF), NS, MB-sMan, US. 

In our variety the flowers are reputedly less brightly co- 
loured and less pubescent than the typical east-asiatic plant. 

85. CALLITRICHACEAE (WATER-STARWORT FAKELY) 
Flower insignificant, without perianth and reduced to an 
ovary or a single stamen. 

1. CALLITRICHE L. HATER STARWORT 

Submerged aquatics with submerged flowers. 

a. Leaves all alike; fr\iit larger 2. C. hermaphroditica 

aa. Leaves \isually dimorphic; fruit smaller 1. C, palustris 

1. Z, palustris L. (C. heterophylla AA. ; C, vema L.) — 
Submerged aqua^dcwith opposite and entire leaves, the latter 
usually dimorphic. Submerged leaves filiform, 1-nerved and xisu- 
ally about 2 cm long. Floating leaves smaller, — spatulate, 
3-nerved, the nerves reticulate. Fruit longer than broad, 
1.0-1.5 mm long, shallowly svilcate, the angles very sharp to nar- 
rowly winged. Summer. Common submerged aquatic. — (G), K-Aka, 
L-SPM, NS-BC, US, (SA), Eur. 

We have examined and revised to C. palustris two (DAG, OT) 
of the three Manitoba collections listed as C. heterophylla 
Pxirsh by L8ve 1959. The other collection was not seen. 

Macoun 1890 also reports C. heterophylla from the Koose Jaw 
Creek but there are no Saskatchewan specimens filed under that 
name to-day at CAN and the original collection has presum=.bly 
been revised since to some other species, possibly C. palustris . 
CIRCAEA 146 " 



1968 Boivin, Flora of Prairie Provinces 8? 

2. C^. tiermaphroditlca L. (C. anceps AA. ; C. autuinnalis L.) 

(EtoilT dTeauT^ir'lfir^aves similar and narrowly linear, 

mostly around 1 cm long. Friiit 1.2-1.5-(2.0) mm wide, as wide 
as or slightly wider than long, deeply svilcate nearly to the 
central axis, being divided into 4 flat lobes. Svimmer. Slow 
moving water. — (G), Mack-(Y)-Aka, (L-NF), NB-BC, US, Eur. 

Order 4?. SAXIFRAGALES 
Resembling the Resales , with free petals and fused sepals, 
but the carpels more or less united and the flower typically 
perigynous . 

a. Carpels (4)-5; mostly fleshy plants 86. Crassulaceae 

aa. Carpels 2 87. Saalfragaceae 

86. CRASSUUCEAE (ORPINE FAMILY) 

Differs from the SaxLfragaceae b y its more n\imerous car- 
pels that are only slightly united at base. 

a. Flowers showy 1« Sedum 

aa. Flowers greenish, without petals 2. Penthorum 

1, SEDUM L. STC»JE-CROP 

Fleshy herbs of dry and rocky habitats with showy flowers 
like those of SaxLfraga , but the carpels more n\imerous, 

a. Leaves mostly opposite or verticillate 5« S. Rosea 

aa. Leaves alternate. 

b. Leaves very thick and less than 3 "nn wide. 

c. Stem leaves less than 5 mm long 1. S. acre 

cc. Longer, mostly around 1 cm long. 

d. Leaves narrowed at base 6. ^. lanceolatum 

dd. Conspicuously larger at base .. 

7. S. stenopetaltun 

bb. Leaves flat and at least 5 "nn wide. 

e. Flowers reddish 4. S. Telephium 

ee. Yellow. 

f. Leaves spatulate, dentate above 

the middle only 2. S. hybridvun 

ff . Lanceolate, serrate their whole 

length 3*S. Aiaoon 

1, S. ACRE L. — Mountain-Moss, Love-Entangle (Gazon d'or. 
Petite joubarbe) — The whole plant yellowish-green and forming 
a carpet less than 1 dm high. Leaves small and short, closely 
imbricated, not falling off in drying. Flowers yellow, few. 
Early summer. Cultivated and rarely escaped in dry or rocky 
places: Pointe-du-Bois , Ft. Qu'Appelle, Ma-Me-0. — (G, NF-SPM), 
NS-BC, US, Eur. 

2. S, HYERIDUM L. — Leaves 5-12 mm iri.de, short-spatulate, 
dentate only in the upper half. About 2 dm high. Yellow flo- 
wers in a terminal cyme. Early summer. Cultivated and rarely 

147 SEDUM 



88 PHYTOLOGIA Vol. 17, no. 2 

escaping to roadaldes and rocky places: Pointe-du-boia , Fort 
Saskatchewan. — Q, sMan, cAlta, Bur. 

3. S, AIZOON L. Leaves 3-10 cm long, lanceolate, ser- 
rate their whole length. Plant 2-6 dm high, lowers yellow in 
a cyme. Early summer. Cultivated and rarely escaped to road- 
sides: Ma-Me-0. — cAltn, Bur. 

It was also reported for Saskatoon by Russell 19^^, and 
Breitiing 1957, but the Justifying collection is likely to be 
only a CTiltivated specimen as it is labelled R.C. Russell , Sas- 
katoon, "U", garden, June 29, 1932 (SASK; DAO, photo). Further, 
It was later revised to S. Telephium . 

4, S. TELEPHIUM L. — Live-Forever, Orpine (Grassette, 
Chou au liJivre) — Flowers reddish in a dense terminal corymb. 
Stem k^7 dm high. Leaves t elliptic, rather iTge and very 
fleshy, coarsely dentate, often densely punctate in piorple. Kid 
summer. Cultivated and rarely escaped to roadsides; reported 
from The Pas, — (NF), NS_0-(Man), BC, US, Eur. 

5* A* ^2SS3 ^^'^ Scop, var. i^tggrifolium (Raf.) Eerger 
(S. Rosevun sphalm,) — Aaron's Rod, Mids\immer-Ken (Killegraine, 
Racine de Rose) — Leaves partly alternate, partly opposite or 
verticillate, entire, ovate to lanceolate. 1-3 dm high. In- 
florescence small, purplish-black. Early summer. Rocky alpine 
habitats. — Mack-Aka, Alta-BC, US, (Eur). 

In the more eastern var. Rosea the fruits are paler, pink 
to red, and the leaves are commonly dentate. 

6. S^, lwice5j^atus, Torrey (S, stenopetalum AA.) — Flower- 
ing stems arising from a dense carpet of sterile shoots. Leaves 
linear, those of the sterile shoots crowded and persisting in 
the herbariiun, the stem leaves not so crowded and falling off in 
drying. Flowers yellow in a terminal cyme. Early summer. Rol- 
ling montane prairies, ftom the Coteau Boise westward. — Y- 
(Aka), swS-BC, US. 

7, S^. s.tenop etalum Ptirsh (S, Douglasii Hooker) — Simi].ar 
but bulbiferous in the upper half of the stem. Leaves drying 
^itish and abundantly rusty-spotted. Bulblets axillary, folia- 
ceous . Early summer . Rocky places at mid altitudes : Water- 
ton, — swAlta-sBC, wUS. 

2. PENTHORUM L. DITCh-STONE-CROP 
Petals lacking and the plant not fleshy, 

1, P, sedoides L. — Perennial herb arising from a creep- 
ing base. Leaves lanceolate, serrate. Inflorescence glandular, 
terminal. Flowers in secund cymes. Filaments 10, persistent 
in fruit. Calyx lobes small and discrete. Mid summer. Shores 
and ditches, rare. ~ NB-seMan, US. 

87. SAXIFRAGACEAE (SAXIFRAGE FAMILY) 
Like the Crassulaceae , but the ovary typically reduced to 
2 carpels, 

SEDDM IkS 



1968 Boivin, Flora of Prairie Provinces 89 

a. Petals lacking 10. Chrysosplenium 

aa. Petals present, 

b. Stamens alternating with standnodia; 

carpels k; leaves entire 11. Pamassia 

bb. Staminodia lacking; carpels usually 2. 
c. Petals trifid to pectinate. 

d. Styles 3; leaves palmatipartite .. 
7. Lithophragma 

dd. Styles 2; leaves shall owly to 

deeply bilobed 8. Mitella 

cc. Petals entire. 

e. Inflorescence a simple raceme... 9. Conimitella 
ee. More branched and not a raceme. 

f. Stamens 5» 

g. Ovary bil ocular; inflores- 
cence cymose 2, Suksdorfia 

gg. Unilocular; inflorescence 
spicate to narrowly pani- 
culate 6. Heuchera 

ff. Stamens 10. 

h. Petals filiform, resembling 
the filaments of the sta- 
mens 5. Tiarella 

hh. Petals broader and more obvious, 
i. Carpels completely fused; 

styles partly fused.... 4. Teles onix 
ii. At least the styles free, 
j. Carpels mostly 

completely free..l. Leptarrhena 
jj. Carpels fused ven- 
trally for the lower 
half or so 3. Saxlfraga 

1. LEPTARRHENA Br. 
As Sajcifraga but the carpels nearly free to the base and 
the caljrx barely adnate to the base of the ovary. 

1. L. pyrpli folia (D. Don) Br. — Rather resembling Saxi - 
fraga rhomboid ea , etc . , but the stem typically bearing one lar- 
ge leaf which is t cordate at base. Basal leaves oblong, thick- 
ish, serrate, the nerves impressed above. Inflorescence densely 
glandular in red. Flowers marcescent. Petals white, narrow and 
inconspicuous, t linear. Early summer. Along creeks and sho- 
res. — Y-Aka, swAlta-BC, US. 

2. SUKSDCEFIA Gray 
Stem arising from a tuft of bulblets. Stsunens only 5 and 
the inflorescence cymose; otherwise as in Saxifraga . 

a. Flowers l-3-(7) 2. S. violacea 

aa. More numeroxis 1. S. ranvmculifolia 

149 LEPTARRHENA 



90 PHYTOLOGIA Vol. 17, no. 2 

1. j^* ran\inculifolia(Hooker) Englar ( Hemiev rinun cull fo - 
lia (Hooker)"Rflif7T'^^^^^''S^ein arising from a cluster of rusty-co- 
loured bulblets. 1-3 dm high and glandular-pubescent. Leaves 
palmatipartlte. Flowers white, usually with a daep red center. 
Late spring and early summer. Wet rocky places in the mountains; 
Waterton — (swAlta)-BC, US. 

2. ^. vioj3ceg. Gray — A delicate herb resembling many 
Saxlfraga , bu^Tthepetals pink to drying violet; they are white 
or yellow, sometimes red, in Saxlfraga, except S. oppositlfolia . 
Stem simple, 1-3 dm high, with few and inconspicuous basal 
biilblets . Herbage glandular-pubescent. Leaves mostly caulljie, 
alternate and palmatilobed to palmatlfid. Flowers few or sin- 
gle. Petals rather showy, oblauiceolate, sometimes nearly white. 
Late spring and early summer. Wet rocky banks and cliffs in 
the mountains; rare: Carbondale River, — swAlta-BC, US. 

3. SAXIFRAGA L. SAXIFRAGA 

The basic type of the family and -^ readily recognized ge- 
nus by its ovary obviously composed of two carpels that are fu- 
sed ventrally below the middle, but quite free in the upper 
half, the two styles conspicuously distinct. Stamens 10. 

a . Leaves opposite l6. S. oppositlfolia 

aa. Leaves alternate or all basal. 

b. Stem leafless below the inflorescence Group A 

bb. Stem leafy , Group B 

Group A 
Foliage mainly basal, the stem leafless, but the branches 
of the inflorescence often subtended by t reduced leaves, 

a. Leaves subcordate to deeply cordate at base. 

b. Many of the flowers replaced by clusters 

of bulblets 2. S. Mertensiana 

bb. Not bulbiferous 1. S. p\mctata 

aa. Leaves broadly to narrowly cuneate at base. 

c. Sepals sharply reflexed and pendent, 

d. Glabrous or slightly puberulent 

above 3. S. Lyallii 

dd. Abundantly glandular-pubescent 

throughout 6. S. ferruginea 

cc. Sepals ascending to more or less spreading. 

6, Petals 2-4 mm long 4, S, occidentalis 

ee. More elongate, 4.0-4,5 u™ long... 5. S. virginlensls 

Group B 
Stem with few to many leaves below the inflorescence. 

a. Leaves trifid to palmately lobed, 

b, Bulbiferous in the upper axils ., 10. S, cemua 

bb. Not bulbiferous. 

c . Leaf lobes ligulate 12. S. cespitosa 

SUKSDORFIA 150 



1968 Boivin, Flora of Prairie Provinces 91 

cc. Ovate to rounded 11. S. riwilaris 

aa. Leaves 3-toothed to entire. 

d. Flowers white; petals punctate or not. 

e. Leaves soft, with a rounded tip.... 9. S. ads pendens 
ee. Leaves stiff, prickly pointed. 

f. Leaves entire 13. S, bronchi alis 

ff. 3-toothed at apex 1^. S. tricuspidata 

dd. Yellow-flowered, the petals not punctate, 
g. Conspicuously long stoloniferous .. 

8. S. flagellaris 

gg. Not stoloniferous. 

h. Leaves all alike, all sessile... 15. S. aleoides 
hh. Basal leaves petiolate 7. S . Hirc\il\is 

1, ^ PSSSisSft ^* '^^'* Porsildiana (Calder & Savile) Boi- 
vin (S. aestivalis AA. ; S. argutTa lAJ)'^-!. Leaves deeply reni- 

f onn and flabellately lobed. Scapose, villous, stoloniferous. 
Flowers white with a red center. Filaments thin. Early summer. 
Wet cliffs near tiraberline: Rockies. ~ K-(Mack)-Y, swAlta-BC. 

Four other intergrading varieties occur in Canada, of which 
one may mention var, arguta (D. Don) Engl. & Irmsch, (including 
ssp, pacifica Htilt&i), with glabrous and larger leaves, the main 
ones 2,5-7.5 cm wide, occurring from southern Alaska to north- 
western B.C. This was also cited for Yukon as ssp. pacifica in 
Bot. Not. 109: 192, 1956,but the Justifying collection, N.J. 
Freeman , QxSu. Creek, 1953 (WIN; DAO, photo), has since been re- 
vised to var. Porsildiana . 

2. ^ Mertensiana Bong,~ Cocoa-Jiuts ~ Flowers partly re- 
placed by clusters of pinky bulblets, Scapose, reddish glandu- 
lar-pubescent. Leaves orbicular, deeply cordate, palmately lo- 
bed, the lobes 3-toothed, Inflorescence very open. Flowers 
white with conspicuously clavate filaments. Early summer. Drip- 
ping cliffs in the movintains : Waterton . — sAkn , swAlta-BC , wDS . 

3. S. lyallii Engler var. Lyallii — Leaves spatulate, 
coarsely "%o£lie3''tn' the upper haif'^ ^ Sc'apose and mostly around 
1 dm high. Inflorescence i racemose. Petals white to red tin- 
ged. Sepals deep red. Filaments clavate. Early summer. Al- 
pine brooks and late snow patches. Rockies, — swA.lta-sBC, (US) 
~ Var, HjOtwiii Calder & Savile — Taller plant, 1-3 dm high, 
with larger basal leaves, broadly obovate to flabelliform. In- 
florescence paniculate, — Y-Aka, swAlta-BC, US ~ Var. laxa 
Engler (S. Lyallii X S. odontoloma AA,) — Also taller, 2l^dm 
high and the basal leaves orbicular, broadly cuneate to sub- 
truncate at base. Inflorescence paniculate. Sometimes reputed 
a hybrid, but one parent is missing over much of the range, ~ 
(swAlta)-sBC, (US), 

4, S^, 5S£i^S2i^i2. Watson var, JJcc^jj^lwrt^a^As^ (S, nivalis 
AA.; S, rhomboidea AA,; S, r\ifidula (SmallJMacoiai; Micranthes 
rhomboidea AA. ) — Quite like the following, but the inflores- 
cence more congested and the flowers smaller. Herbage commonly 
reddish glandular-puberulent . Petals obovate to oblong. First 

151 sahfraga 



92 P H Y T L G I A Vol. 17, no. 2 

half of summer. Dry montane prairies on slopes: Cypress and 
Rockies. — (seAka), swS-swAlt«-sBC, wL'S. 

Fxirther south there occurs a number of rather weak varia- 
tions, of i^ich var. idahoensis (Piper) C.L. Hitchc. has stronj?- 
ly clavate filaments and var. latipetiolata C.L. Hitchc. h<is a 
short and broadly winged petiole. 

S. rhomboidea Greene is a Colorado and Wyoming species with 
a semi-inferior ovary, while in our S. occidentalis the ovary 
is almost completely superior. AH Canadian specimens met with 
under S. rhomboidea have been studied and revised to £. occi - 
den talis . 

5. _^. virginiensis Mx. — Everlasting, Sweet Wilson ( Pas - 
se -pi err e ) — Leaves typically rhomboid-ovate and serrate. Sea- 
pose, commonly 1-2 dm high, mostly glandular-villous . Petals 
oblanceolate. Mid spring. Open sandy or rocky places where it 
may be quite conspicuous at flowering time. — NB-seKan, Jo. 

6, S. fS^^iiginsS. Graham — Leaves rather l=rge, commonly 
3-10 cm long, cioneate-oblanceolate and remotely serrate above 
the middle only. Inflorescence diffuse. Flowers white. Petals 
unguiculate, lanceolate. First half of summer. Wet shaded 
rocks, at the middle altitudes: Waterton. — (n^ack), sAka, 
(swAlta)-BC, DS — F. yreelandii (Sm^ll) St. John & Thayer (var. 
Macounii Engler & Innscher) — Flowers p^^rtly replaced by green 
leafy bulblets, their leaves obovate to spatulate. — (sAka), 
swAlta-BC, US. 

7. S^. Hirculus L. — (Faux-ciste) — Flower yellow, usually 
solitary. Riifous -villous above. Stem leaves niiinerous, sessile, 
narrowly linear, the basal ones lanceolate, with a petiole about 
as long as the blade. Petals ± 1 cm long. Kid summer. Wet 
arctic tundra. — G-Aka, nQ-nMan, wDS, Eur. 

The many reports, new and old, from Saskatchewan, Alberta 
and B.C., are not substantiated by any specimen that w© could 
locate and were presumably based on old misidentifications or were 
speculative additions . 

8, 3. flagellaris W. var. flagellaris — Spider -Plant — 
Producing - 6 conspicuous superf iclar"stolons . Stem leafy, so- 
litary, with 1 to a few yellow flowers. Herbage ± glandular- 
pubescent. Stolons filiform, naked, about 1 dm long, rooting 
at tip. Mid summer. High alpine on polygons or solifluction 
soils: Rockies. — wMack-Aka, swAlta-nBC, US, Eur. 

The glandulosity is clear to light brown in ours but the 
glands aire purple black in the arctic var. platysepala Trautv, 

9. S^. a^cgjdeni, L. var. orggojignsis (Raf.) Breitung — 
Leaves mostly 3-toothed or 3-lobe37Dut soft -ynd not spinescent. 
Biennial, less than 1 dm high, glandxilar-puberulent throughout. 
Flowers \rtiite. Kid summer. Talus slopes and permafrost soils 
at high altitudes. — sY-seAka, swAlta-BC, wUS, 

The tjrpical eurasian phase is generally larger, with l^ir- 
ger flowers and larger stem leaves, 

10, S^, cernua L, — With clusters of fleshy, deep purple 
bulblets in the axils of the upper leaves . Glandular-villous , 
Leaves palmatilobed, the lower ones on very long petioles, 

SA.XIFRAGA 152 



1968 Boivin, Flora of Prairie Provinces 93 

Flower lAite, tsrpically single and terminal. Mid summer. Wet 
cliffs and mountain summits . — (G)-F-A.ka, L, Q, swAlta-BC, US, 
Eur, 

11. S. rivularis L, -- Similar to the proceeding, but not 
bulbiferouT ancTtKe^w flowers usually on very long pedicels, 
commonly longer than half the height of the plant. Leaves (3)- 
5-(7) lobed, not bulbiferous. Petals white. Early stimmer. 
Crevices of outcrops in arctic regions and in the mountains. — 
G-Aka, L-NF, Q, nKan, swAlta-BC, US, (Eur). 

12. S^ SSfEiSSS^ "^^ (^*^* groenl^J^dic^ (!-•) Pursh; var. 
minima Blai3c.T'"^^^-'T!^eaves digitately lobed, the lobes lig\ilate. 
Glandular -puberxil en t, forming dense cushions, the stems about 
1 dm high. Leaves cut into 3-(5) lobes. Flower white, often 
single. First half of svimmer. Alpine shale slopes and arctic 
gravels. — G-Aka, L-NF, Q, nMan, swAlta-seBC, US, Eur. 

13. S. bronchialis L. var. austromontana (Wiegand) G.N. 
Jones — Formiiig'^dwrse^ushions ofentireTstiff and spines cent 
leaves. Leaves marcescent, stiffly ciliate. Stem thin, glan- 
dular puberulent. Petals 5.0-6.5 mm long, not unguicvdate , 
^rtiite, with about 6 deep-red dots. Early to mid summer. Rocky 
alpine meadows. — sw/U.ta-BC, US, 

Replaced to the northwest by a var. pvirpureomaculata Hul- 
ten with unguiculate and someirtiat larger petals, typically 7-8 
mm long, 

14. ^. tri^usgidata Rottb. ( Leptasea tricuspidata (Rottb.) 
Haw.) ~ Leaves'^nesKyp^-toothed at apex, the teeth spiny. 
Carpet forming perennial, similar to the preceeding. Leaves li- 
gulate, stiffly ciliate. Flowers white, the petals with 10-15 
magenta dots . Early siimmer . Rocky outcrops in northern regions . 
— G-Aka, nL, nQ-BC. 

15. S. aizoides L. — Yellow- flowered carpet-making peren- 
nial. Stem densely puberulent, 1 dm high or less. Leaves all 
alike, sessile, narrowly lanceolate, marcescent, slightly fleshy. 
Mid summer. Alpine and arctic gravels and other loose soils. ~ 
G-Mack-(Y) L-NF, NS, Q-(nO)-nMan, swAlta-eBC, US, Eur. 

16. S^, oppositlfolia L. — Mayflower — Leaves opposite; 
flowers purpleV'^'Sensely^eafy carpet-making perennial. Leaves 
obovate, long ciliate, marcescent, turning blackish. Flowers 
solitary at the end of the branches. Early siimmer. Exposed 
rocky or gravelly places in arctic or alpine regions. — G-Aka, 
L-NF, Q, nKan, swAlta-BC, US, Eur. 

Reports of S. Aizoon Jacq. from Saskatchewan by many authors 
are probably based on the distribution given by Hooker 1832. The 
latter mention may have been based on collections from the Great 
Slave Lake or possibly the Great Bear Lake. 

4. TELESCaJIX Raf . 
Differs from Saxifra^a in the carpels being fused ventrally 
their i^ole length and the styles often partly fused. 

1. ^. "JsgesiA (Torrey) Raf. var. {jgjicheriforais (Rydb.) 
Bacigalupi ( Boykinia heucheriformis (Rydb7)RosrT^^^With the 

153 TELESONIX 



9li PHYTOLOGIA Vol. 17, no. 2 

general habit of a Heuchera , but the flowers reddish and the pu- 
bescence also often reddish, especially near the base of the flo- 
wer. Glandiilar-pubeecent throughout, 1-7 dm high. Leaves orbi- 
cvilar, lobed and dentate, cordate at base. Calyx ± reddish. 
Early sunmer. Rock crevices at the Hot Springs of Roche Klette. 
— 8wAltA-(BC), US. 

In ours the petals are obovate to spatulate and mostly 3 "■ 
long. The typical phnse, restricted to the Rockies of Colorado, 
Is somewhat larger flowered, the petals 3-5 nmi long and somewhat 
larger, broadly obovate to suborbicular . 

5. TIARELLA L. FALSE KITREWORT 
Flower slightly irregular. Upper calyx lobe somewhat 

longer than the others. Cairpels unequal in size the lower one 
often becoming as much as twice as long as the l-(2) upper ones 
in fruit. 

a. Leaves simple 1. T. unifoliata 

aa. Trifoliate 2. T. trifoliata 

1. J^ HCHSiJl^ Hooker (f . trisecta Lakala) — Petals 
insignif icantpaSoutaJs narrow as the filaments of the anthers . 
Glandular-pubertilent perennial, the leaves mostly basal, tri- 
lobed to tripartite, the lobes irregularly crenate-dentate . 
Flowers vrtiite in a narrow panicle. Early siumner. Kountain 
woods in the Rockies and Swan Hills. — Aka, Alta-BC, nwUS. 

More deeply lobed specimens have been called now a mere 
form, f. trisecta , now as an interspecific hybrid to T. trifo - 
JLlata , The last assumption seems rather improbable since the 
form was originally described from the albertan Rockies, an 
area ^ere one of the postulated parents is not known to occur. 

2, T. tj^g^jiata^L. — Similar, but the leaves trifoliate. 
Tending to* be taller and more abundantly flowered. Early sum- 
mer. Wetter coniferous forests, rare: Whltecoiart, — sAka, 
wcAlta-BC, nwUS. 

6. HEUCHHIA L. ALUM-ROOT 
Stamens only 5 *s in Suksdorfia, but the carpels fused in- 
to a unilocular ovai^r. Cthei-wise as in Saxifraga . Flower often 
somewhat asynetrical. 

a. Calyx 2-4 mm long. Including the semi-inferior ovary. 

b. Leaf -teeth acute 1 . H . glabra 

bb. Leaves broader, their teeth broadly 

roTonded 't. H. parvifolia 

aa. Flowers larger, the calyx 5-12 mm long. 

c. Stamens Included in the calyx ........ 2. H. cylindrica 

CO. Stamens exserted; leaves and flowers 

larger • 3. H. Richardsonii 

1. H. giants. ^* — Pedicels rec\irved, mostly longer than 
the flowers. Leaves sharply dentate, at least one well developed 
TIARELLA l5U 



1968 Boivin, Flora of Prairie Provinces 95 

leaf borne on the stem or subtending the lowest branch. Panicu- 
le open, sometimes sec\md. Mid summer. River cliffs, rare: 
Mt, Edith Cavell. — sAJca, svAlta-BC, US. 

2. H^. c^^Jidrica Douglas var. ^.i^bella (T. & G.) Wheelock 
(var. septentrionalis R., B. & L.) — Petals linear, included 
and inconspicuous, but the cal^x lobes yellowish, Scapose pe- 
rennial 3-6 dm high. Leaves broadly ovate, lobed, the lobes 
crenate. Inflorescence a narrow racemiform panicle. Late 
spring to mid summer. Open rocky slopes in the moxintains, — 
swAlta-s3C, wUS, 

Petioles glabrous or somewhat glandular-puberulent, never 
hirsute. The tjrpical phase occiirs west of \is and is readily 
recognized by the dense and mixed pubescence of the petioles, 
partly long hirsute, partly gl^ndular-pubervilent. 

3. H^. Richardsonii Br, var, Richardsovy. (var, hispidior 
R,, B, & L, ;''fT ^hisi^x[r 'AA, ) — Alum-Root — Much like the pro- 
ceeding, but the calyx strongly asymetrical and the stamens ex- 
serted. Calyx barely petaloid. Petals pink, spatulate, about 
as long as the calyx lobes. Early summer. Common on rolling 
prairie. — Mack, 0-sMan-neBC, US. 

In CMTS the capsvile is included, the stamens barely exsert- 
ed and the petals are merely papillose. We have submerged var. 
hispidior as being a mere sporadic extreme of pubescence. 
Further south one may find v^r, Grayana R., B. & L. (including 
var. af finis R., B. &L, , a smaller-flowered extreme) with a 
somewhat exserted capsule, more strongly exserted stamens and 
petals at once glandular and papillose. 

4. H^, par^iifolis Nutt. var. dissecta M.E. Jones (H. fla - 
bellifolia RydbTT'lI^^lowers small and the white petals exserted 
as in H. glabra , but the panicle narrow and racemiform. Gene- 
rally smaller, the leaves only 1-3 cm wide. Late spring to 
early summer. Foothill prairies, — (swS)-swAlt^'-(seBC), US, 

7, UTHOPHRAGMA Nutt, 
Petals conspicuoxosly and digitately lobed. The gender of 
this genus was discussed in Taxon jL2: 208, I963. 

a, Bulbiferous in the upper axils ..•..••• 1, L, gl»brum 

aa. Not bulbiferous •*••• • 2, L, pgrviflorum 

!• ii» £i2^;ii2 Nutt. ramulosum (Suksd.) Boivin (L. bulbi - 
ferum Rydb. ; L. tenellum A ATP^^^^'^wer flowers replaced by clust- 
ers of deep-pvirple fleshy bulblets. Otherwise, quite like the 
following. Calyx campanulate, elongating up to 5 nnn in fruit. 
Petals somewhat smeller, mostly trifid. Late spring. Prairies 
near springs: Cypress Hills and Rockies. ~ swS -swA.lt a-BC, US. 

The more restricted var. glabrum from the western United 
States lacks any bulblets. 

2. L. ^^CGfiSSffl (Hooker) Nutt. — Leaves palmatipartite. 
Flowers few, in~T'^!ermiiipl raceme. Calyx more elongate, cuneate 
at base, elongating to 6-10 mm in fruit and becoming sc»ne^at 
tubular. Petals white, mostly 5-lobed. Early svimmer. Moist 

montane prairies. — swAlta-BC, US, 

155 LITHOPHRAGMA 



96 PHYTOLOQIA Vol. 17, no. 2 

8. MTTELU L. MrTRHTz/ORT, BISHOP'S CAP 
Petals trifid to pectinateLy divided into filiform seg- 
ments . Styles 2 . 

a. Petals dipitately trifid, wnite U. M. trifida 

aa. Petals pectinate. 

b. Pedicels 1-2 mm long; petioles villous 

witn lone rufous hairs 3- M. 3reweri 

bb. Longer; pubesceiice wfiite. 

c. Stamens 10; leaves broadly rounded 

at tip I. M. nuda 

cc. Stamens 5; leaves obtuse at tip; 

larger plant 2. M. pentandra 

1. M. nuda L. — Small delicate forest herb with yellow- 
ish-petals pectlnately divided. Smaller, l-(2) dm nigh. Lea- 
ves smaller, l-3-(!)) cm wide, suborbicular, deeply cordate, 

± crenate. Stamens 10. Seeds black, small,but conspicuous on 
the cup-like fruit wall. Early summer. Common forest species. 
~ (K)-Mack-Y-(Aka), L-SFM, NS-BC, US, (Eur). 

2. M. P£2J^a£idr£ Hooker — Stamens only ? and opposite tne 
greenish petaYs/Leaves broadly cordate, shallowly lobed, tne 
lobes crenate. Summer. Wetter spots in montane and subalpine 
forests and meadows. — Y-Aka, wAlta-BC, US. 

3. M. 3reweri Gray — Much as in the preceeding, but tne 
leaves broader and reniform. and the stamens opposite the calyx 
lobes . Leaves merely crenate or sometimes weakly lobed . Mid 
summer. Wetter areas in the upper montane zone in Waterton. 
— swAlta-3C, US. 

Ii. M. trifida Graham (M. violacea Rydb.) — Calyx lobes 
whitish and the trifid petals white . Stamens 5j opposite tne 
calyx lobes. Leaves more like tnose of M. pentandra. First 
half of summer. Mountain springs and we^ cliffs . — (swAlta)- 
BC, US. 

9. CONIMITELLA Rydb. 

Differs from Mitella by its entire petals and almost com- 
pletely inferior ovary. 

1. C. Williamsii (D.C. Eaton) Rydb. — Bracts petaloid, 
white and pink, 1-2 mm long and fimbriate. Herbage densely 
glandular-puberulent . Leaves reniform, all basal. Scape ra- 
ther long, bearing only 5-10 subsessile flowers. Petals white, 
narrowly oblanceolate, U-5 inm long including a claw nearly as 
long as the blade. Calyx lobes i 1 mm long, petaloid, white 
and pink. Early summer. Rich montane forests: Crownest Fo- 
rest. — swAlta , wUS. 

10. GHRYSOSPLEMIUM L. GOLDEN SAXIFRAGE 
Petals lacking. Carpels 2, united into a unilocular ovary, 

the two styles far removed to opposite sides of the ovary. Sta- 
mens marcescent and present even in fruit. 
MITELLA 156 



1968 Boivin, Flora of Prairie Provinces 97 

1. C. aliernifoliAom L. var. tetrandruin (Th. Fries) Lund 
(C. americanum AA.; C. tetrandrum Th. Fries) — (Cresson dore, 
Cress on de roche) — Small erect herb, usually less than 1 dm 
high, with reniform and crenate leaves. Most leaves and flo- 
wers clustered near the top of the plant. Sepals all alike, 
green, erect. Stamens U, opposite the sepals. Early summer. 
Wet shaded places. — (G)-F-K-(Mack-Y)-Aka, (L), Q-(0)-Man-BC, 
wUS, (Eur) — Var. iowense (Rydb.) Boivin (C. iowense Rydb.) — 
Sepals of two sizes"ptneouter ones somewhat wider. Sepals yel- 
lowish-green, recurved at tip. Stamens 5 to 8, the additional 
ones alternating with the sepals. — svMack, sMan-sAlta, (ncUS, 
Eur). 

Var. iowense is very close to var. sibiricum Ser., the main 
distinction of the latter being that the stamens are always in 
8's. 

11. PARNASSIA L. GRASS OF PARNASSUS 
With $ clusters of staminodia, each cluster oorne on a fla- 
bellate base. Carpels h- Herbs with entire leaves and a sin- 
gle terminal flower. Stem scapose or unifoliate. 

a. Leaves reniform U» P« fimbriata 

aa . Leaves ovate, longer than broad. 

b. Petals small, about as large and as long 

as the sepals 1. P. Kotzebuei 

bb. Much larger, at least twice broader than 
the sepals. 

c. Stem leafless 3* P» glauca 

cc. Stem unifoliate 2. P. palustria 

1. ^. j^tzebuei Cham. var. IJotzebuei^ — Smaller, usually 
around 1 dm hi^. Stem leafless. Flower small, the petals el- 
liptic-lanceolate and about as long as the calyx lobes. Before 
mid summer. Wetter alpine and arctic meadows. — (G-F)-K-Aka, 
L-NF, Q-(nO)-nMan-(nS)-Alta-BC, US, (Eur). 

A dwarf var. pumila Hitchc. & Ownbey with much reduced 
staminodia has been described from a limited area in the Okana- 
gan Valley. 

2. P. palijstris L. var. tenui^ Wahl . (var. neogaea Fern.; 
P. multiseta^LedTT^ern.) — Grass^f Parnassus, White Butter- 
cups (Fleur du Pamasse) — Tufted herb, each stem bearing a 
single smaller, cordate and sessile leaf towards the lower 
third. Stem usually 2-h dm high. Leaves ovate, broadly rounded 
to cordate at base. Petals about 1^ times as long as the se- 
pals. Staminodia cluster typically with more than 10 segments. 
Mid to late summer. Wet meadows and marshy places. — K-(Mack)- 
Y-Aka, (L-NF), Q-BC, US, (Eur) — Var. mOTtanensis (Fern. & Rydb.) 
C.L. Hitchc. (P. montanensis Rydb. & Fenii )' — "Sonevihat smal- 
ler. Petals only slightly longer than the calyx lobes. Stami- 
nodia with less than 10 segments. Not always clearly distinct. 

— (Y), Alta-(seBC, US) — Var . jarvifloj^ (DC . ) Boivin (P. 
parviflora DC.) — Still smaller. Typically 1-2 dm high. Stem 

157 PARNASSIA 



98 PHYTOLOGIA Vol. 17, no. 2 

and basal leaves usually cuneate or rounded at base. Petals 
less than 1 cm long. Staminodia witn lesc than 10 segments. — 
(Mack-Aka, L)-^iy, N3-PEI, Q-nllan-seBC, US. 

The inclusion of P. montanensis in Saskatcnewan lists oy 
Russell 195U and Breitunf 1957 is credited to Raup 1936. How- 
ever the latter eives only tnree localities, two of them, Calu- 
met and Shelter Point, deinp in Alberta while Great Slave LaKe 
is in Mackenzie District. There was no Saskatcnewan sheet at 
GH in 196$. 

3* X.' Z^33St Raf. (P. america na Muni.; P. carollniana AA.) 
— Flowering Plantain — Leaves all oasal, broadly ovate to 
elliptic, rounded at base. Calyx lobes snort, only 3-5 ™n long. 
Petals 10-18 mm long, more than twice as long as the calyx lo- 
bes. Staminodia mostly with 3 coarse and reddish segments. Late 
summer. Wetter prairies. — NF, NB-cS, US. 

Canadian reports of the soutnem P. carollniana Mx. are ge- 
nerally based on specimens of P. glauca, but Gardner's 19U6 re- 
ports for Cnurcnill and Labrador are undoubtedly based on some- 
tning else still. The corresponding specimens could not be 
found at DAG or QFA in 1965 and 1966. 

Ii. ^. fi jn br ia ta Konig var. fjjTbria^g — Petals coarsely 
fimbriate on each side in the lower nail . Leaves broader than 
long, reniform and deeply cordate. Stem leaf small, borne to- 
wards the middle. Mid summer. Brooksides and springs near 
timberline. — (ffv*Iack)-Y-Aka, swAlta-BC, US. 

The staminodia are snort, stubby and not capitate in our 
variety, but they are longer, thinner and capitate in two other 
varieties from the western U.S. 

Order U8. SAPPACSNIALES 
Carnivorous and capturing insects in a variety of ways. A 
primitive type of flower witn the parts mostly in 5's and free, 
except for the fused carpels. 

a. Inflorescence a raceme 88. Droseraceae 

aa . Flower solitary 8?. Sarra^'eniaceae 

88. DROSEPACSAE (S^JNOF// FAMILY) 

Single genus with us. Styles 3-5* Insects trapped by 
hair-like processes. 

1. DROSERA L. SUNDE".-/ 

Leaves covered with coarse hair -like processes, capitate, 
glutinous and in which the insects become trapped to be eventual- 
ly digested. Heros with the leaves all basal and flowers in a 
raceme borne on a scape. 

a. Leaves linear, the limb •*• 2 mm wide 2. D. linearis 

aa. Broader. 

b. Leaves ± obdeltoid, sligntly broader 

than long 3« D. rotund if olia 

bb. Leaves obovate to broadly oblanceolate. . . 1. D. anglica 
PARNASSIA 158 ~ 



1968 Boivin, Flora of Prairie Provinces 99 

^» J^ SfiiiiSS Hudson (D, Intermedia AA.) — Leaves 1-3 cm 
long, 2.5-^.0 mm wide, narrowly obovate to narrowly oblamceola- 
te, elongating in age. Mid summer. Northern bogs, usually in 
wetter and pioneer habitats. ~ Mack-Aka, L-NF, Q-BC, US, Eur, 
(Oc). 

Sometimes treated as the hybrid of D. linearis X rotundi - 
folia but the Canadian distribution of D. anglica extends much 
further north than that of D. linearis and the solution of hy- 
bridity does not seem very plausible, 

^' ^* ii£2S£i5 Goldie — Leaves 2-4- (6) cm long, 2- (3) mm 
wide, long linear, erect. Mid svunmer. Bogs, rare. — NF, Q-S- 
(Alta)-BC, US. 

3, JD, rotmj^ifolia L, var, £^t}Jgi<iifolia — Dewgrass, Eye- 
bright (HTrbeTlagoutte, Petit Saint-Sacrement) — Leaves 
wider and more spreading, more or less obdeltoid to suborbicu- 
lar, (5)-8-10-(12) mm wide and usually slightly wider than long. 
Early to mid summer. Sphagnvim hummocks in bogs. ~ G, seK-Aka, 
L-SPM, NS-BC, US, Eur. 

89. SAItRACENIACEAE PITCHER-PLANT FAMILY 
Insects trapped in hollowed out petioles half- filled with 
digestive liqiiids. Stamens n-otierous. Style 1. 

1. 3ARRACEKIA L. SIDE-SAD^^LE FLOWffi 
Style tinusually large, shaped like an umbrella, and wider 
than the ovary or fniit, which it covers. 

1. ^. ^iJJ^urea L. var. mrgurea — Indian Pipe , Frog's 
Trous ers ( Sabot , Cochon de pele ) — A single, large, drooping, 
deep red flower on a long scape, arising from a rosette of lea- 
ves half- buried in Sphagnum . These shaped like "horns of plen- 
ty", and half full of water. Sepals 2.5-4,0 cm long. First 
half of sxunmer. Sphagnum bogs. — L-SPK, NS-neAlta,US — Var. 
rira^cola Boivin — More superficial, the rhizome very short or 
njnoistinct , the whole plant not buried in moss. Sepals 
shorter, 1.5-2.2 cm long. Wet terraces and shores, rare: Ni- 
pawin and Prince Albert. — cC, cC. 

The only Alberta collection seen was from Anzac (ALTA; 
DAO, photo). It is made up of 3 separate leaves only and its 
varietal determination remains tentative. 

Order 49. UMBELLALES 
Related to the Araliales . Carpels 2, maturing into a dry 
fruit which splits into a pair of achene-like fruits. Achenes 
borne on a central structxire termed carpophore. Single family. 

90. L^^PEI.IIFERAE (PARSLEY FAMILY) 
Flowers in ijmbels and the ovary inferior. Flw-rers 5-raerous, 
the perianth parts free, but the sepals much reduced. Flowers 
typically unisexual. Genpric characters in this family are 
often rather obscurely technical. 

159 SARRACENIA 



100 PHYTOLOGIl Vol. 17, no. 2 

a. Flowors in bluelah heads; foliage splnescent... 2. c.ryn/.ium 
aa. Flowers in umbels. 

b. Leaves digitately compound 1. ^anic\ila 

bb. Not digitate, although sometimes trifoliate, 
c. Leaves divided progressively into 
numeroiis small and rather narrow 

ultimate segments Group 1 

cc. Leaves simple or divided into fairly 
well defined leaflets. 

d. Stem leaves simple to trifoliate Group B 

dd. Leaflets more mimerous Group C 

Group A 
Leaves deeply and progressively divided into many and ra- 
ther narrow segments; leaflets not obvious or poorly defined. 

a. Flowers mostly replaced by bulblets 9. Cicuta 

aa* Not bulbiferous, 

b. Involucre of large and pectinately 

dissected bracts 22, Daucus 

bb. Bracts much smaller and little if at 
all dissected, or even lacking. 

c. Umbell simple and few flowered 3. Scandlx 

cc. Compound and the flowers very nimerous, 
d. Leaves all basal, or at least the 
lower pair opposite, 
e. Fruit not winged, but finely 

tuberculate 6, Muslneon 

ee. Fruit winged, not tuberculate. 
f , Fruit winged along the 

marginal nerves only ..... 19 • Lomatium 
ff. Conspicuously winged along 
both the marginal and 

dorsal nerves 18. Cymopterus 

dd. Stem leaves all alternate, sometimes 
opposite in the Inflorescence, 
g. Segments very few (mostly 5) » 

very narrow and very long.. 13. Perlderidla 
gg. Segments much more nvmerous 
and shorter, 
h. Stem with irregularly 

scattered purple blotches.,., 5» Cani\m 
hh. Stem not maculate. 

1, Native perennial; fruit 

very flat 19. Lomatium 

11, Annual or biennial weeds; 
fiTilt slightly compressed, 
j. Flowers white; the 
shorter pedicels 
shorter than the 

fruit ,., ,.... 11. Carum 

UMBELLIFERAE 160 



1968 Boivin, Flora of Prairie Provinces 101 

jj. Yellowish-green; all 
pedicels many times 
as long as the fruit .. 
15* Anethvun 

Group B 
Leaves simple, entire or merely dentate to lobed or trifo- 
liate, the leaflets rather broad, 

a. Leaves entire • ?• Buplevtrum 

aa. Serrate to trifoliate. 

b. Leaflets huge, at least 1 dm wide 21, Heraclemn 

bb. Much smaller or the leaf simple, 

c. Flowers yellow; primary raj^s of the 

umbel nearly vmiform in length ••,, 8, Zizia 

cc. Flowers white; \unbel rays very 

uneven •.••,. 10, Cryptotaenia 

Group C 
Leaves coiqjound, the leaflets more than 3 and all or most 
of them discrete and well defined, 

a. Leaves pinnate, 

b. Leaflets -i: linear 14. Sium 

bb. Leaflets ± oblong , 20, Pastinaca 

aa. Leaves temately divided. 

c. Leaflets not serrate, but entire or with 
a few lobes, 

d. Stem tall and leafy 17. Levis tjcum 

dd. Stem short, the leaves all basal or 

near basal , ,,,, 19, Lomatium 

cc. Finely to deeply serrate, 

e. Firuit strongly flattened dorsally 

and xdjiged , l6, Angelica 

ee. Fruit slightly flattened laterally, 
wingless, 
f , Leaves symetrically divided into 

(3) or 9 leaflets «.,,,,,,, 12, Aegopodium 

ff. Central segment more divided than 
the lateral ones , the leaflets com- 
monly 5 or 15 or 21, etc, 
g. Fruit over 1 cm long, us\ially 

setose-strigose ,, 4, Osmorhiza 

gg. Fruit glabrous, much shorter, 
h. Flowers yellow, the central 

pistillate one subsessile ,,,, 8, Zizia 
hh. White and all pedicelled ,,,, 9. Cicuta 

1, SANICULA L. SANICLE 

Fruit catchy, being covered with numerous hooked prickles. 
Calyx nearly as large as the corolla, 

161 SANIGUU 



102 PHYTOLOGIA Vol. 17, no. 2 

1, S^, mari^andica L, (S, marylandlca ephalm.) — 3nako- 
Root, BlAcic Snake^oot"^ Coramon deciduous foreat species with 
digitate leaves. Leaflets 5, obovate to oblanceolate, seasile, 
serrate, the larger 2 often bifid to bipartite. Stem simple, 
the branching of the inflorescence tending to be opposite. Early 
summer. Nearly ublqvdtoTos in deciduous woods. — KF-SPK, K5-BC, 
US, (SA). 

2. ERYNGIUM L. 
Flowers in dense heads, much simulating a Composite. Fruit 
densely covered with membranous scales. 

1. E. PLANUM L. — (Herbe aux serpents) — Stiff herb, 
bluish above. Foliage spiny-toothed. Leaves alternate, but 
the main branches of the inflorescence verticlllate. Heads with 
a spinescent involucre. Flowers bl\iish. Mid summer. Casual 
escape from cultivation, ~ Q-0, S-BC, (US, Eur). 

3. SCANDIX L. 
Body of the fruit prolonged into a much longer cylindrical 
beak. 

1, S, PECTEU -VENERIS L. — Venus* Comb, Lady's Comb (Pei- 
gne de Ventis, Aiguille de berger) — Fruit longest, k-7 cm long. 
Annual with the leaves finely dissected into very numerous and 
narrow segments. Umbels simple, of less than 10 flowers and 
subtended by an involucre of ± connate bracts. Flowers white, 
Frtiit scabrous. Carpophore needle-like. Late spring to mid 
summer. Rare weed: Golbum, — 0, S, EC, US, (SA) , Eur, (Afr, 
Oc). 

k, OSMORHIZA Raf , SWEET CICELY 

Except for one atypical species, fruit catchy by appressed 
and acicular hairs, especially numerous towards the base, the 
latter prolonged into a sharp and fairly long point, 

a. Flcfwers yellowish or greenish; fruit 

glabrous 1. 0. oceiden talis 

aa. Flowers white or pink; fruit coarsely 
strigose, 

b. Involucre and involucels lacking ,,.,,, 2. 0, chilensis 
bb. Involucre and involucels present 3« 0, aristata 

1, ^, occidfflvtalis (Nutt,) Torrey — Atypical, the black- 
ish acheries UnearT'glabrotis , and devoid of a sharp basal beak. 
Main leaves typically iri.th 15 or 21 leaflets, the latter lanceo- 
late to elliptic-lanceolate, puberulent. Involucre and involu- 
cels lacking. Fruit 12-18 mm long, longer than its pedicel. 
Late spring. Open woods and rocky slopes at lower altitudes, 
— swAlta-seBC, wDS, 

^« 2f S^^^tSE^iB. ^* * ^» '^^^* chilensis (0. bravipes (C, & 
R.) Suksd,; 0, divaricata (Britton)"Sulcsdn^~ Usually with one 

ERYNGIUM 162 



1968 Boivin, Flora of Prairie Provinces 103 

stem leaf below the inflorescence, of 9 leaflets, the latter 
triangular -lanceolate, serrate above, gradually more deeply cut 
below. Flowers white. Fruits (1.5)-2.0-(2.5) cm long, all or 
mostly longer than their pedicel, the latter 0.5-2.0 cm long 
and widely divergent. Early siumner. Woods, ~ sAka, NF, NS, 
NB-0, swS-BC, US, (SA) — Var. purpurea (C. & R.) Boivin (O. 
purpurea (C, & R.) Suksd.) — Flowers^ink or at least with a 
pink center, rarely white. Fruit shorter, (0.8)-1.0-(1.5) cm, 
stubbier at tip, shorter than its pedicel, ~ sAka, swAlta-BC, 
nwUS — Var. cuprgssi^MvJgna Boivin (0. depauperate Phil,; 0, 
obtusa (C, & R/T F^rnr)--Fiowers white. Fruits not so short, 
± 1.5 cm long, yet all or most of them shorter than their pedi- 
cel, the latter (l)-2-3 cm long. Stem Tisually leafless below 
the inflorescence, the lower leaf of the latter usually with 
9 leaflets. — seK, sAka, (sL-NF) , NS, (NB)-Q-BC, US, (SA). 

3» 2f Sd^t^ (ThTinb.) Mak, & Yabe var. brejistylis (DC.) 
Boivin (0. Claytonii (Mx.) C.B. Clarke) — SweelJarvxi*^ Com- 
monly with one stem leaf of i 2? leaflets , the latter as in 0. 
chilensis . Herbage villous. Flowers white. Pedicels mostly 
0,5-1.0 cm in fruit. Friiit ± 1,5 cm long. Styles 0.5-2,0 mm 
long. Late spring. Poplar woods at Moon Lake in Riding Kotin- 
tain ~ NF, NS-sKan, US — Var, J^ngigtylis (Torrey) Boivin (O. 
longistylis (Torrey) DC.). ~ )^iiseiRoo€^ Paregoric-Root — 
Stem glabrous, the foliage glabrous to villous. Styles longer, 
2.0-3.5 nim long. Oak bluffs and galerie-forests. — NS, NB-Alta, 
US. 

Reports of var. brevistylis (= 0. Claytonii ) from western 
Canada appear to be all based on specimens with the longer sty- 
les and lesser pubesceaice typical of var, longistylis . Except 
for the Riding I-'ountain and perhaps also for the Cypress Hill 
reports. The Ilacoun collection (QK; DAO, photo) from the Cypress 
Hills was typical indeed of var, brevistylis , but in the absence 
of later confirmation, we are inclined to suspect the possibili- 
ty of mixed labels. 

Our two varieties are not shai^sly disjunct morphologically 
and consequently a number of intermediate types based on vinusual 
associations of diagnostic characters have been described and 
named. Specimens vrith styles of intermediate size are not un- 
common and one is then left with pubescence as the only usable 
distinction. Further the asiatic 0, aristata is more or less 
intermediate between our two types , the herbage being villous 
(as var, brevistylis ) but the beak rather longish (like var. lon - 
gistylis ) or not infrequently intermediate in size. However, 
var. aristata is best distinguished by its commonly longer pedi- 
cels, these being l-2-(3) cm long in fruit while they are usual- 
ly about 0.5 cm long in our two american varieties, sometimes 
longer, but never averaging more than 1 cm on any plant. 

The rank of variety seems most appropriate for these inter- 
grading and morphologically overlapping taxa. The varietal rank 
also reflects most obviously their londeniable and very close 
affinity. ^^^ OSMCRHIZA 



lOU PHYTOLOGIA Vol. 17, no. 2 

Var. brevistylis (DC.) stat. n., 0. breviLtylii; DC., Prod. 
U: 232. 1(530; Urospermum aristat'jw (Tniinb.) Ktze. var. bre vie - 
" Tyle (DC) Ktze., Rev. Gen. 1: 270. 1991; Osmorniza ClaytonlT " 
TMx!) C.3. Clarke. ^ 

Var. l onpistylis (Torrey) atot. n., Myrmis lonr^iatylis 
Torrey, Fl. U .S. 310. ld2U; Uroapennm n ar iatatum (Tnunb.) Ktze. 
var. lorif^iatyle (Torrey) Ktze., Rev. Tien. 1: 270. I89I. 

^. CONTUM L. POISON' HEJ1L0CK 

Ribs of the fruit preeminent and strongly sinuous . Carpo- 
phore not becominf^ bifid. Stylopodium very broad. Otnerwise 
the fruit resembles Cicuta . 

1. C. MACULATUM L. — Poison Hemlock (Cipue d 'Europe) — 
Stei.i sparsely to densely and irregularly purpj.e-blotcned. Lea- 
ves divided into very mamerous small segments, the mairi ones 
alternate, becoming opposite in the inflorescence. Bracts of 
the involucre (and involucels) broadly margined, tending to be 
fused and usually ref lexed . Early to mid suirjner. Estaolished 
along roadsides at Maclean. — NS, Q-0, S, swBC, US, Eur. 

6. MUSINEGN Raf . 

Rather resembling Lomatium , but the fruits wingless and 
only sligntly ccmprei-sed laterally. 

1. M. divaricatum (Pursh) Nutt. (var. Hookeri T. & G.; 
M. trachyspermum Nutt.) — Conspicuous in early spring on dry 
Hillsides, a low nerb with an umbel of yellow flowers and at 
least one pair of opposite leaves. With a deeply buried tap- 
root and much dissected leaves. Puberulent to scabrous, espe- 
cially the stem and inflorescence. Up to 2 dm hi^. First half 
of spring. Hillsides. — sv^an-sAlta, US. 

7. BUPLEURUM L. THOROUGH -WAX 
Fruit resembling the preceeding but smooth and the stylo- 
podium especially broad. 

1. 3. americanum CAR. — Leaves simple, entire, linear- 
lanceolate. Involucre and involucels rather large and conspi- 
cuous. Flowers pale yellow with the stylopodia foming a cons- 
picuous brown center. Fruits (and ovary) strongly glaucous, 
rather bluish. Mid sumirier. Gravelly and rocky prairies: Water- 
ton. — nwMack-Aka, swAlta-seBC, nwUS. 

The inclusion of B.C. in the distribution is based solely 
on a collection by Dawson at the head of tne Kootenay River in 
1871 (CAN). This has never been confirmed and we have also 
come to appreciate that the geographical data on Dawson's la- 
bels are accurate only within a rather broad margin of approxi- 
mation. It could be that Dawson's collection came from the 
Alberta side. 

CONIUIl 16U 



1968 Boivin, Flora of Prairie Provinces 10$ 

8. ZEIA W. D. J. Koch ALEXANDERS 

Fruit slightly ccsnpressed laterally as in the last few ge- 
nera, but the stylopodium wanting. Each umbellule of pistil- 
late flowers shows a central flower sessile or nearly so. 

a. Basal and lower leaves simple, the middle and 

upper trifoliate 1. Z. aptera 

aa. Basal and stem leaves biternate, with 9-11 ~ 

leaflets 2. Z. aurea 

^* ^* ££i5££ (Gray) Fern. (Z. cordata AA.) — Alexanders 
— A common yellow -flowered herb conspicuous in early summer in 
ditches and other wettish places. Basal and lower leaves cor- 
date, crenately serrate. Leaflets ovate to lanceolate, serrate. 
Leaves thickish. Early summer. Chernozem prairies and wetter 
places. ~ swY, swQ-BC, US. 

The recent extension of range to Yukon by Boivin 1966 was 
based on j^. FouiTiier . Haines Junction, 25 juillet 19$8 (QFAj 
DAO, photo). 

2. Z^. aurea (L.) W.D.J. Koch (Thaspium barbinode A A.) — 
Golden Alexanders, Meadow-Parsnip — Similar, the leaves thinner 
and more divided, mostly with 9 or 11 leaflets. Often taller, 
5-10 dm high. Leaflets rhomboid to lanceolate, serrate. Early 
summer. Galerie -forests. Oak islands and low chernozems. — NS, 
NB-sMan, US. 

Despite numerous Saskatchewan reports of Z. aurea, all of 
the U or 5 collections found under that name in various herbaria 
turned out to belong to Z . aptera . All Manitoba specimens under 
Thaspium barbinode (Mx,)~Nutt. at CAN and DAO also proved to be 
i . aurea . 

9. CICUTA L. WATER -HEMLOCK 

A middling type \-rith small, slightly compressed and vring- 
less fruit. Flowers white. Involucre much reduced or absent. 
Base of stem slightly bulbous and fistulous with numerous cross- 
plates. Very poisonous plants. 

a . Flowers mostly replaced by clusters of 

bulblets 1 . C . bulbifera 

aa. Not bulbiferouE. 

b. Fruit depressed globose 2. C. mackenzjeana 

bb. Ovoid J leaflets broader ~3. C. maculata 

1. ^. buljjjfei^ L. — A rather sparse herb with at least 
one terminal white umbel and numerous bulblets scattered along 
the branches. Annual or perennial, 5-12 dm high. Foliage dis- 
sected to filiform segments, about 1 mm wide and entire or so- 
metimes very remotely serrate. Fruit infrequent, suborbicular, 
about 1,5 mm long and about as wide. Second half of summer. 
Swampy ground or snores. — sMack, L-tiF , NS-BC, US. 

2. j^ mackenzjeana Raup — Like a narrow-leaved form of 
the following. Tuberous roots poorly developed or lacking. 

165 CICUTA 



106 PHYTOLOQIA Vol. 17, no. 2 

Rather thick -steatuned for its sparae foliage and tending to be 
faatigiate in habit. Leaflets linear-elongate, about 10-1$ ti- 
mes as long as broad, usually less than $ irjn wide. Fruit oroad- 
ly orbicular, 2.0-2.^ mm long, as vd.de or wider than long. Mid 
summer. Marshes and bogs northward; mainly subarctic iri dis- 
tribution. — Mack— Aka, wcQ-neBC. 

3. C. maculata L. var. anfUitifolia Hooker (C. Douglasii 
AA.; C. occidervtaTiE Greene) -"^CowbaneToeaver-Poison ( Carotte 
h Moreau) — A tall herb with flattith, wtiite umbels, cons pi- 
cuous around most sloughs just before mid summer. Seme of the 
rootlets tuberous; base of the stem enlarging, beconing fleshy 
and tuberous towards the end of the season. Commonly abojt 1 m 
high. Leaflets narrowly lanceolate, (0.5)-1.0-(1.5) cm wide, 
about U-6 times as long as wide, most of the lateral nerves 
ending at the bottom of the sinuses. Fruit 2.5-3*0 rrjrj wide and 
SOTievrtiat narrower. Mid summer of somewhat earlier. Open marshy 
places. — swMack-sY, wQ-neBC, US — Var. macula t a — Leaflets 
broader, 1-3 cm wide, ovate to lanceolate, "^^-Utimes as long as 
large. Fruit a bit longer, 3-U iwi long. Prairie CCteau at 
Notre -Dame -de-Lourdes. — NS-sMan, (eUS). 

10. CRYPTOTAENIA DC. HONETa'CRT 

Fruit elongate as in Osmorhiza, but glabrous and not pro- 
longed into a sharp point at base. Involucre lacking. 

1. ^. can a den sis (L.) DC. var. canadensis — Honewort 
( Cerfeuil sa uvage) -- Lea ve s trifoliate, tne leaflets doubly 
serrate. Inflorescence vaguely paniculate. Flowers white. Pe- 
dicels very conspicuously uneven in length. First half of sum- 
mer. Rare in alluvial woods: Portage, Morden. — NB-sMan, US, 
(Eur). 

The Far Eastern var. japonica (Hassk.) Makino has more open 
umbels subtended by better developed involucres and involucels, 
each of 2-5 bractleta. 

11. CAP.UM L. CARA!^L^Y 

Closely related to the preceeding. Involucre typically of 
a single bract which is often lobed. Fruit slightly compressed 
laterally. 

1. C. CARVI L. — Caraway ( Anis , Anis batard) — Leaves 
pinnately dissected into numerous small and linear se^ents. 
Annual. Terminal umbel usually overtopped by tne lateral ones 
by fruiting time. Flowers white. First half of summer. Often 
cultivated and a casual escape to roadsides, shores, snelter- 
belts, etc. — G, NF-(SPM), NS-Q-(0)4^an-Alta-(BC), US, Eur — 
F. RHODOCHRANTHUM A.H. Moore ~ Flowers pinK. Infrequent. — 
NS, Q, Man-Alta. 

12. AEC-OPOriU>t L. 
Fruit without oil tubes, merely dark green between the 
thin nerves . 

CRYPTOTAENIA 166 



1968 Boivin, Flora of Prairie Provinces 107 

1. A. PODOraiARIA L. — Goutweed, Ground -Elder (Herbe aux 
goutteux, Petite Angelique) — Main leaves with 9 leaflets, the 
lateral ones strongly asymetrical. Stoloniferous perennial. 
Leaflets ovate to oblong, often broadly margined in wnite. Flo- 
wers white. Styles rather long, pendent in fruit. Early summer. 
Cultivated and sometimes spreading out of control: Morden. — 
NF, NS, NB-sMan, BC, neUS, Eur. 

13. PFRTDYRlViJA Reichenbach SQUAW-ROOT 

A segregate of Carum , perhaps mainly based on habit. 

1. P. Gairdneri (H. & A.) Mathias ( Atenia montana (Blank.) 
Rydb.) --Squaw-Root — Foliage unusually sparse; main leaves 
about 1 dm long and divided into a few (mostly 5-7) remote leaf- 
lets, these very narrow, l-(3) inni wide, very long, and usually 
deciduous by fruiting time. Perennial from a cluster of tuberous 
roots. Flowers white. Mid summer. Submontane prairies, mainly 
in draws and aroimd bluffs. — sw3-BC, US. 

lU. SIUM L. WATER -?APi>N IP 

Leaves pinnate, otherwise much as in Cicuta. 

1. S. suave Walter (S. cicutifolium Schrank) — Leaves 
pinrate; otherwise quite slriilar to Cicuta maculata with wnich 
it often grows. Reputedly perennial"! Leaflets linear, 1 cm 
wide or less, finely dissected when submerged. Involucre of 
numerous lanceolate and reflexed bracts. Flowers wJriite. All 
summer. Common around sloughs and on marshy shores. — sKack, 
(Aka), NF, NS-BC, (US, Eur). 

1$. AHZTHUM L. 
In this and the following genera the fruit is dorsally com- 
pressed, hence each achene is as \ride as the whole fruit. Fruit 
strongly flattened and narrowly winged marginally. Involucre 
and involucels lacking. 

1. A. ORAVEOLENS L. ~ Dill (Fenouil, Aneth) ~ Stem pale, 
finely striate longitudinally in white and green. Resembles 
Cari:im Carvi, but the flowers yellow and the pedicels nearly uni- 
form in length. Annual. Leaves finely divided into linear to 
filiform segments. Inflorescence most often becoming glandular- 
punctate first in deep green, then in black. Mid to late suirjner. 
Waste places. — Q-Alta, US, Eur. 

16. ANGELICA L. ANGELICA 

Fruit as in Anetnumj leaflets broad and distinct; flowers 
usually wnite. Involucre usually lacking. Involucels small. 

a. Flowers yellow; involucral bracts about as long 

as tne peduncles 3« A. Dawsonii 

aa . Flowers wnite to pinkish; involucre lacking. 

167 ANETHUM 



106 PHYTOLOGIA Vol. 17, no. 2 

b. Leaf racnis straight, its branches 

ascending; 2. A. ap^uta 

bb. Leaf racais geniculate, its branches 

widely spreading to reflexed 1. A. t^enuriexa 

1. A. penuflexa Nutt. var. genuflexa — Primary divisions 
of the lea*f racni£''aaout equally spreading' from tne petiole and 
more or less radiating from its tip. Coarse perennial often 

1 m tall. Involucels of filiform bracts nearly as long as the 
pedicels. Inflorescence densely puberulent, but tne fruit be- 
coming nearly glabrous, with a deep green centre and wnitisn 
wings. Mid summer. Low spots in semi-open forest. — (sAka), 
cAlta-3C, (wUS). 

Stem glabrous and the leaflets eciliate. Involucels snort- 
er than the pedicels. In the Far Eastern vicariant var. multi- 
nervis (Koidz.) Hiroe (including A. refracta F. Schmidt) the 
stem is puberulent above, the leaTlets ciliate and the involu- 
cel longer than the pedicels. 

2. j^. argiita Nutt. (A. Lyallii Watson) — Resembles tne 
above, but qurteglabrous and slightly glaucous, or sligntly 
scabrous. Subterminal leaflets often proximally adnate in the 
manner of the following. Involucels lacking or much reduced. 
Mid summer. Mountane forests, rare: Rockies. — swAlta-seBC, 
wUS. 

3. A. Dawsonii Watson — Mountain -Parsnip — Involucre 
conspicuous,"'*of~Sracts mostly. 2-3 cr long, their margins laci- 
niate and their base i petiolate. Less than 1 m ni^ and gla- 
brous. Leaflets 9-1$, the intermediate ones often sessile and 
cuneate on the distal side, broadly adnate to the rachis on the 
proximal side. Umbel solitary, on a rather elongate peduncle 
2-U dm long. Late spring. Rare in wettish montane woods: Wa- 
terton. — swAlta-se3C, (nwUS). 

17. LEVISTICUM Hill 

Fruit as in Anethum; leaflets broad and distinct; flowers 
yellow . Involu ere present . 

1. L. OFFICINALE V/.D.J. Koch — Lovage ( Herbe k cocnons , 
Celeri batard) — Leaflets lanceolate and entire to rhomboid 
and few -toothed or few-lobed towards the middle. Coarse peren- 
nial about 1 m high. Involucre of broadly membranous bracts. 
Involucels of broadly membranous and fused bractlets. Early 
summer. Sometimes planted and long persisting to slowly spread- 
ing around abandoned homesteads: Langham. — N3, Q-0, 3, (US), 
Eur. 

18. CYMOPTEF.US Raf . 

Each aohene with h bro-id win^s, otherwise similar to Lona - 
tium. 

ANGELICA 168 



1968 Bolvln, Flora of Prairie Provinces 109 

1. C. acaulis (Pursh) Raf . ( Cymopteris acaulig sphalm.) — 
Low herb^with habit of Nusineon and Lomatium , but the leaves 
all basal, the inflorescence more congested, the flowers white 
and the fruits with more v;ings. Perennial with a deeply buried 
fleshy taproot connected to the rosette by a thin and fragile 
pseudoscape. Leaves much dissected into linear lobes. Inflo- 
rescence congested, - puberulent. Involucre lacking. Involu- 
cels palmatifid, the tips of the lobes overtopping the white to 
pinkish flowers. Pedicels of the pistillate flowers veiy short, 
shorter than the ovary and partly adnate to the involucel. Early 
to mid spring. Dry hills, mainly along the major coulees. — 
s^an-sAlta, US. 

Previous reports of Cymopterus montanus (Nutt.) T. * G. we- 
re discussed by Scoggan 1957. The only herbarium sheet located 
was N. Griddle 1033 , Aweme, prairie sbche, 2U mai 1909 (MT; DAO, 
photo) and it turned out to be the rare Lomatium orients le . 

19. LOMATIUM Raf. 
Rather polymorphic. Typically low herbs with a taproot, 
the fruit dorsally flattened and winged around the edge. No in- 
volucre. Fruit nearly always at least as long as its pedicel. 

a. Leaf divided into well defined leaflets...?. L. triter n atum 
aa. Leaf finely divided into numerous small 
ultimate segments, 
b. Ovary and fruit densely puberiilent. 

c. Involucel simple and palmately lobed .. 
2. L. foeniculaceum 

cc. Involucel of several free and slender 

bractlets $, L. Sandbergii 

bb . Glabrous . ~ 

d. Bractlets broadly oblanceolate 1. L. Gous 

dd. Narrowly lanceolate, broadest nearer ~ 

the base. 

e. Stem glabrous 6, L. dissectum 

ee. Densely puberulent. ~ 

f . Stem with at least one pair of 
opposite leaves near the base .. 

U« L. macrocarpum 

ff . Stem with a single leaf in the 
lower half, or sanetimes the 
leaves more numerous and alternate, 
rarely all basal 3. L. orientale 

•^* Jk* SiS}iS> (Watson) C. & R. (L. montanum C. & R.) — Cous 
( Cahous ) — Taproot with a subglobula'r enlargement . Commmonly 
glabrous. Leaves usually all basal. Flowers yellow. Primary 
branches of the inflorescence few and very uneven in fruit. 
Early spring. Dry hillsides, rare: Gypress Hills. — swS, 
nwUS. 

169 LOMATIUM 



110 PHYTOLOGIA Vol. 17, no. 2 

2. L. foenicu lacji^tTi. (Nutt.) CAR. var. fo*3r.icnIac*ji«ri (L. 
daucifolfujn AA.; L. villo3U/T Haf.; Cot^swellia vrflosa (RafT) 
Schultes) — (RacTne biscuit ) — Short vilTdus throuf^ojt. Lea- 
ves all basal ve^yTinely divided, about quadripinnatipartite 
into very numerous and narrow ultimate segjnents. ocape about 
1 dm fiifrh. Bractlets fused into a strone;ly asymetrical involu- 
cel, the latter peltate, palmatilobed and oroadly membranous 
along the edp;es . Flowers yellow. Early to mid sprint. Dry 
hills along major coulees. — swHan-sAlta-(neBC), Ui. 

There are a number of more southern varieties sucn as var. 
fimbriatum (Theobald) stat. n., ssp. fimbriatijm Theobald , 3rit- 
tonia lb: 15, 1966, with pubescent petals. Also var. inyoense 
(Math, gc Const.) stat. n., L. inyoense Katii. & Const. , El Aliso 
3: 120, 1955 in wtiicn the umbels are reduced to a single pedicel. 
^ }, L. orienta_le C . & R . ( Cogswellia orientalis (C . 4 R . ) 
M.E. Jones) — Quite similar to the above, tne leaves not quite 
so deeply divided, the herbage puberulent, but the pedicels and 
fruit glabrous. Stem nearly always bearing one leaf in tne low- 
er half. Flov/ers white. Early spring. Steppes on the bluffs 
of the Souris, rare: Minto, Aweme, Bienfait. — sw}-!an-se3, US. 

Peucedanum nudicaule (Pursh) Nutt. as used oy older authors 
and, presumably, by Kacoiun I89O, usually refers to specimens of 
Lomatiun orientale. 

U. L. macrocarDUin (H. & A.) C. & R. var. macro carj)um (Cogs- 
wellia ma^rocarpa^ THr^ A . ) M.E. Jones) — The stout stem typi- 
cally bearing one pair of opposite leaves near the base. Stem 
1-3 dm high. Herbage lightly to densely villous tcmentose. 
Bractlets fused near the base. Flowers white. Fruit largest, 
narrovrly oblong, 8-I3 mm long. Spring. Steppes and hillsides, 
mainly along coulees. — swMan-BC, US. 

The more southein var. ellipticum (T.& G.) Jepson has longer 
peduncles and fruits. 

5. L. Sandbergii C. & R. — Resembles L. foenic ulaceum 
but merely scabrous puberulent and the leaves smaller, the limb 
5 cm long or less. Stem more or less clearly leafy near the 
base, the leaves alternate. Flowers yellow. Bractlets free, 
few, narrowly elongate, the larger ones often digitate at tip. 
Mid summer. Shale slides above timberline. Waterton. — 
3M&.lta-seBC, nwUS. 

6. L. djLSsectum (Nutt.) Math. 4 Const, var. multifidum 
(Nutt.) I^h r'&^'onst . ( Leptot aenia multifida NuttTT''""^^^^""TalTe3t, 
6-I5 dm hif^ and the leaves mo"sl' "divided, tri pinnate to quadri- 
pinnate with the segments pinnatifid to bipinnatipartite. Stem 
leafy, the leaves alternate, puberulent below, much less densely 
30 to glabrous above, the plant otherwise glabrous or nearly 30. 
Involucels strongly reflexed. PT.owers yellow or purplish. Fruit 
elliptic, 1 cm long or less, nearly sessile or at least longer 
than its pedicel. (Early spring?). Sheltered montane prairies. 
— 3wS-swAlta-sBC, nwUS. 

In the more western typical phase the leaf is less finely 
dissected, the ultimate segments often over 2 mm wide, and the 
fruit is always subsessile. 

LOMATIUM 170 



1968 Boivin, Flora of Prairie Provinces 111 

7. L. triternatum (Pursh) C.& R. var. tritematuin (L. nu- 
dicaule aT. ; L. simplex AA., var. leptophyllum THooker) Mathia¥) 
— With (3)-9-15-(35) distinct leaflets, entire, narrowly lan- 
ceolate to long linear. Stem leafless, thickened below the um- 
bel. At least the stem, and usually the whole plant including 
the fruits, finely puberulent. Flowers yellow. Late spring to 
early summer. Low ground in regions of steppe. — sAlta-sBC, 
nwUS. 

The more western var. platycarpum (Torrey) Boivin is known 
in Canada only from the Okanagan valley. It has a larger fruit, 
the wings being about as \n.de as the body, and a less variable 
leaf, the narrowly linear leaflets being nearly always 9-15 in 
number . 

Despite many Alberta reports of L. n udicaule (Pursh) C. &R., 
only one collection was found under tKat~hame : A .H . Brinkman 
300£, near Beaver Creek, June U, 1928 (NY; DAO, photojl It 
turned out to be L. triternat um. 

20. PASTINAGA L. PARST^P 
Fruit flattened and marginally winged in the manner of Lo- 

matium. Involucre and involucels lacking. 

1. P. SATIVA L. — Parsnip (Pa nais sa uvage ) — Leaves pin- 
nately divided into a few broad lea^ets. Strongly scented herb. 
Stem 1-2 ra high, fistulose, polygonal rather than cylindric. 
Leaflets irregularly serrate, toothed and lobed. Flowers yellow. 
Mid summer. Cultivated and occasionally escaped, sometimes in 
great abundance. — Y-Aka, NF-SBl, NS-BC, US, Eur. 

21. HERACLEUM L. CCW-PARSNIP 
Peripheral flox^ers larger j the petals bifid. Fruit simi- 
lar to Lomatium. 

1. H. la na turn lAx . — Wild Parsnip , Cow -Parsnip (Cigtte) — 
Leaves trifoliate ,^he huge leaflets 1-U dm wide. A huge herb 
in many ways, leaves, stem, umbels, etc. Biennial, 1-2 m high, 
the herbage copiously villous. Flowers white. Early summer. 
Wetter woods, usually semi-open, and frequently in the periphe- 
ral shrubbery. ~ seK-Aka, L-SPM, NS-BC, nUS, (eEur) . 

22. DAUCUS L. CARROT 

Fruit densely covered with bristles borne in rows along 
the nerves of the achene. Peripheral flowers larger and irre- 
gular. 

1. D. CAROTA L. — Wild Carrot , Queen-Anne's Lace (Ca- 
rotte sauvag e) — Umbel with a conspicuous involucre of bracts 
about as long as the rays and pectinately dissected. Coarsely 
hirsute biennial with finely dissected leaves. Umbels stri- 
kingly contracted after flowering and until the maturity of 
the fruits. Flowers white, the central one often pinkish. Mid 
summer. Wild progenitor of the cultivated carrot, occuring 
with ua only as a rare roadside weed: Brandon, Indian Head. — 

171 DAUCUS 



112 PHYTOLOGIA Vol. 17, no. 2 

L, NS-S, BC, US, Eur. 

Foenicul mn vulgare Miller was mentioned for Colinton, Al- 
berta, by Oroh 19Li7, but there is no correspondinf^ apecimen un- 
der that name at DAO and in 19^0 Oroh now mentions the species 
only for B.C. Presumably the original sheet was in the inter- 
val revised to something else. 



FOENICULUM 172 



REVIESr 
otto & Isa Degener 



Bernhard Zepernick of Berlin, Germany, in Baessler-Arch, 
Beitr. VOlkerk. Bd. 15: 329-36$. 1967, deals with "Bemerkungen 
zur Farbei^i der Polynesier" or, roughly translated, "Remarks 
about Polynesian Dye Plants". The article deals with about 100 
species, giving their cori^ct scientific names (without authori- 
ties) and indicating when necessary the synonyms used by about 60 
authors in over 90 articles. The commonest dyes are gained from 
Curcuma longa, Ale\irites moluccana and Morinda citrifolia . The 
author describes the plants used for certain dyes (blue and green 
are rare), in what island groups they aire used, on what materials, 
and their vernacular names . The reviewers wish to alert the 
reader that Solanum nigrtm was native in Polynesia long before 
the coming of the Caucasian explorers, and that Ricinxia communis 
is a common, naturalized weed. Two endemic species of Rubus ex- 
ist in the Hawaiian Islands and hence the name of one should not 
be a synonym of the other. Mr. Zepernick, with aid of five 
tables, has given us in less than $0 pages what the usual author 
might give us in a booklet of 1^0 or more. The study is of 
general interest to botanists as well as anthropologists dea li ng 
with the islands of the Pacific. 



NOTES ON NEW AND NOTEWDRTHI PIANTS. L 
Harold N, Moldenke 



CITHAREXTLUM HIRTELLUU var. GUATMALENSE Moldenke, var. nov. 

Haec varietas a forma typica speciei laminis folioram subtus 
in reticulo venule rum parcissime setulosis recedlt. 

This variety differs from the typical form of the species in 
having the vein and veinlet reticulation on the lower leaf- 
surface very sparsely setulose with whitish, stiff, straight, 
unbranched, sharp-pointed, spi^ading hairs, and the lamina it- 
self glabrate. 

The type of the variety was collected by Julian Alfred 
Steyermark ( no. UlSlS ) along the Rio Yameja, at about 00 meters 
altitude, Cerro San Gil, Izabal, Guatemala, on December 2k, 
I9I4I, and is deposited in the Britton Herbarium at the New York 
Botanical Garden. 

LYSIMACELA QUADRIFOLIA f . RUBESCENS Moldenke, f , nov. 

Haec fonna a forma typica speciei caulibus foliisque in statu 

113 



llJi P H Y T L G I 1 Vol. 17, no. 2 

Juvenile plusminusve rubris recedlt. 

This form differe from the typical form of the speciea in 
having the upper portions of its stems and all the uppier leaves, 
or sometimes the entire plant, red when young. 

The type of the form was collected by Alma Lance Uoldenke and 
Harold Norman k'oldenke ( no. 21x25$ ) on an open roadbank at Moose 
Meadow, Tolland County, Connecticut, on May 31, 1968, and is de- 
posited in the herbarium of the Dotanisk Institut at JUrhvis Uni- 
versitet, Aarhus, Denmark. This form sometimes grows in very 
extensive purestand colonies, while at other times it is inter- 
spersed with the typical green form of the species in precisely 
the same envirormental conditions of soil, drainage, exposure 
to sunlight, etc. The type irtiere the entire plant is red from 
top to base was not collected, but occurred in purestand 
colonies on roadbanks only a few miles from where the tyj)e 
specimens were gathered. 

There is another form of the species known, L. quadrifoHa f . 
variegata (Peck) House, in which the tips of the petals sire 
orange. It is described in Bull. N, Y. State kus. Ii7: 157 
(189U) and 2$li: $$9 (1921;). 

PRIVA lAPPULACEA f. ALBIFLORA Moldenke, f . nov. 

Haec forma a forma typica special coorollis albis recedit. 
This form differs from the typical form of the species in 
having irtiite corollas. 

The type of the form was collected by Walter H. Lewis, Jr., 
John Duncan Dwyer, T. S. ELias, and K. R. Robertson ( no. £26) 
at the edge of a river and adjacent rainforest and railway, 
Changuinola to 5 miles south at the Junction of Rio Changuinola 
and Rio Terebe, at an altitude of 100 to 200 feet, Bocas del 
Tore, Panama, between December 17 and 19, 1966, and is deposited 
in the herbarium of the Missouri Botanical Garden at St. Louis, 

SVIDA CONTROVERSA (Hemsl.) Moldenke, comb, nov, 

Cornus cont rovers a Hensl. in Curtis, Bot. Mag. 135 [ser. h, 
5]: pi. 8261. 1909; Kew Bull. Misc. Inf. 1909: 331. 1909. 

XYLOSTEON MORROffI (A. Gray) Moldenke, comb, nov. 

Lonicera morrowi A, Gray in Perry, Narr, Exped. Chin. Jap. 
2: 313. 1856. 



ADDITIONAL NOTES ON THE GENUS VITEX . IX 
Harold N, Uoldenke 



VITKX TRIFOLIA var. SIMPLICIFOLIA Cham. 

Additional bibliography: Moldenke, Phytologia 17: 11 — 13, U5, 



1968 Moldenke, Notes on Vltex 115 

ii7, 50, & 51—56. 1968. 

Van Steenis (1957) prefers to regard this plant as a subspe- 
cies, Tihich he calls V. tidfolia subsp. litoralis . He comments 
that the plant was considered as a valid species by Thunberg and 
by Blanco, later as a variety by Chamisso, Schauer, Makino, Rid- 
ley, and Bentham, "This evaluation as a variety has been main- 
tained by later monographers (Lam & Bakh\iizen van den Brink, 

Merrill, and Moldenke) , Backer , Comer , and following 

him Backer & Ueeuse have again treated it as a good, dis- 
tinct species. And Comer has taken great pains to give arguments 

for this view. Contrary to Ridley who suggested to have 

seen it change into normal V. trifolia after transplantation to 
Singapore, Comer maintains that it maintains its habit and 
characters in cultivation and is no mere phenotype. He trans- 
planted ten specimens to the Botanic Gardens, Singapore, where he 
also had living shrubs of V. trifolia and V. negundo, and has 
found that they retain their habit. As to the constancy of that 
character there remains hence little doubt, though additional ex- 
peidjnents in raising inland plamts from seed of the prostrate 
form and crossing it with V. trifolia are still a desiderat\im. In 
addition Comer assumes to have found differences with V_. tri - 
folia in the corolla, fruiting calyx, and the fruit. I have 
tried to verify these differences with many sheets preserved at 
Leyden but I cannot corroborate these statements. The fruits of 
V, trifolia and V. ovata offer no differences in size, shape, and 
internal tissue structure. That the inflorescences of V. ovata 
are smaller than the average size in V. trifolia I deem not sig- 
nificant, as they are bome on small side-branches. The only 
chsa*acteristics holding are vegetative in nature, viz the typical 
prostrate, rooting, runner-like branches, and the obovate, small, 
simple leaves, and geographic: its exclusive growth on the sandy 
beach." 

Ohwi (1965) gives the distribution of the variety as "Honshu, 
Shikoku, Kyushu. — Korea, Bonins, Ryukyus, Formosa to se. Asia, 
Pacific Islands, and Australia." Bryan says that on Johnston Is- 
land it was "planted by man or introduced by some other means 
since 1923". Taniguti (I963) records it from Hemizima Island, 
Japan, while Hatusima (1962) records it frcm the Amaml Islands in 
the Ryukyu Archipelago, 

Nobuhara (1967) tells us that "The shorter the distance to the 
coast line, the less the cover of Canavalia and the more, to some 
extent, that of Vitex rotundi folia expands," Nobuhara, Okada, & 
Fujihira (1962) report that our plant has average tolerability 
toward salt spray from typhoons. Wilson found is common on Quel- 
part Island, while Chiao refers to it as a "rare bush along sea- 
shore" in Shantung and Ching describes it as "a low dense sand- 
binding shrubby perennial herb on active sand, up to 1 1/2 ft. 
tall" in Chekiang. 

A letter to me from Berta Cerin, dated April 29, 1962, announces 
that she plans to study the chemical constituents of this plant. 

Additional vernacular names recorded for the plant are "hii-po- 



116 P H Y T L I A Vol. 17, no. 2 

kiu", •♦hamacfi" , "hamag5", "hama-gB" ["hama" - the sea], "hiama- 
sikimi", "kolokolo-kahakai", •taoa quite sage", "pch-po-kiu", 
"pohinahina" , •^olinalina" , "simple-leaf chiaaW-tree", 
"Bimple-leaif shrub chaate-tree", ajid "taiT»an-haniag8", 

The Lam (192U) reference given in the bibliography of this 
plant is often dated "1925", but the latter date is merely th« 
title-page date for the volume; the page involved actually waa 
issued in 192U. Van Steenis (195?) gives the date of publication 
of Bentham's name (IdTOJ for this taxon as "1376". Hara (19U3) 
cites Merrill's Enum. Philip. PI. (1923) as page "3U7" in error. 
The Hooker & Amott (1336) references in the bibliography and 
list of illustrations listed previcjusly are sometimes dated "iSUiy 
but pages 193 to 288 and plates UO to $9 of this work were actual- 
ly issued in 1336, 

Lam (I92U) cites Kotara s.n. from the Bonin Islands and Koch 
s.n. from Dutch New Guinea. Hatusima (I966) cites his no. 28$6$ 
and gives the general distribution of the variety aa "Japan to 
Malaysia, Australia and Polynesia". Li (1963) cites Faurie U$2 & 
1169 , Gressitt £23, A_. Henry s.n. , Oldham 382 , Owatarl s.n. . 
Price k9h & 650, Takenouchl s.n. , E. H^ Wilson 10973 , and Yamano- 
to s.n. from Formosa. Miquel (18707 cites Oldham 1 [specimen?] , 
Bttrger 7 [specimens?], Keiske 1, Maximowicz 1, and Siebold 3« 

The A. Henry 12302 and Saint John ^ Fosberg 16976 , distributed 
as this variety, are actually var. subtrieecta (Kuntze) Moldenke. 
On the other hand, many collections of var, simplicifolia have 
been distjributed in herbaria as typical V. trifolla L, 

Additional citations: CHINA: Chekiang: Ghlao iWiS [Herb. Univ. 
Nanking IkShh] (Bi, ¥--lli27017) ; R. C. Ching I967 (W— 12li6828) . 
Shantung: Chiao 277U (W— l59623li) . CHINESE COASTAL ISUNDS: Hai- 
nan: Fung 20500 (Mi)} Liang 62926 (W— 1670956) . Lantau: McClure 
s.n. [Herb. Lingnan Univ. 13095] (W--1298l0)i Taam 1702 (W— 
22U;609); Tsang s.n. [Herb. Lingnan Univ. 166U9] (W--12U9810) . 
HONGKONG: Bodinier 679 (W— 2ii9712l4); C. Wright s.n. [Hong Kong] 
(W— Ui911). THAILAND: Larsen . Smitinand , & Wamcke 121^6 (Ac, Rf ) . 
INDOCHINA: Tonkin: P^telot 317 (W— 1716990) . KOREA: R. K^ Smith 
s.n. [Aug. 23, 1932] (Bi)} Mrs. R. K^ Saith s.n. [0-10-31] (W— 
1757013) . KOREAN COASTAL ISUNDS: Quelpart: In-cho 11 21^ (Mi, S)} 
E. H. Wilson 9392 (W~-105U.88) . WESTERN PACIFIC ISLANDS: JAPAN: 
Anashima: Koidztmii s.n. [5.8.1922] (Mi). Honshiu: Collector un- 
determined 361; (W— 73901), s.n. [Sagami, 17 Juli 1910] (W— 
1133035); Ichikawa 200661 [122] (W— 13U7laU) ; Kirono 762 (S, W— 
233630U) } Maruyama & Okamoto I6OO (W— 23l576ii) } Maximowicz 90 
(W— 73900)} Sasaki & Tagasl 606 (Mi, W— 2156562)} Savatier s.n. 
[Yokaska] (W--2li97127) . Kiushiu: Hurusawa 202 (W— 2038128)} Take- 
nouchl 1728 (W — 1271675) . Shikoku: Collector undetermined s.n. 
[Susaki, Tosa, Aug. 16, I892] (W— 206183 ) . FOxHUOSA : Gressitt 523 
(N)} A. Henry s.n. [Takow] (W— U55205)} Takenouchl s.n. [Aug. 5, 



1968 Moldanke, Notes on Yitex 117 

19liO] (W~2063U01); E. H. Wilson 10978 (W--1052371) . PHILIPPINE 
ISLANDS: Luzon: Haenke s.n. [Luzon, 1792] (Bi). Mindoro: H. H_. 
Bartlett 13708 , in part (Mi). Sibuyan: Kljner 12135 (Bi). BONIN 
ISLANDS: Anijlma: Kondo 115 (Bi). Chichijlma: Kondo 33 (Bi) , 
Imajima: H. L. Porter 3 (Mi). Island undetermined: C, Wright s. 
n. [Bonin Islands] (W~73896) . VOLCANO ISLANDS: Itrojlma: H. L. 
Porter 3 (W-— 19Uli269) . MELANESIA: NEW HEBRIDES: Aneityum: Kajew- 
ski 690 (Bi). AUSTRALIAN REGION: AUSTRALIA: Queensland: Brass 
19l9" TBi) » POLYNESIA: HAWAIIAN ISUNDS: Hawaii: A. F. Judd s.n, 
(Bi). Kauai: F. R. Fosberg 1273ii (Bi, Bi) ; A, A. Heller 2731 
(Bi, Ms — 30950); Saint John , Hosaka, Hume , Inafuku, Lindsay , Ma- 
suhara , Mitchell , & Wong lOSlJ. (Bi) j C. Skottsberg 10^9 (Bi) . 
Lamai: G. C. Munro 90 (Bi), 122 (Bi), s.n. [Kaena Point, 12/2/ 
1$] (Bi). Maxii: Topping s.n. [0. Degener 950li] (Bi, Lb~l5779, 
Mi). Molokai: 0. Degener 9506 (Bi, Mi), 9507 (Bi) . Niihau: 
Handy s.n. [Aug. lli, 1931] (Bi) i ^1s9jL Stokes s.n. [Kiekie] 
(Bi). Oahu: 0. Degener 10018 (Bi, Mi), 112U5 (Bi), 112U7 (Bi); 
F. R. Fosberg 8881 (Bi), IO36O (Bi), 131U8 (Bi, Bi), llqSU (Bi); 
J. A. Harris C.2i;2lUO (Bi), C.2J4220I (Bi); Hathaway & Caindec 
139 (Bi); Meebold s.n. [Pauraalu, May 1932] (Bi); H. N. Moldenke 
"^^08 (Bi, Ca, Fg, Mi); jJ, W. Moore s.n. [July li+, 1929] (Bi, 
Bi); J. F. Rock Ji3 (Bi, Bi); £• F. 2^ Stokes s.n. [Alaopjq)a, 
Jvme 1-2, 1920] (Bi); Topping 3012 (Bi); M. M. Townaend s.n, 
[Oct. 20, I9I1O] (Mi); D. P. Wilder s.n, [Leilehua Plain, 1912] 
(Bi) . Island undetermined: 0. Degener 112U6 (Bi); C, N. Forbes 
s.n, (Bi); Hillebrand & Lydgate s.n. (Bi); G, P^ Wilder s.n. 
[1913] (Bi). CULTIVATED: Johnston Island: E, H« Biyan s.n, [Au- 
gust 30, 19lih] (Bi) . 

VITEX TRIFOLIA var. SIMPLICIFOLIA f . ALBIFLORA (I. Matsuraura) 
Moldenke 
Additional bibliography: Moldenke, Phytologia 6: 197. 1958; 
Moldenke, R6sum6 173, 388, & U79. 1959. 

VITEX TRIFOLIA var. SUBTRISECTA (Kuntze) Moldenke 

Additional bibliography: Warb. in Engl,, Bot, Jahrb. 13: 1^29. 
I89I; Kuntze, Rev. Gen. PI. 2: 510 & 5ll. 1891; Mak., 111. Fl. 
Nipp. 186. I9UO; Moldenke, Phytologia 3: 178. 19U9; Moldenke in 
Humbert Fl. Madag. 17U: 72, 82, & 273. 1956; Moldenke, Phyiiolo- 
gia 8: 88—90. I96I; Moldenke, Biol, Abstr. 37: 1062. 1962; Hock- 
ing, Excerpt. Bot, A,6: 53li. 1963; Neal, In Gard. Hawaii, ed. 2, 
727 & 728. 1965. 

It is worth recording here that Makino's original Japanese 
description of his var, heterophylla has been rendered in Latin 
by Hara (I9I18) as "Folia aut simplicia aut tripartita". The co- 
rolla is described as "purple" on M, S, Clemens 11067bis and on 
Native collector PI. 11^9 [Herb, Roy, Forest Dept. 3567], "reddish- 



U8 PHTTOLOOIA Vol. 17, no. 2 

purple" on S^ K^ I^ 270 , and '^lue" on Rock 78 3Q . Rock refera 
to the plant as a "common shrub along banks" in the Southeastern 
Shan States of Bunaa; it is also said to be conmon on the plains 
in Thailand, where the bark and roots are employed as a febrifuge 
and where the plant is known as "phi-suae" . The plant has also 
been collected in sandy areas behind the beach on outer sandhills 
in Thailauid, at 3000 feat altitude in Mew Guinea, and between 
3000 and UOOO feet altitude in Yllnnanl It has been collected in 
anthesis in February and June, 

R. K, Godfrey $9186 bears a notation "locally naturalized in 
sandy lots" in Pinellas County, Florida. P_. 0^ Schallert 23077 
is var, variegata Moldenke in most herbaria, but the specimen 
of this number preserved in the Berlin herbarius shows no vaii.«ga- 
tion, although the leaf -edges are irregularly turned over, «hich 
may be an indication of variegation. 

Additional citations: FLORIDA: Pinellas Co.: R. jU Godfrey 
59186 (Hi--l51i7l8) . BURMA: Shan States: £. F. C. Rock 232$ (W— 
12111807). CHINA: Tttnnan: A. Henry 12302 (W—U59013) ', £. F. C. 
Rock 2669a (W— 121ii89l) , 2969 (W—12132$2) , 7838 (W— 133211iOyT 
CHINESE COASTAL ISLANDS: Hainan: S. K. Lau 270 (W~162911a2) . 
THAILAND: Larsen , Smitinand , & Wamcke I3ZL (Ac, Plf ) ; Native col- 
lector PI .11^9 [Herb, Rojr. Forest Dept. 3567] (W— 2061i79$) . WES- 
TERN PACIFIC ISLANDS: RYUKYU ARCHIPELAGO: OKINAWAN ISUNDS: Okin- 
awa: Field & Loew 21v (Mi) . PHILIPPINE ISLANDS: Mindoro: H. H. 
Bartlett 13708 , in part (Mi). INDONESU: GREATEK SUNDA ISLANDS: 
Sumatra: Hamel & Toroes $$1 (Ui)j Schlffner 2h^h (Bl); Toroea 910 
(Mi); Yates ^ (Mi). MELANESU: NEW GUINEA: Northeastern New 
Guinea: M. K. Clemens 11067 bis (Mi), U1503 (Mi). SOLOMON IS- 
LANDS: Bougainville: Waterhouse 60 [Herb. Mus. Yale Sch. Forest. 
2266U] (Bi, Bi). lASAWA FIJI ISLANDS: Vitl Levu: J. W. Gillespie 
U38O (Bl)} A. C. Smith U$$9 (Bi), 6078 (Bi). POLYNESIA: LINE IS- 
LANDS: Palmyra: E. Y, Dawson 1982$ (Bi). MARQUESAS ISLANDS: Is- 
land undetermined: Quayle 1281 [ 2181] (Bi). TUAMDTU ISLANDS: 
Anaa: H. Saint John 1J;2$2 (Bi) . Rarola: Doty & Newhouse 11721; 
(Bi). SOCIETY ISUNDS: Raiatea: J. W. Moore 696 (Bi). AUSTRAL 
ISLANDS: Rimatara: Saint John & Fosberg 16976 (Bi). Rurutu: 
Chapin 8g3 (Bi); F, R. Fosberg II98I (Bi)} H. Saint John 16573 
(Bi)} A. M. Stokes 1 (Bi). CULTIVATED: Baker Island: E. H. Bry- 
an 1315 (Bl) . Florida: P. 0. Schallert 23077 , in part (B) . 
Hawaiian Islands: J, F. Rock s.n. [S. Kona, April 28, 1957] (Bl). 
Johnston Island: K. P. Fosberg 15 (Bi), Marshall Islands: F. R. 
Fosberg 36709 (Bi). 

VITEX TRIFOLIA var. SUBTRISECTA f . ALBIFLORA Moldenke 

Bibliography: Moldenke, Phytologia 8: 90—91. 1961; Moldenke, 
Biol. Abstr. 37: 1062. 1962} Hocking, Exceipt. Bot. A.6: 53U. 
1963. 



1968 Moldenke, Notes on Vitex 119 

Additional citations: POLYNESIA: AUSTRiO. ISUNDS: Rurutu: H, 
Saint John 1670^ (Bi — isotype) , 

VITEX TRIFOLIA var. VAKEBIAIA Moldenke 

Synonymy: Vitex trl folia variegata [Moldenke] ex Lord, Shrubs 
& Trees Austral. Gard., rev. ed., 232, 196U. 

Additional & emended bibliography: Neal, In Card. Hawaii, ed, 
1, 6Ia. l9U8i L, H. Bailey, Man, Cvlt, PI., ed. 2, QUk & mli, 
I9li9i Kuck & Tongg, Mod. Trop, Gard, 77 & 236. 1955} Moldenke, 
Phytologia 8: 91. 1961; Menninger, Seaside PI. 151) & 155. 196U; 
E. E, Lord, Shrubs & Trees Austral, Gard., rev. ed., 232, 196Uj 
Neal, In Gard. Hawaii, ed. 2, 728. 1965} Moldenke, R6sum6 Suppl. 
15: 15. 1967} Moldenke, Phytologia 17: 52. 1968. 

Illustrations: Menninger, Seaside PI. pi. 223. 196U. 

Lord (I96U) describes this variety as " Vitex trlfolia variega - 
ta with the leaves broadly cream-margined, a very showy shrub", 
and recommends it for coastal areas in Australia, Kuck k Tongg 
(1955) state that the plant is very wind-resistant. 

The Berlin specimen of P. 0, Schallert 23007 does not show any 
variegation, although its leaf -margins are turned over, and is 
cited by me herein under V, trifolia var, subtrisecta (Kuntze) 
Moldenke, It is very possible that the turning over of the leaf- 
margins is an indication that they were variegated there and that 
the specimen should, therefore, be cited here under var, variegata . 

Additional citations: CULTIVATED: Florida: H, N, Moldenke 
2li09U (Ac, Rf). Hawaiian Islands: Ito s,n, [Schofield, May 1936] 
(Bi); Cj^ Sj^ Jizdd s.n. [Puunene, Feb. 8, I9U0] (Bi)} Neal s,h. 
[Nov, 19, I9U+] (Bi), s.n, [July 9, 19U5] (Bi)} J, A, Price 3,n. 
[May 10, 19ii3] (Bi) . 

VITEX TRIPINNATA (Lour,) Merr, 

Additional & emended bibliography: Jacks, in Hook, f, & Jacks,, 
Ind, Kew,, pr, 1, 1: 582 (1893) and 2: IO36 & 1121, 1895} A, W. 
Hill, Ind. Kew, Suppl, 6: 219 (1926) and 9: 297 & 298. 1938} 
Merr, Sl Chun, Sxinyatsenia 5: 178, I9UO; Jacks, in Hook, f , & 
Jacks,, Ind, Kew,, pr, 2, 1: 582 (I9U6) and 2: IO36 & 1121 (I9I16) 
and pr, 3, 1: 582 (I960) and 2: IO36 & 1121, I96O} Moldenke, Phy- 
tologia 8: 91—92, I96I} Moldenke, Biol, Abstr, 37: 1062. 1962} 
Hocking, Excerpt. Bot. A, 6: 53U. 1963. 

Recent collectors have found this plant growing in evergreen 
forests, at 150 meters altitude, fruiting in August, The corollas 
are described as having been "yellow" on Clemens & Clemens 339h» 

The Bejaud 223 , in part, in the Berlin herbarium, cited by me 
previously as V_, tripinnata , proves actually to be var, clemensorum 
Moldenke . 

Additional citations: CHINESE COASTAL ISLANDS: Hainan: How 
72997 (Bi). THAILAND: Larsen, Smtinand, & Wamcke 1385 (Ac, Rf ) . 
INDOCHINA: Annam: Clemens & Clemens 339h (W— ll;2 71^99 ) . Tonkin: 
Pgtelot 6398 (W— 1759U57) , 61il9 (W— 1759li67) . State undetermined: 
Eberhardinil32 [Hoa-Binh] (W~Si97092) . 



120 PHYTOLOOIA Vol. 17, no. 2 

VTTEX TfOPINNATA var. CLmniSOIOJV Moldenke 

Bibliography: Uoldenke, Phytologia 8: 92. 1961; Uoldenke, fciol. 
Abstr. 37: 1062. 1962 j Hocking, Excerpt. Bot. A.6: 53J4. 1963. 

The Berlin specimen of Bejaud 223, previously cited by me aa 
typical V_, tripinnata , has been re-exainined and proves to be var. 
clemensor'um . It is, however, mixed with somethdng not verben«»- 
ceous. 

Additional citations: INDOCHIHA: Cambodia: bejaud 223 , in 
part (B). 

VTTEX TRISTIS S. Elliot 

Additional bibliography: Durand ^ Jacks., Ind, Kew. Suppl. 1, 
pr. 1, U57 (1906) and pr. 2, U57. 19U.i Moldenke in Humbert, Fl. 
Madag. 17U: 7h, 113—11^, & 273, fig. 17 (1). 1956; Moldenke, 
Phytologia 6: 200—201. 1958; Moldenke, Rfesumfe 157 & U79. 1959; 
Durand & Jacks., Ind. Kew. Suppl. 1, pr. 3, 1457. 1959. 

Illustrations: lioldenke in Humbert, Fl. Madag. 17U: 115, fig. 
17 (1). 1956. 

VTTEX UBANDHENSIS A, Chev. 

Additional bibliography: Prain, Ind, Zen, Suppl, 5, pr. 1, 
273. 1921; Moldenke, Phytologia 6: 201. 1958; Moldenke, Rfisumfe 
II4O & U79. 1959; Prain, Ind. Kew, Suppl, 5, pr, 2, 273. i960. 

VITEX UMBROSA Sw, 

Additional synonymy: Nephrandra dubia Willd, in Cothen,, 
Disp, Veg. 8, 1790. 

Additional bibliography: J, F, Qnel. in L,, Syst, Nat. Veg., 
ed. 13, pr, 1, 2: 963 (1789) and pr, 2, 2: 9U6 & 963. 1796; 
Pers,, Sp, PI, 3: 361, 1819; Steud,, Norn, Bot, Phan,, ed. 1, 
888, 1821; Griseb,, Cat, PI, Cub, 216, 1866; Jacks, in Hook, f, 
& Jacks,, Ind. Kew,, pr, 1, 2: 308 (1891) & 121ii (1895) and pr, 
2, 2: 308 & I21U. 19li6; Asprey & Robbins, Ecol, Monog, 23: 385 & 
I4II, fig, 20. 1953; Hocking, Diet, Tenns Pharmacog, 32, 1955; 
Jacks, in Hook, f. & Jacks.. Ind. Kew,, pr, 3, 2: 308 & 12lU, 
I96O; Moldenke, Phytologia 8: 92, I96I; Moldenke, R6sum4 Suppl, 

15: a, 1967. 

Additional illustrations: Asprey & Robbins, Ecol, Monog, 23: 
385, fig. 20. 1953. 

Recent collectors describe this species as a tree, 12 m. 
tall, the stem diameter 50 cm, at breast height, the flowers 
scented, and the fruit orange, growing on steep wooded hillsides, 
at 1000 feet altitude. The corolla is described as "purple with 
yellow blotch at top of lower lip" on Steam 976. Hocking 
(1955) reports the common names "boxwood" and "South American 
boxwood" for this species. 

Additional citations: JAMAICA: Proctor 19783 (N); Steam 976 
(S), 



,6\. \^ 



r« o 



PHYTOLOGIA 

Designed to expedite botanical publication 
Vol. 17 September, 1968 No. 3 

LIBRAR 

CONTENTS SEP 24 1968 

NEW YORK 
BOTANICAL GARDE 

MYINT, T., & WARD, D. B., A taxonomic revision of the genus 

Bonamia (Convoliulaceae) 121 

MOLDENKE, H. N., Additional notes on the genus Vitex. X 240 

ViTJNDERLIN, R. P., A note on Bauhinia hagenbeckii Harms 245 

MOURA, C. A. F. de, .4 new species of Piriqueta Aublet (Turneraceae) 

from Mato\Grosso, Brazil 247 



Published by Harold N. Moldenke and Alma L. Moldenke 

303 Parkside Road 
Plainfield, New Jersey 07060 
U.S.A. 

Price of this number, $2; per volume, $6.75, in advance, 
or $7 at close of volume 

V 



A TAXONOMIC REVISION OF THE GENUS 
BONAMIA (CONVOLVULACEAE) 

Tin Myint and Daniel B. Ward ^ 

Table of Contents 

Page 

List of Figures 123 

List of Maps 123 

Introduction 124 

History 125 

Morphology 127 

Systematic Treatment 134 

Bonamia 134 

Key to Sections of Bonamia 136 

Regional Keys 143 

Key to the African and Asian Species 143 

Key to the Australian Species 144 

Key to the American and Hawaiian Species 145 

I. Section: Bonamia 149 

1. Bonamia alternifolia 150 

2. Bonamia spectabilis 153 

3. Bonamia d ens i flora 156 

4. Bonamia thunbergiana 157 

5. Bonamia mossambicensis 160 

6. Bonamia cordata 162 

7. Bonamia semidigyna 163 

8. Bonamia elegans 168 

9. Bonamia dietrichiana 169 

10. Bonamia menziesii 170 



1 Art and Science University, Mandalay, Burma; and the 
University of Florida, Gainesville, Florida. 

This study is derived from a doctoral dissertation 
written by the senior author, University of Florida, 1964, 
under the direction of the junior author. The authors' 
gratitude is hereby expressed to the Government of the Union 
of Burma, who met important expenses by a State Scholarship 
grant, to the curators of those herbaria who generously sent 
their specimens, including types, for examination, to the 
Center for Tropical Agriculture, University of Florida, for 
preparation of the typescript, and to the Department of 
Botany, University of Florida, for meeting the direct cost 
of publication. 

121 



122 P H Y T L G I A Vol. 17, no. 3 

11. Fjonamia grand i flora 1^4 

12. Bonamla flliiitiCf) 177 

13. l'.ont?nn"a su3;jliiirf'.-j 178 

14. Bonamiii ovnl i fo] i;i 379 

15. Bonamia mult i caul is 181 

16. nonarm' a soricca ^82 

17. Bonamia holiviana 1^^ 

18. Bnnamia ho] li i 1^5 

19. Bonamia mgri]ioidns 186 

20. Bonamia broy i pfd ] crdlata 188 

21. Bonamia ffrrugiir-a 191 

22. Bonamia umbr^lr-ita • • • ^''^ 

23. Bonam i a sphar-rocf^i'hala 193 

24. Bonamia kuhlmanni i 195 

25. Bonamia ponivJana 196 

II. Section: Brewr-ria 197 

26. Bonamia linearis 198 

27. Bonamia oblongifolia 200 

28. Bonamia brevlfolia 203 

29. Bonamia media 204 

30. Bonamia rosea 209 

31. Bonamia pannosa 211 

32. Bonami a volutina 214 

III. Section: Trichantha 215 

33. Bonamia ti-jchantha 216 

34. Bonamia balansae 221 

35. Bonamia corumbaensis 222 

36. Bonamia agrostopolis 224 

37. Bonamia burchellii 225 

38. Bonamia tomentosa. 228 

39. Bonamia subsessilis 229 

40. Bonamia mattogross ens is 230 

Little-Known Species 231 

41. Bonamia abscissa 231 

42. Bonamia boivinii 232 

43. Bonamia langsdorffii 232 

44. Bonamia capitata 233 

45. Bonamia sedderoides 233 

Doubtful and Excluded Species 233 

Bibliography 237 

Appendix: New Names and Combinations 239 



1968 



Myint & Ward, Revision of Bonamia 



123 



Figure 1. 
Figure 2. 
Figure 3. 

Figure 4. 

Figure 5. 
Figure 6. 



List of Figures 

Variations in sizes and shapes of leaves in 
Bonamia spectabilis 

Leaf shapes and sizes in the two varieties 
of B. menziesii 



Leaf shape, inflorescence and floral parts 

of B. breviped icellata , and inflorescence 
and floral parts of B. maripoides . . . . 

Variations in sizes and shapes of leaves in 
B. media , B. brevifolia , B. oblongifolia 
and B. linearis 

Variations in sizes and shapes of leaves in 
B. pannosa , B. dietrichiana and B^ rosea 

Leaf shapes and sizes in Bonamia trichantha . 



Page 

155 

173 

189 

201 

213 
219 



Map 1. 

Map 2. 
Map 3. 



Map 4. 
Map 5. 

Map 6. 



Map 
Map 


7. 
8. 


Map 


9. 


Map 


10 


Map 
Map 


11 
12 


Map 


13 


Map 


14 



List of Maps 

Page 
Distribution of B. alternifolia , B. specta - 
bilis , B. mossambicensis , and B. velutina . . 152 

Distribution of B. thunbergiana 159 

Distribution of B. semidigyna var. semidigyna 

and var. farinacea , B. elegans , and B. 

dietrichiana 164 

Distribution of B. menziesii var. menziesii 

and var. rockii 171 

Distribution of B. grand i flora , B. elliptica , 

B. sulphurea , B. ovalifolia, B. multicaulis , 

and B. breviped icellata 176 

Distribution of B. boliviana, B. holtii , and 

B. sericea var. sericea and var. lati folia . 183 
Distribution of B. maripoides and B. ferruginea . 187 
Distribution of B. umbellata , B. sphaerocephala , 

B. kuhlmannii , and B. peruviana 194 

Distribution of B. linearis , B. brevifolia , and 

B. oblongifolia 199 

Distribution of B. media var. media, var. villosa 

and var. emarginata 206 

Distribution of B. pannosa and B. rosea 210 

Distribution of B. trichantha var. trichantha , 

var. oblonga , var. ovata f. ovata and f. 

glabrata 217 

Distribution of B. balansae , B. corumbaensis 

and B. agrostopolis 223 

Distribution of B. burchellii , B. tomcntosa , 

B. subs ess ilis and B. mattogross ens is . . . . 227 



12U P H I T L G I A Vol. 17, no. 3 

Introduction 

The genus Bonamia belongs to the Convolvulaceae, a family of 
flowering plants. According to the concept presented in this 
study, it is a genus of forty-five species cind eleven varieties. 
Bonamia is a fairly large genus compared with some of its close 
relatives, Stylisma (represented by six species), Calycobolus 
(represented by eleven or twelve species) and Seddera frepresen- 
ted by about fifteen or twenty species). In contrast to the 
narrow distribution of these genera, Bonamia occurs throughout 
the tropical and warm temperate regions of both hemispheres, with 
a concentration of species in South America, Australia and Mada- 
gascar. Several species are known only by type collections or 
only from type localities and are poorly represented in the 
herbaria of the world. Several others are known only by a modest 
number of specimens. Only fourteen species are known from ten 
or more collections, and only five species are not restricted 
to a narrow geographical region. 

The only synopsis of the genus Bonamia , that of H. Hallier 
(1897), contained twenty-eight species, one of which definitely 
belongs to the genus Seddera and another to the genus Metaporana ; 
no key was given. Recent studies of the genus (van Ooststroom, 
1954, O'Donell, 1959, Verdcourt, 1963 and Myint, 1968) deal only 
with a few species of particular areas or countries. Other 're- 
cent students (Meeuse, 1957, Wilson, 1960, and Shinners , 1962), 
have applied the name Bonamia in a somewhat broader sense by in- 
clusion of some species here considered as belonging to related 
genera. In contrast to these authors, Roberty (1952) breaks Bo- 
namia into more than one genus. These conflicting treatments 
create doubt as to generic limits and invite a thorough investi- 
gation of the entire genus. That some clarification of the species 
and varieties comprising the genus Bonamia is needed is evident 
from the large number of misidentified or misplaced specimens ex- 
tant in most herbaria. With the addition of many species to 
the original genus, and due to the inclusion of several species of 
related genera, it becomes desirable to redefine the generic limits 
of Bonamia , to form three sections within the genus, to evaluate 
some characters that have not been used in its classification, and 
to determine boundaries of several species which were inadequately 
described and are poorly known. 

Bonamia is characterized technically by the possession of 
free or partially free styles, nonaccrescent sepals, and ovate, 
obovate or ovate-cordate cotyledons. It is not surprising that 
Asa Gray, recognizing the overall similarity of Bonamia and Sty- 
lisma , put the members of the latter in Bonamia . However, a re- 
cent monographic study of Stylisma by the senior author (Myint, 1966) 
together with additional information gathered during the present 
investigation, strongly indicates the feasibility and desirability 
of treating them as separate genera, while admitting the existence 
of connection between them. 



1968 Ityint & viard, Revision of Bonamia 125 

In the present study, one new species, two new varieties and 
one new form are described; three new combinations are made; and 
three sections are proposed. A key to sections, a general key 
to all species, and three regional keys to species are presented. 
All specimens (assigned to the genus Bonamia and its related genera) 
available from the following herbaria were examined. The abbre- 
viations are listed here according to Lanjouw and Stafleu (1964). 

A- Arnold Arboretum, Cambridge, Massachusetts. 

B - Botanisches Museum, Berlin-Dahlem, Germany. 

BM - British Museum of Natural History, London, Great Britian. 

BRI - Botanic Museum and Herbarium, Brisbane, Queensland, Australia. 

EA - The East African Herbarium, Nairobi, Kenya. 

F - Chicago Natural History Museum, Chicago, Illinois. 

G - Conservatoire et Jardin Botaniques , Geneve, Switzerland. 

GH - The Gray Herbarium of Harvard University, Cambridge, Massa- 

chus etts . 
HBG - Stattsinstitut fur allgemeine Botanik, Hamburg, Germany. 
K - The Herbarium, Royal Botanic Gardens, Kew, Great Britian. 
L - Rijksherbarium, Leiden, Netherlands. 
MEXU - Herbario Nacional del Instituto de Biologia, Mexico, D.F. 

Mexico 
NY - New York Botanical Garden, New York, New York. 
R - Divisao de Botanica do Museu Nacional, Rio de Janeiro, Brazil. 
RB - Jardim Botanico, Rio de Janeiro, Brazil. 

UC - Herbarium of the University of California, Berkeley, California. 
US - National Museum, Smithsonian Institution, Washington, D.C. 
W - Naturhistorisches Museum, Wien, Austria. 

In addition to the species from the above-mentioned herbaria, 
specimens of B. grand iflora , B. multicaulis and B. ovalifolia from 
the following herbaria were examined. 

DUKE - Duke University, Durham, North Carolina. 

FLAS - Herbarium of the University of Florida, Gainesville, Florida. 

FSU - Florida State University Herbarium, Tallahassee, Florida. 

GA - Herbarium of the University of Georgia, Athen, Georgia. 

MICH - University Herbarium, University of Michigan, Ann Arbor, 

Michigan. 
NCU - Herbarium of University of North Carolina, Chapel Hill, 

North Carolina. 
NSC - Department of Botany, North Carolina State College, Raleigh, 

North Carolina. 
PH - Academy of Natural Sciences, Philadelphia, Pennsylvania. 
SMU - Herbarium of Southern Methodist University, Dallas, Texas. 



History 

The generic name Bonamia was established by DuPetit-Thouars 
(1804) in honor of Francois Bonami (1710-1786), a French physician 
and botanist who wrote the Flora of the Environ of Nantes in 1782. 



126 PHYTOLOQIA Vol. 17, no. 3 

It was basod on a woody virif of Madagascar, ]ator described as B. 
altornifolia by Jaumn Saint-Hilaire (1805) and as B. madagascar - 
iensis by Poirnt flSlO). 

The generic name IJreweria was proposed by Robert Brown ("1310) 
in honor of Samuel Brewer ("1670-1743) , an English amateur botanist. 
The generic description was based on three Australian species, 
Breweria linearis , Br . media and Br. pannosa (i.e. Bonamia linearis , 
B. media and B. pannosa of the present treatment). Before the com- 
parative study of the Convolvulaceae by Hallier C1893), Breweria 
was most commonly treated as a genus distinct from Bonamia . 

The generic name Trichantha was described by Karsten and Triana 
(1856), based on a woody vine of Columbia, described by them as 
Trichantha ferruginea (=Bonamia trichantha of the present treatment). 
However, this generic name is invalid, since it is preoccupied by 
Trichantha Hooker (1844) of the Gesneriaceae. 

The name Peris permum was established by Otto Degener (1932), 
based on a woody vine, Peris permum albiflorum ( =Bonamia menziesii 
of the present treatment) , which is endemic to the Hawaiian Islands. 

A fifth generic name, Breweriopsis , was proposed by G. Roberty 
in his new and strange system of classification of the Convolvula- 
ceae (1952). It was based on Breweriopsis elegans (= Bonamia elegans 
of the present treatment), an endemic of lower Burma. 

Although these five generic names were described from different 
plants from various parts of the world (Madagascar, Australia, Co- 
lumbia, Hawaii and tropical Asia), they all are characterized by 
essentially similar floral features. Other generic names of close 
nomenclatural association, especially to the names Breweria and 
Bonamia , are Stylisma Rafinesque (1818), Seddera Hochst, (1844), 
Calycobolus Willd. ex Roem. and Schult. (1819), Prevostea Choisy 
(1825), Dufourea H.B.K. (1818), Reinwardtia Spreng. (1825), De- 
thardigia Nees et Mart. (1823), Codonanthus G. Don (1856), and 
Metaporana N.E. Brown (1914). 

The treatment of these genera in the past has varied widely. 
Choisy (1845) treated Bonamia , Breweria , Stylisma , Seddera and 
Prevostea ( =Calycobolus ) as distinct genera. Gray, in his earlier 
manual (1856), treated Stylisma as a distinct genus; but later 
(1862) he questioned the validity of Breweria and Stylisma , and 
suggested their reduction to Bonamia . Bentham and Hooker (1876) 
did not accept Gray's suggestion and treated Bonamia as a monotypic 
genus of Madagascar and Breweria in a very broad sense by in- 
cluding species previously assigned to Seddera , Stylisma and Caly- 
cobolus , in addition to species described under Breweria and 
Trichantha. Peter (1897) slightly modified Bentham and Hooker's 
classification by treating Bonamia as containing two species 
(B. alterni folia and B. menziesii) , members of Calcycobolus and 
one species of Bonamia (B. ferruginea) under the generic name 
Prevostea, and the rest in three subgenera (namely Seddera , 



1968 Hyint & Ward, Revision of Bonaniia 127 

Stylisma and Eubreweria ) under the generic name Breweria . 

Hallier (1893) was the first to call attention to the weak- 
ness of the differences observed between Bonamia and Breweria sen. 
str. He combined the two groups into a single genus, and the ol- 
der name Bonamia was substituted for the later Breweria . He re- 
tained Stylisma , Seddera and Prevostea as distinct genera from 
Bonamia (although his choice of the name Prevostea rather than 
Calycobolus was incorrect). The generic delimitation thus adopted 
and revised by Hallier was accepted by House (1907), but he treat- 
ed three species of Bonamia under Calycobolus , since he missed the 
fact that members of Calycobolus are characterized by accrescent 
sepals rather than by unequal sepals.. Amongst the authors of some 
local floras, Baker and Rendle (1906), Hutchinson and Dalziel 
(1931), Small (1933), van Ooststroom (1932; 1954), O'Donell (1959) 
and Vercourt (1963) followed Hallier, whereas Clarke (1883), 
Bailey (1901), Baker and Wright (1904) and Fernald (1950) appar- 
ently followed Bentham and Hooker. In the more recent studies 
Meeuse (1957), working on south African species, referred a spec- 
ies of Seddera to Bonamia , and Wilson (1960) and Shinners (1962) , 
independently working on the North American species, referred all 
species of Stylisma to Bonamia. 

Roberty's treatment of Bonamia and its related genera in his 
new system of classification (1952) is so different from all other 
authors mentioned above and so artificial in selection of the dis- 
tinguishing characters that it is nof at all acceptable and de- 
serves no special attention except a short comment. His proposal 
of the new genus Breweriopsis and treatment of Bonamia and Breweria 
as distinct genera are based on insufficient knowledge of the 
plants, as is evident from the fact that he included B. spectabilis 
in Breweriopsis (under B. elegans ) and B. minor in Bonamia (under 
B. cymosa ) , whereas these two are definitely conspecific (the dif- 
ference being only in the pubescence of stems, which is variable). 
Further, he included three species of Bonamia in Stylisma humi strata 
in addition to all known species of that genus; he also associated 
Bonamia ferruginea with Dipteropeltis ferruginea , an entirely dif- 
ferent plant of tropical Africa. Several other serious errors 
have been pointed out by Verdcourt (1957; 1963). 



Morphology 

The morphological characteristics of species of Bonamia are 
poorly known because of the infrequent or rare occurrence and spor- 
adic or limited distribution of many species. Most previous stu- 
dies, except Hallier's comparative study of the family and synopsis 
of Bonamia , have been directed mainly to the descriptions of es- 
sential features for delimiting different species. 

HABIT ; Plants of Bonamia are perennial, woody, suffrutes- 
cent or rarely herbaceous vines, occasionally small shrubs or 



128 P H Y T L G I A Vol. 17, no. 3 

suljshrubs, growing from deep-spated roots. The roots arf- mostly 
woody and never tubfjrous as they are in some species of I porno ea , 
the largest genus of the Convolvulaceae. The tap roots are thick, 
at least near the bases of the shoots, and gradually taper down- 
ward. Adventitious roots at the nodes are not common, even in thr- 
prostrate or trailing species. 

The stem is generally weak and slender or occasionally woody, 
mostly prostrate, twining or scandent, infrequently procumbent, 
suberect or erect. Members of section Trichantha are consistently 
woody and high-climbing vines or small erect shrubs, as in B. 
corumbaensis . The habit of members belonging to section Bonamia 
is quite variable from species to species, from suberect or pro- 
cumbent as in B. sericea and B. ovalifolia , prostrate, twining 
or scandent as in most other species, to very high climbing as 
in B. brevipedicellata , B. maripoides , B. ferruginea , and B. semi - 
digyna . B. brevipedicellata has been recorded as "50 ft. high, 1 
inch in diameter." Members belonging to the section Breweria are 
generally smaller, somewhat herbaceous, suffrutescent or becoming 
woody. They are mostly prostrate, procumbent or erect, rarely 
twining. The stem is usually slender, as in B. linearis , B. media 
and B. brevifolia, or thick and erect as in B. rosea and B. velu - 
tina. In general, section Trichantha and section Breweria show 
the extreme types of habit, whereas section Bonamia is somewhat 
intermediate. 

STEM ; The stems are mostly terete or slightly angular, smooth, 
minutely striated, punctate or provided with lenticels, glabrous, 
sparsely pubescent, glabrescent, moderately to densely sericeous, 
villous, tomentose, velutinous or ferrugineous. Internodes, high- 
ly variable in length from species to species, are one or two cen- 
timeters long, as in most species of section Breweria and some 
species of section Bonamia , to several centimeters long, as in 
most species of section Trichantha and a few species of section 
Bonamia . The colors are light green, greyish green, silvery grey 
or brownish grey depending on the absence or presence of a dense 
coating of hairs. Underground stems have not been seen, although 
they might be present in several species. Branching is alternate, 
and frequent or occasional. Extent of branching is not a constant 
feature and is variable even in a single species. In some species 
there is a little or no secondary branching. 

LEAVES : In general, the leaves show a homoblastic series, 
with all leaves similar in shape, although with the upper smaller 
than the lower. B. elegans shows a heteroblastic series, with 
the juvenile leaves ovate, elliptic or ovate-elliptic, and the 
upper leaves (on flowering branches) oblong. 

Leaf shape in members of the genus Bonamia varies from ellip- 
tic, ovate, cordate, or orbicular (with length-width ratios of one 
or very close to one) to oblong, lanceolate or linear (with length- 
width ratios of two or higher). Leaf size also varies from one 



1968 Ityint & Ward, Revision of Bonami a 129 

centimeter of slightly longer as in B. brevifolia , B. media and 
B. rosea , to several centimeters long as in B. agrostopolis , B. 
ferruginea, B. mattogrossensis , B. kuhlmannii , B. subsessilis 
and B. trichantha . In general, the leaves are smaller in members 
of section Breweria and larger in those belonging to section Tri- 
chantha, while most species of section Bonamia possess leaves of 
somewhat intermediate sizes. 

Leaves are sessile, subsessile or shortly petiolate in the 
members of section Breweria , whereas they are distinctly petio- 
late or long-petiolate in those of section Trichantha . Leaves in 
members of section Bonamia may be sessile, subsessile, shortly 
petiolate or long-petiolate. Leaves are thin, herbaceous or slight- 
ly subcoriaceous as in section Breweria , thin or thick, soft or 
subcoriaceous , as in most species of section Bonamia, or thick, 
subcoriaceous, coriaceous or leathery as in section Trichantha . 
Leaf base and apex vary inconsistently from acute and attenuate 
to obtuse, rounded, truncate and cordate or emarginate; leaves 
are frequently mucronate or mucronulate. Such wide variations 
of leaf base and apex are shown in all three sections. 

Leaves are entire in most species , slightly undulate in B. 
alternifolia and slightly crenate or somewhat wavy in B. burche - 
llii . They are mostly green, dark green or greyish green on the 
upper surface, pale or light green on the lower surface. Leaf 
surfaces are glabrous, sparsely pubescent, puberulous , sericeous, 
velutinous, tomentose, strigose, or f errugineous , frequently more 
densely so on the lower surface or on the veins. Veins are thin 
and obscure as in some species of section Breweria or very promi- 
nent as in most species of section Trichantha , in which even the 
intercostal veins are prominent. In the species of section Bonamia , 
veins are not distinct as in B. d ens i flora , B. multicaulis , and 
B. sericea , or are very prominent as in B. alternifolia and B. 
ferruginea . Lateral veins are few (2-5) , as in some Australian 
species, or several (6-11), as in most South American species. 

INDUMENTUM ; Hairs, except glandular ones, in all species of 
Bonamia are two-celled, and are usually appressed. The stalk-cell 
is extremely small, and the terminal cell bears two elongated arms. 
Both stalk and two-armed cells are mostly thick-walled or rarely 
thin-walled. The two arms of hairs on the vegetative parts and on 
the sepals are equal or slightly unequal and usually point in oppo- 
site directions. However, hairs on the margin of the sepals, when 
present, possess erect arms, thus pointing approximately in a 
single direction (i.e. away from surface); hairs on interplicae 
of the corolla and upper part of the tube possess unequal arms, 
with one extremely long arm pointing toward the apex of the pe- 
tals and another very short or almost indistinguishable arm point- 
ing toward the base of the corolla; hairs on the filaments of the 
stamens are curly and soft and occasionally glandular; hairs on 
the ovary, mostly at the apex, have two straight arms pointing to- 
ward the mouth of the corolla. 



130 P H Y T L G I A Vol. 17, no. 3 

The typn of hairs is generally uniform in the whole genus, 
although their density, length or arms and occurrence on different 
parts vary from species to species and are taxonomically impor- 
tant. They are distinctly appressed when their occurrence on 
certain parts is sparse or scattered; but when the hairs are 
dense, they are less appressed and sometimes crisped. 

INFLORHSCCNCF, : The flowers are axillary and mostly soli- 
tary or in simple dichasial cymes in the members of section Bre - 
weria. In the members of section Trichantha , the flowers are 
mostly in axillary compound or simple cj^es , or in terminal pan- 
icles, rarely solitary or in simple cymes. The flowers in the 
members of section Bo nam i a are variable from axillary and soli- 
tary to dense clusters in an axillary or terminal position. Sol- 
itary flowers seem to be a result of reduction of compound in- 
florescences, since the less advanced species possess inflorescences 
of large number of flowers, whereas the more advanced species 
generally possess solitary flowers or simple cymes. This is evi- 
dent in some species in which the aborted floral buds occur in the 
axils of bracts or bracteoles. 

FLOWER : The flowers are sessile, shortly pedunculate or 
shortly pedicellate, distinctly pedunculate and pedicellate or 
long-peduculate. Such a wide variation is most evident in the 
section Bonamia. In B. ovalifolia , B. grand i flora , B. multicaulis , 
and B^ sericea , the flowers are sessile, subsessile, shortly pe- 
dunculate or shortly pedicellate, whereas in B. semidigyna and B. 
cordata the peducles are comparatively very long. In B. peru- 
viana and B. kuhlmannii the pedicels and secondary peduncles are 
nearly as long as the primary peduncles, whereas in B. maripoides 
and B. sulphurea the pedicels are fairly long, sometimes longer 
than the short peduncles. This feature is rather uniform in 
the members of section Breweria and section Trichantha . 

Bracts and bracteoles are usually small, reduced and scale- 
like, with a few exceptions. In most species they are linear, 
linear-lanceolate or subulate and mostly shorter than pedicels. 
However, bracts in B. pannosa are long-linear, distinctly longer 
than pedicels; bracts in B. brevifolia are as long as the pedi- 
cels; bracts in B. cordata and B. mossambicensis are foliaceous, 
mostly ovate-lanceolate or elliptic-lanceolate in shape and as 
long as or longer than the sepals. Bracteoles also show the same 
range of shapes, although their size is smaller. Both bracts and 
bracteoles in most species are alternate. The alternate position 
is not conspicuous in B. mossambicensis and B. sphaerocephala , 
whose inflorescences are multiflorus and dense because of ab- 
sence of pedicels. The bracts and bracteoles are persistent in 
most species but deciduous in some species of section Trichantha . 

CALYX : All species have calyces composed of five quincun- 
cially imbricated sepals, which are free or slightly fused at 
their extreme base. In all species of the genus the sepals are 



1968 Myint & Ward, Revision of Bonamia 131 

not accrescent. This is the most important characteristic dis- 
tinguishing the genus Bonamia from the genus Calycobolus . The 
sepals in Bonamia are coriaceous, subcoriaceous or rarely soft and 
somewhat herbaceous, but never membranous. The shape and size are 
highly variable from species to species. In general, they are 
lanceolate, ovate, ovate-lanceolate, oblong-ovate, elliptic, or- 
bicular or rarely ovate-subcordate, obovate or oblique-ovate. They 
are acute, obtuse, acuminate, obtuse-mucronate, rounded, truncate 
or slightly emarginate at the apex. They are equal or subequal in 
most species and distinctly unequal in some species, such as B. 
cordata , B. f erruginea , B. mossambicensis , B. pannosa , B. peru - 
viana and~B. kuhlmannii . In all these species, the outer two 
sepals are large, ovate, ovate-orbicular, ovate-cordate, or rarely 
ovate-lanceolate, densely tomentose, f errugineous , or pilose 
outside, densely sericeous or velutinous inside except at the 
center, or glabrous. The in-out sepal (third sepal) is smaller 
like the inner two, or large and oblique-ovate in shape and similar 
to the outer sepals in its pubescence (except at the inner margin). 
In the members of section Breweria , sepals are mostly ovate-lanceo- 
late, small and equal or subequal except in B. pannosa . They are 
sericeous, tomentose, pilose or velutinous outside in all members 
of this section. In the members of section Trichantha , sepals 
are ovate, ovate-orbicular, oblong-ovate or orbicular, equal or 
slightly unequal, and tomentose, sericeous, sparsely pubescent, 
puberulous or rarely glabrous outside. The sepals are glabrous 
inside except in B. mattogross ens is . In the members of section 
Bonamia the sepals are highly variable from species to species 
in their shape, from lanceolate to ovate or orbicular, from equal 
to unequal, in their size from small to large, in their apices 
from acute and acuminate to obtuse and rounded, and in their sur- 
face from glabrous or sparsely puberulous to densely tomentose, 
villous or f errugineous. Sepals are generally thin, herbaceous or 
rarely subcoriaceous in section Breweria , thick and coriaceous in 
Trichantha and variable in section Bonamia . 

COROLLA : The corolla is sympetalous, funnel-shaped, shortly 
tubular campanulate, narrow-campanulate or campanulate-funnelform, 
and plicate in the bud stage. It is shallowly lobed as in B. al- 
ternifolia , somewhat lobulate, entire or subentire as in most 
species of the genus. During the bud-stage the lateral edges of 
each petal are hidden by being folded inwardly along the line of 
fusion between the petals. The infolded areas, termed plicae, are 
roughly triangular with their apices extending toward the tubular 
portion of the corolla, with the lateral angles of adjacent plicae 
nearly meeting at the apex of each petal. Between the plicae, the 
central portions of the petals or mesopetaline bands, termed inter- 
plicae, form the exposed surface of the bud. Each interplica is 
narrowly triangular tapering toward the apex of the petal, with 
the base merging with adjacent interplicae to form the tubular por- 
tion of the corolla. The size of the corolla varies from small 
to large. In the section Breweria , the corolla is usually small, 
8-15 mm. long, rarely longer. In the section Trichantha , the 
corolla is somewhat larger, mostly 18-25 mm. long. In the section 



132 PHYTOLOGIA Vol. 17, no. 3 

Bonamia, the corolla is large or smal]. 

The pigmentation of the corolla varies from species to species, 
but it is constant within a single species — blue, deep blue, pur- 
plish blue, or purple in B. grand i flora , B. elliptica , B. ovali - 
folia, B. multicaulis , B. elegans , B. spectabilis , B. mossambicen - 
sis , B. mattogrossensis , B. pannosa , and B. media ; red, rose, or 
pink in B. abscissa and B. rosea ; yellow or yellowish in B. balan - 
sae and perhaps in B. corumbaensis and B. menziesii . In the re- 
maining species, the corolla is white, or, in a few of them, un- 
known. The color of the corolla is variable from blue to white 
in B. media and B. pannosa , and from red to pink or white in B. 
rosea . 

ANDROECIUM ; The stamens are five, alternate to the petals. 
They are mostly inserted, or rarely exserted as in B. altemifolia . 
The filaments are epipetalous , being adnate to the corolla at the 
basal portion. They are usually slightly longer than the styles, 
but may be appreciably longer or shorter. They are equal, subequal 
or unequal in length, and generally filiform above and flattened 
or widened near the base. The filaments are glabrous, sparsely 
or densely villous, at least the lower part. The anthers are 
bilocular, introrse or partially introrse, dehiscing by longitudinal 
slits. Their attachment to the filaments is mostly dorsal or basi- 
dorsal, and sagittate, cordate or subcordate at the base. They 
are oblong, oblong-ovate or lanceolate and usually 1.5-5 ram. long. 

GYNOECIUM ; The gynoecium is composed of two carpels, fused 
except for the stigmas and a portion of the styles. The degree 
of fusion of the stylar branches, termed stylodia, is variable 
from species to species; but in general the stylar branches are 
free at least for the upper one-fourth and in many species they are 
free to the middle, nearly to the base or readily separable to the 
base. The stylar branches are filiform, glabrous or rarely with 
scattered hairs, and equal or unequal in length. Each stylar branch 
is provided with a single vascular strand, which branches near the 
stigmas in the members of section X nichantha ; in the members of 
two other sections it is unbranched. The stigmas are globose, sub- 
globose, globose-capitate, capitate, conical, reniform, or bilobed , 
rarely peltate. They are usually wider than the tips of stylar 
branches, and rarely small and not distinctly distinguishable. 
The surfaces of stigmas are smooth, warted or occasionally rugose 
or rugulose. In the section Breweria stigmas are usually large, 
globose, subglobose or globose-capitate, rarely peltate and mostly 
smooth. In the section Trichantha stigmas are large, reniform, 
bilobed or capitate and smooth. In the section Bonamia stigmas are 
variable from small to large, globose to capitate or peltate, and 
from smooth to rugose or rugulose. 

The ovary contains two chambers, each of which encloses two 
erect ovules in axile placentation. It is ovoid, ovoid-conical, 
oblong or conical. It is sparsely long-pilose, densely long-pilose, 
tomentose or glabrous, often pilose only at the apex. The ovary 



1968 Mylnt & Ward, Revision of Bonamla 133 

at its base is usually surrounded by a thin or thick annular disc, 
very prominent in several species and frequently enlarged in the 
fruit. 

FRUIT ; The fruits in all species are valvular capsules with 
persistent sepals. They are most frequently two- to four-seeded, 
but occasionally one-seeded. Although the number of seeds per 
capsule is variable and is not a good characteristic for distin- 
guishing Bonamia from Stylisma (as was done by Hallier) , the 
mean number differs between the two genera (higher in Bonamia 
and lower in Stylisma ) and in several species two- or one- 
seeded capsules are rare. The capsules are thin-walled or thick- 
walled and four- or eight-valvular, rarely two-valvular. In 
some species the individual valves may break again into smaller 
pieces , and thus the capsules may superficially appear to be 
multivalvular. In the members of section Trichantha , the capsule 
walls are thick and hard , frequently breaking into two pieces , 
although they are four-valvular in reality. Occasionally the 
capsules may remain indehiscent for a long time as in B. menziesii . 
The septum in the capsule is thin in most species of section 
Breweria and section Bonamia , whereas it is thick and hard in 
the members of section Trichantha . The capsules are small in 
section Breweria and are large in section Trichantha , whereas 
they are variable in size in section Bonamia . 

SEED : Seeds are somewhat similar in shape from species to 
species, but are variable in size, color, surface and indumentum. 
In section Bonamia seeds are small or large, varying from 3-6 (7) 
mm. in length, brown, dark brown, or black in color and smooth or 
punctate and glabrous on the surface. In section Breweria , seeds 
are small, 2-4 mm. long, brown or dark brown, smooth or punctate 
and glabrous. In section Trichantha , seeds are small or large, 
4-7 mm. long, brown or dark brown, and long-haired along the edge 
and villous or long-haired on the ventral and dorsal surface. 
The seedcoat is hard in all species and is frequently covered with 
a thin perisperm. 

The embryo is embedded in cartilaginous endosperm, which on 
wetting swells into a gelatinous mass. The embryonic axis is short 
with an indistinguishable hypocotyl or elongate with a short hypo- 
cotyl. The plumule is located between the two cotyledonary petioles 
or on the side of the stalk formed by the fusion of the two cotyle- 
donary petioles. The cotyledons are petiolate, foliaceous , thin 
and herbaceous. They are generally ovate, obovate, ovate-cordate, 
orbicular, obscurely bilobed or rarely linear-bifid, rounded, trun- 
cate or emarginate at the apex, cordate or truncate at the base, 
and mostly symmetrical or rarely somewhat oblique. The two coty- 
ledons are closely appressed to each other and the two fold 
together repeatedly (thus appearing to be raultiplicate and corrugate) 
and also fold against the radicle. In some species the closely 
appressed cotyledons are flat and folded once or twice and then 
against the radicle. 



13li P H Y T L G I A Vol. 17, no. 3 

Systematic Treatment 

Bonamia DuPetit-Thouars , Hist. Veg. Isl. France Reunion, Madagas- 
car 1:33, pi. 8. 1804, nom. cons. 
Breworia R. Br. Prodr. Fl. Nov. Holl. 487. 1810. Typo: 

Bonamia linearis fR. Br.) Hall. f. (as Breweria linearis 

R. Br. 1810 — BM; W.'). 
Trichantha Karst. et Triana, Linnaca 28:437. 1856. Type: 

Bonamia trichantha Hall, f . Cas Trichantha ferruginea 

K. et T. 1856--G; BM.' W.' ) ; not Trichantha Hooker, Icon. 

PI. tt. 666,667. 1844. 
Perispermum 0. Degener, Flora Hawaiiensis, Fam. 307. 1932. 

Type: Bonamia menziesii Gray (as Perispermum albiflorum 

Degener, 1932— MO.'). 
Brewer iops is G. Roberty , Candollea 14:31. 1952. Type: 

Bonamia elegans (Wall. ) Hall. f. (as Breweriopsis elegans 

(Wall.) Roberty, 1952— BM.' G.'). 

Perennial, herbaceous, suffrutescent or woody, twining, prostrate 
or trailing, occasionally procumbent or suberect, shubby vines or 
erect subshrubs; shoots a few to several, simple or branched, grow- 
ing all year around or arising annually from old shoots, bases of 
previous shoots, crowns, horzontally spreading subterranean stems, 
or from roots. Roots deep-penetrating, often becoming thick in 
some, frequently with pulpy bark, never bulbous nor fleshy. Stems 
mostly thin, elongate, occasionally becoming as thick as 2.5 cm. 
at the base, as long as a few decimeters to several meters in 
height, smooth or lenticellate, glabrous to densely pubescent, 
villous, sericeous or ferrugineous. Leaves petiolate, subsessile 
or sessile, estipulate, herbaceous or subcoriaceous , occasionally 
leathery; blades simple, entire, occasionally undulate or slightly 
wavy, ovate, ovate-cordate, elliptic, ovate-elliptic, oblong-ovate, 
lanceolate, oblong, linear or linear-lanceolate, often large, acute, 
obtuse, acuminate, acute-mucronate , obtuse-mucronate or slightly 
emarginate at the apex, acute, attenuate, cordate, rounded or trun- 
cate at the base; veins prominent to inconspicuous except the mi- 
dribs, mostly impressed above, often with distinct intercostal 
veins; hairs appressed, two-armed , straight or crisped, very fine 
to distinctly long, scattered or dense, silvery grey, greyish 
white, pale brown or grey, often becoming rusty brown when dry. 
Inflorescences axillary or terminal, pedunculate or subsessile, 
simple or compound dichasial cymes of few to several flowers, 
often solitary or terminal panicles composed of several dichasial 
cymes; peduncles short or elongate, usually shorter than leaves, 
or absent; pedicels usually short, occasionally elongate (becoming 
as long as 2 cm.); bracts small and linear or distinctly folia- 
ceous, mostly two for each individual flower, opposite or slightly 
alternate, sometimes crowded in congested clusters. Sepals five, 
quincuncially imbricate, free or rarely united at the extreme base. 



1968 L^'int & Vifard, Revision of Bonamia 135 

mostly ovate, ovate-lanceolate, broadly lanceolate, orbicular, or 
oblong-orbicular, equal or unequal, acute, acuminate, obtuse, 
rounded or slightly emarginate at the apex, sericeous, tomentose, 
pilose, velutinous , ferrugineous or glabrous on the inner surface, 
persistent in capsules. Corolla white, blue, bluish purple, pink 
or red, yellow, yellowish white or greenish white, funnel-shaped, 
campanulate or shortly tubular campanulate, plicate in bud, sym- 
petalous with entire, subentire, lobed or lobulate limb, outside 
sparsely or densely pilose on interplicae (midpetaline bands), 
glabrous on plicae (infolded areas); individual hairs on inter- 
plicae with two unequal arms (long arms directed toward apices 
of petals). Stamens five, alternate with petals, inserted or rare- 
ly exserted, all fertile; filaments epipetalous (being adnate to 
the lower, narrow part of the corolla), straight, filiform or 
somewhat dilated below, dorsiventrally flattened, unequal, sub- 
equal or equal in length, glabrous or thinly to densely villous 
or gland ular-villous (with crisped or curly hairs), frequently 
villous only on the basal dilated portions; anthers two-celled, 
oblong or oblong-lanceolate, dorsifixed or apparently basifixed, 
frequently sagittate or cordate at the base, introrse or partially 
extrorse by vertically dehiscing slits; pollen colpate and punc- 
titegillate, not spiniferous. Ovary superior, bicarpellate, bilo- 
cular, long-pilose or hirsute with two-armed hairs (both arms of 
each hair directing toward the mouth of the corolla) or glabrous, 
surrounded by annular disc at the base; ovules two in each loculus, 
erect, anatropous , in axile placentation, appearing to be basal; 
styles terminal, two, almost free to partially united, included 
in the corolla to partially exserted; stylar branches (or stylodia) 
equal to unequal, filiform, mostly glabrous, occasionally with 
scattered hairs; stigmas large or small, globose, subglobose, 
capitate, reniform, bilobed conical or rarely peltate, smooth or 
rugose, occasionally lobulate. Fruits 1- to 4-seeded, 4- to 8- 
valvular, rarely 2-valvular capsules with thin and chartaceous 
or thick and ligneous walls, ovoid, globose or conical-ovoid, 
apiculate at the apex, glabrous or with scattered hairs, two-celled 
with thin or thick septum, with persistent sepals, dehiscing by 
valves, occasionally dehiscing by basal circumcision, rarely re- 
maining indehiscent for a long time after ripeness. Seeds brown, 
dark brown or black, smooth or punctate, glabrous or lanate, oval 
and plano-convex or roughly three-angled , with hard or rarely 
soft seedcoat, covered with thin transparent perisperm; endosperm 
scanty or copious, cartilagenous , swelling on wetting. Cotyledons 
thin, foliaceous, ovate, obovate, ovate-cordate or orbicular, 
rarely linear-bifid, mostly symmetrical, rarely slightly oblique, 
corrugate-plicate and folded against radicle or simply flat 
or slightly folded along central line and folded against radicle; 
cotyledonary petioles free or fused. Flowering from summer to 
winter. 

Type: B. madagascariensis Poiret, in Lamarck, Encycl. Meth. 
Bot. suppl. 1:677. 1810, nom. illeg. (B. altemifolia J. St. Hi- 
laire, Expos. Fam. 2:349. 1805.) 



136 PHYTOLOGIA Vol. 17, no. 3 

Dry sandy soils of various types, raroly moist or wet ground, 
frequently coastal plains and dunes, occasionally inland high 
ground, open forests, grassy plains, scrubby lands, edges of dense 
forests, frequently along streams and riverbanks ; of tropics, sub- 
tropics and warm temperate regions of both hemispheres, with a 
concentration of more species in South America, Australia and 
Madagascar, and fewer species in Asia, North America emd mainland 
Africa. 

The genus is heterogeneous and is roughly separable into 
three sections. 



Key to Sections of Bonamia 

1. Seeds glabrous; fruits with thin walls, dehiscing into 
four or eight valves, rarely indehiscent, with thin or 
membranous septa; flowers solitary, in simple or com- 
pound cymes, umbellate or capitate heads (axillary or 
terminal) . 2 

1. Seeds fulvous -villous on the ventral and dorsal sides, 
long-haired along the edge; fruits with ligneous and 
thick walls, dehiscing into two or four valves, with 
ligneous or thick septa; flowers mostly in panicles or 
pseudopanicles , rarely solitary or simple cymes. 

Sect. Trichantha 

2. Corolla 1.8 cm. or longer, if shorter, flowers in dense 
capitate clusters; peduncles or pedicels or the two to- 
gether consistently as long as 1 cm. or longer, rarely 
shorter; leaves 3 cm. or longer; flowers mostly in 
cymes, dense clusters, or occasionally solitary. 

Sect. Bonamia 

2. Corolla shorter than 1.8 cm.; peduncles or pedicels or 
the two together mostly 5 ram. or shorter, rarely long- 
er; leaves shorter than 3 cm. , narrower than 2 cm. , if 
longer or wider, corolla shorter theui 1.8 cm.; flowers 
mostly solitary, occasionally in simple cymes, rarely 
5 to 7-flowered cymes. Sect. Breweria 



General Key to Species of Bonamia 

1. Outer sepals larger than inner sepals, as wide as or wider 
than twice the width of inner sepals and more densely 
tomentose. 2 

1. Outer and inner sepals equal, subequal or slightly un- 
equal (outer sepals 1.5 X inner sepals or narrower). 7 



1968 Myint & Ward, Revision of Bonamia 137 

2, Leaves shorter than 4 cm. ; flowers solitary or rarely 
in simple cymes, sessile or subsessile, rarely short- 
pedunculate or short-pedicellate. 31. B. pannosa 

2. Leaves longer than 4.5 cm.; flowers mostly in compound 
cymes , usually numerous or in capitate cymes , or long- 
pedunculate and/or long-pedicellate. 3 

3. Pedicels longer than 1 cm. or peduncles 3 cm. or longer; 
inflorescences loose cymes or few-flowered cjmies ; leaves 
ovate or ovate-cordate. 4 

3. Pedicels very short, rarely as long as 7 mm. ; inflores- 
cences dense capitate, sessile or shortly pedunculate; 
leaves ovate-lanceolate, ovate-elliptic or oblong-ovate. 6 

4. Bracts foliaceous , ovate or ovate-lanceolate, 5 mm. or 
longer; pedicels short; peduncles 3 cm. or longer. 

6. B. cordata 

4. Bracts minute, scale-like, linear or subulate, 2-4 mm. 
long; pedicels longer than 1 cm.; peduncles 2.5 cm. or 
shorter. 5 

5. Leaves 5-12 cm. by 3-8 cm. , cordate or subcordate at the 
base; outer sepals 2 cm. by 1.7 cm.; styles free almost 

to the ovary. 24. B. kuhlmannii 

5. Leaves 5.5-7 cm. by 3-4 cm., obtuse or truncate at the 
base; outer sepals 8-12 mm. by 7-10 mm. ; styles fused 

at least lower one-third. 25. B. peruviana 

6. Sepals ferrugineous or tomentose (with short hairs); 
outer sepals thick, ovate or ovate-subcordate, obtuse 
at the apex; leaves obtuse or acute at the apex; bracts 
inconspicuous. 21. B. ferruginea 

6. Sepals long-sericeous or hirsute (with long hairs); 
outer sepals thin and somewhat foliaceous (with dis- 
tinct venation), ovate-lanceolate, acute or acuminate at 
the apex; leaves acute or acuminate at the apex; bracts 

1 cm. long. 5. B. mossambicensis 

7. Sepals consistently (both outer and inner) acute or 
acuminate at the apex; if obtuse then 10 mm. or longer. 8 

7. Sepals (at least the inner or the outer) obtuse, rounded, 
obtuse-mucronate or emarginate. 29 

8. Flowers in dense clusters, capitate or dense umbellate 
cymes, mostly sessile, subsessile or rarely shortly 
pedicellate. 9 



138 P H Y T L G I A Vol. 17, no. 3 

8. Flowers solitary or in axillary loosp cymos of 3-7; if 
morf, podicols or poduncles 2 cm. or longer. 11 

9. Leaves elliptic, oblong or oblong-elliptic, 4 cm. or 
shorter, 2 cm. or narrower, long-mucronate at the apex, 
dense lanate on the surface; corolla 1 cm. or shorter. 

23. B. sphaerocephala 

9. Leaves elliptic-lanceolate or ovate-lanceolate, longer 
than 4 cm. or wider than 2 cm., attenuate at the apex, 
sericeous, hirsute or glabrate on the surface; corolla 
longer than 1.5 cm. 10 

10. Sepals hirsute with long hairs, unequal; corolla blue; 
bracts pilose or hirsute with long hairs. 

5. B. mossambicensis 

10. Sepals pubescent with short hairs, equal or subequal; 
corolla white; bracts puberulous , finely sericeous or 
nearly glabrous. 18. B. holtii 

11. Pedicels or poduncles or the two together as long as 

1 cm. or longer consistently. 12 

11. Pedicels or peduncles or the two together shorter than 

1 cm. (at least a few of them) . 23 

12. Sepals ovate or ovate-lanceolate, usually shorter than 
1. 5 X width, tomentose, villous, densely sericeous or 

f errugineous , mostly 12 mm. or shorter, rarely longer. 13 

12. Sepals lanceolate or oblong-lanceolate, usually as long 
as 1. 5 X width or longer, glabrous or puberulous, mostly 

12 mm. or longer, rarely slightly shorter. 19 

13. Sepals 7-12 mm. wide, densely villous or f errugineous. 14 

13. Sepals 4-6 mm. wide, softly pubescent, tomentose, 
sericeous or nearly glabrous. 18 

14. Leaves linear-lanceolate, 10 mm. or narrower, with 
length-width ratio of 2.5 or higher; corolla blue. 

15. B. multicaulis 

14. Leaves ovate, cordate, elliptic, oblong-elliptic or 
rotund, wider than 10 mm. and/or with length-width 

ratio of 2 or lower; if higher, corolla white. 15 

15. Leaves cordate at the base, 4 cm. or wider; peduncles 3 

cm. or longer. 16 

15. Leaves rounded, obtuse or rarely subcordate at the base, 

3 cm. or narrower; peduncles shorter than 3 cm. 17 



1968 lilyint & Ward, Revision of Bonamia 139 

16. Corolla white or yellowish white, with slightly lobu- 
late limb; flowers mostly in simple or compound cymes. 

7. B. semidigyna 

16. Corolla red or pink, with entire or subentire limb; 
flowers mostly solitary, rarely in simple cymes. 

41. B. abscissa 

17. Leaves ovate or orbicular, 2.5 cm. or shorter; corolla 
blue, 3 cm. or longer; stem 1 m. or shorter, procumbent 

or suberect. 14. B. ovalifolia 

17. Leaves oblong-elliptic, elliptic, oblong-lanceolate or 
rarely rotund, mostly longer than 2.5 cm.; corolla white 
or yellowish, 2.5 cm. or shorter; stem longer than 1 m. , 
scandent or twining, rarely prostrate. 10. B. menziesii 

18. Leaves 4 cm. or shorter, 1.8 cm. or narrower, acute or 
attenuate at the base; flowers solitary or in simple 
cymes, never in dense clusters; stem 1 m. or shorter. 

16. B. sericea 

18. Leaves longer than 4 cm. , 2 cm. or wider, obtuse or 
truncate at the base; flowers in dense clusters (com- 
posed of numerous simple or compound cymes); stem 

longer than 1 cm. 4. B. thunbergiana 

19. Styles free for no more than one-third of length; stigma 
capitate or peltate; leaves, subtending flowers, nar- 
rowly oblong or oblong-lanceolate, with length-width ratio 
of 2 or higher. 8. B. elegans 

19. Styles free to the middle or lower; stigma globose or sub- 
globose; leaves, subtending flowers, ovate or elliptic, 

with length-width ratio of 1.8 or less. 20 

20. Corolla shorter than 2.5 cm., 2.3 cm. or narrower at the 
limb. 13. B. sulphurea 

20. Corolla 3 cm. or longer, wider than 2.5 cm. at the limb. 21 

21. Flowers in simple or compound cymes; pedicels as long as 
1 cm. or longer; leaves elliptic or ovate-elliptic, 3.5- 

6 cm. by 2.5-4 cm. 12. B. elliptica 

21. Flowers solitary or in simple cymes of 2 or 3; pedicels 
shorter than 1 cm. ; leaves ovate, rotund or ovate-sub- 
cordate, 2-3 cm. by 1.7-2.5 cm; if larger, long-mucronate 

at the apex. 22 

22. Leaves ovate or ovate-subcordate, widest near the base, 
with long raucros ; petioles 5 mm. or longer; stem slender, 
shorter than 2 m. 9. B. dietrichiana 



liiO P H Y T L G I A Vol. 17, no. 3 

22. Leaves orbicular or ovate-orbicular, widest at the 
middle; mucros, if present, very minutf; petioles 1-4 

nm. ; stem mostly 2 m. or longer. 11. B. grand i flora 

23. Corolla 3 cm. or longer; sepals wider than 5 mm. 

15. B. multicaulis 

28. Corolla shorter than 2 cm. ; sepals narrower than 5 mm. 24 

24. Leaves orbicular; corolla red, pink or white; stem 

erect or suborect. 30. B. rosea 

24. Leaves linear, oblong", linear-oblong, ovate, elliptic 
or rarely somewhat orbicular; corolla white or blue; 

stem prostrate or twining; if erect, loaves elliptic. 25 

25. Leaves cordate at the base, acute at the apex, with 
length-width ratio 1 or close to 1. 28. B. brevifolia 

25. Leaves obtuse, acute, truncate or subcordate at the base, 
obtuse, acute or emarginate at the apex, with length- 
width ratio more than 1; if 1 or close to 1, emarginate 

or obtuse at the apex. 26 

26. Leaves linear, linear-lanceolate or narrow-oblong, 

usually 5 mm. or narrower. 27 

26. Leaves ovate, ovate-oblong, obovate, elliptic, usually 
wider than 5 mm. 28 

27. Leaves narrow-oblong, rounded at both ends, sessile or 
subsessile, densely sericeous or villous. 

27. B. oblongifolia 

27. Leaves linear or linear-lanceolate, acute, obtuse or at- 
tenuate at the base, obtuse or acute at the apex; petiolate, 
sparsely or rarely densely sericeous. 26. B. linearis 

28. Leaves elliptic, 2.5-4.5 (5.5) cm. long, 1-2.5 cm. broad; 
stem erect or suberect; sepals spathulate. 32. B. velutina 

28. Leaves ovate, ovate-oblong, rarely elliptic, 1-2.3 cm. 
long, rarely longer, 8-15 mm. wide; stem prostrate or 
climbing; sepals ovate or ovate-acuminate. 29. B. media 

29. Sepals glabrous or merely ciliate at the margin, rarely 
sparsely pubescent; leaves glabrous or sparsely pubescent. 30 

29. Sepals, at least inner sepals, pubescent, tomentose or 
sericeous; leaves tomentose, sericeous or villous at 
least on the lower surface, rarely glabrate. 32 



1968 Lfyint & Ward, Revision of Bonamia liil 

30. Leaves oblong or oblong-elliptic; inflorescences umbel- 
late cymes of 5 or more flowers. 22. B. umbellata 

30. Leaves ovate or ovate-cordate; flowers solitary or 

in simple cymes of 2 or 3. 31 

31. Stem longer than 1 m. , twining or scandent; styles 
fused to the middle or higher; leaves glabrous, at- 
tenuate or acute at the apex. 34. B. balansae 

31. Stem 70 cm. or shorter, erect or suberect; styles 
free nearly to the base; leaves with scattered hairs, 
obtuse or truncate and mucronate at the apex. 

35. B. corumbaensis 

32. Corolla distinctly lobed ; stamens exserted; leaves 
with very distinct minor venations, undulate at the 

margin. 1. B. alterni folia 

32. Corolla entire, subentire or merely lobulate or angu- 
lar; stamens inserted; minor veins, except intercostal 
veins, scarcely distinct. 33 

33. Leaves narrowly elliptic, narrowly oblong or oblong- 
elliptic, narrower than 2 cm.; if wider, length-width 

ratio 1.5 or more and acute or attenuate at the base. 34 

33. Leaves ovate, ovate-cordate, broadly elliptic, oblong- 
ovate or oblong-cordate, 2 cm. or wider; if narrower, 
length-width ratio less than 1.5 and obtuse, truncate or 
subcordate at the base. 35 

34. Sepals 2-3 mm. long; corolla white, 1.8 cm. long or 
shorter; leaves with scattered hairs or nearly glabrous, 
obtuse or truncate at the base. 3. B. d ens i flora 

34. Sepals 4-6 (8) mm. long; corolla blue, 2 cm. or longer, 
rarely shorter; leaves distinctly pubescent at least on 
the lower surface, acute or cuneate at the base. 

2. B. spectabilis 

35. Individual flowers sessile or subsessile, occasionally 
with pedicels up to 1-2 (3) mm. long; peduncles of 
individual cymes absent. 36 

35. Individual flowers pedicellate or solitary and peduncu- 
late, with pedicels 3 mm. or longer; if shorter, pedun- 
cles present. 38 

36. Corolla 1.2 cm. or shorter; leaves glabrous above. 

20. B. brevipQdicellata 



1U2 P H Y T L G I A Vol. 17, no. 3 

36. Corolla 1.5 cm. or longer; leaves sericeous or tomen- 

tose above. 37 

37. Corolla white. 39. B. subsessilis 

37. Corolla purple, violet or with purple eye. 

40. B. mattogrossensis 

38. Stem 1 m. or shorter; if longer, leaves 3.5 cm. or 

shorter and rounded or obtuse at the apex. 39 

38. Stem mostly longer than 1.5 m. 40 

39. Stem erect or suberect, 30-60 cm. long; leaves sparsely 
pubescent or becoming glabrous. 35. B. corumbaensis 

39. Stem slender, prostrate, procumbent or climbing, longer 
than 70 cm. ; leaves densely sericeous or villous. 

17. B. boliviana 

40. Flowers solitary, rarely in simple cymes. 

43. B. langsdorffi i 

40. Flowers in simple or compound cymes, pseudopanicles 

or racemose panicles. 41 

41. Leaves glabrous on the upper surface, rarely with 
scattered hairs. 42 

41. Leaves sericeous or tomentose on the upper surface. 45 

42. Corolla 1.2 cm. or shorter; flowers in dense capitate 
clusters; styles shorter than 1 cm. 20. B. brevipedicellata 

42. Corolla 1.5 cm. or longer; flowers in loose cymes, 
panicles or pseudopanicles. 43 

43. Pedicels 1 cm. or longer; inner sepals rounded at 
the apex; outer sepals uniformly appressed sericeous. 

19. B. maripoides 

43. Pedicels shorter than 1 cm. ; inner sepals truncate or 
slightly emarginate at the apex; outer sepals tomentose 

or glabrate. 44 

44. Leaves with veins distinctly impressed above; inter- 
costal veins prominent at least on the lower surface; 
leaves mostly elliptic, 9 cm. or longer. 33. B. agrostopolis 

44. Veins not impressed on the upper surface; intercostal 
veins obscure, if distinct not impressed; leaves mostly 
ovate, if oblong or elliptic, blades 8 cm. or shorter. 

33. B. trichantha 



1968 l.*yint & V('ard, Revision of Bonamla lii3 

45. Leaves obtuse, rounded or truncate at the base; acuminate 

or acute, rarely obtuse at the apex. 46 

45. Leaves subcordate or rarely truncate at the base; ob- 
tuse, rounded or slightly emarginate at the apex. 

38. B. tomentosa 

46. Leaves elliptic or oblong-elliptic, 9 cm. or longer, 
sparsely pubescent or glabrescent above; ovary glabrous. 

36. B. agrostopolis 

46. Leaves ovate or ovate-elliptic, shorter than 8 cm., 

densely tomentose or sericeous, rarely sparsely so on 
the upper surface; ovary pilose at least at the apex. 

37= B. burchellii 



Regional Keys 

The following regional keys are given as supplementary to the 
general key because the identification of a specimen belonging to 
Bonamia is much simplified if the geographical source is known. 



Key to the African and Asian Species 

1. Outer sepals distinctly larger than inner sepals or with 
indumentum of dense, long, spreading hairs (drying golden 
brown); bracts mostly foliaceous. 2 

1. Outer and inner sepals equal, subequal or slightly un- 
equal; bracts mostly small, occasionally foliaceous 

(as in B. semidigyna ) 3 

2. Sepals, at least outer ones, obtuse, with short appressed 
hairs; leaves cordate or ovate-cordate; peduncles long, 
mostly 1 to 7-flowered; corolla white. 6. B. cordata 

2. Sepals acuminate or acute, with long spreading hairs; 
leaves elliptic-lanceolate to oblong-ovate; peduncles 
short, multi-florous ; corolla blue. 5. B. mossambicensis 

3. Sepals, both outer and inner, consistently acute or 
acuminate at the apex. 4 

3. Sepals obtuse or rounded, rarely broadly acute, at the 

apex. 8 

4. Peduncles 3 cm. or longer, rarely slightly shorter; 
leaves cordate or ovate-cordate, usually long-attenuate 

at the apex, 4 cm. or wider. 5 



]M PHYTOLOGIA Vol. 17, no. 3 

4. Peduncles shorter than 2 cm. ; leaves elliptic, ovato- 
elliptic, oblong-elliptic or oblong, rarely slightly cordate 
at the base, obtuse, acute or acuminate at the apex, 
narrower than 3.5 cm. 6 

5. Corolla white or yellowish white, with slightly lobulate 
limb; flowers mostly in simple or compound cymes. 

7. B. semidigyna 

5. Corolla red or pink, with entire or subentire limb; 
flowers mostly solitary, rarely in simple cymes. 

41. B. abscissa 

6. Flowers in cymes of few to several, usually forming 
secund dense clusters; leaves glabrous or sparsely 
pubescent on the upper surface. 4. B. thunbergiana 

6. Flowers solitary or in cymes of two or three; leaves 
densely velutinous or pilose on both surfaces. 7 

7. Corolla blue, longer than 2.5 cm.; sepals longer than 
10 mm. ; stems long, climbing or prostrate, weak. 

8. B. elegans 

7. Corolla white, shorter than 2 cm. ; sepals shorter than 

10 mm. ; stems short, erect. 32. B. velutina 

8. Stamens exserted; corolla distinctly lobed ; leaves 
undulate at the margin, strongly nerved, shortly 
petiolate or subsessile, with length-width ratio less 

than 2. 1. B. alternifolia 

8. Stamens inserted; corolla subentire or merely sub- 
lobulate; leaves entire at the margin, with indistinct 
minor venation, distinctly petiolate, with length-width 
ratio 2 or more. 9 

9. Sepals orbicular or ovate-orbicular, 3.5 mm. or shorter, 
sparsely pubescent or glabrescent; leaves sparsely 
pubescent or becoming glabrous, obtuse, rounded or 
truncate at the base; corolla white. 3. B. densiflora 

9. Sepals ovate or oblong-ovate, 4.5 mm. or longer, densely 
sericeous; leaves mostly sericeous, rarely becoming glab- 
rous, cuneate, subcuneate or acute at the base; corolla 
blue or bluish white. 2. B. spectabilis 



Key to the Australian Species 

1. Outer sepals obtuse, large, as wide as or wider than twice 
the width of inner sepals; third (or in-out) sepal 
oblique. 31. B. pannosa 



1968 l^nt & Ward, Revision of Bonamia lU5 

1. Outer and inner sepals equal, subequal or slightly un- 
equal; third (or in-out) sepal symmetrical. 2 

2. Corolla large, longer than 2.5 cm.; leaves mostly 3 cm. 
or longer, rarely shorter, 2 cm. or wider, mucronate at 

the apex. 9. B. dietrichiana 

2. Corolla small, 2 cm. or shorter; leaves 2.5 cm. or shorter, 
if longer, blades linear or oblong and narrower than 1.5 
cm., acute, obtuse or emarginate at the apex, if mucronate, 
mucro minute, shorter than 1 mm. 3 

3. Leaves linear, linear-lanceolate or narrow-oblong, usually 

5 mm. or narrower; length-width ratio mostly 3 or more. 4 

3. Leaves orbicular, ovate, ovate-elliptic, oblong-ovate 

or ovate-cordate, with length-width ratio less than 2.5. 5 

4. Leaves linear, acute or acuminate at both ends, rarely 
rounded at the base, with length -width ratio of 4 or 

more; petioles 1-4 mm. long. 26. B. linearis 

4. Leaves oblong or linear-oblong, obtuse or rounded at both 
ends, with length-width ratio less than 4; petioles 

0-1 mm. long. 27. B. oblongifolia 

5. Plant erect or suberect; leaves orbicular, rarely broadly 
ovate, emarginate, rounded or obtuse at the apex; sepals 
densely long-haired; corolla rose, pink or rarely white. 

30. B. rosea 

5. Plant prostrate or procumbent; leaves ovate, ovate-ellip- 
tic or ovate-subcordate, acute or obtuse, rarely emarginate 
at the apex; sepals finely pubescent or villous; corolla 
white or blue. 6 

6. Leaves cordate at the base and acute at the apex. 

28. B. brevifolia 

6. Leaves rounded, obtuse or subcordate at the base, ob- 
tuse or emarginate at the apex; if acute then base not 
cordate. 29. B. media 

Key to the American and Hawaiian Species 

1. Outer sepals twice the width of inner sepals or wider, obtuse 
or rounded in contrast to inner acute or acuminate sepals. 2 

1. Outer and inner sepals equal, subequal or slightly un- 
equal, acute, obtuse or acuminate at the apex, rarely 
slightly different. 4 



11^6 P H Y T L G I A Vol. 17, no. 3 

2. Pedicels longer than 1 cm. ; inflorescences loose cymes 
of 3-7 flowers , rarely more than 7 flowers ; peduncles 
long; secondary peduncles longer than 1 cm. 3 

2. Pedicels very short, rarely as long as 7 mm. ; inflores- 
cences dense capitate clusters of more than 10 flowers; 
peduncles short; secondary peduncles absent. 

21. B. ferruginea 

3. Leaves 5-12 cm. by 3-8 cm. , cordate or subcordate at the 
base; outer sepals 2 cm. by 1.7 cm.; styles free almost 

to the ovary. 24. B. kuhlmannii 

3. Leaves 5.5-7 cm. by 3-4 cm., obtuse or truncate at the 
base; outer sepals 8-12 mm. by 7-10 mm. ; styles fused 

at least lower one-third or half. 25. B. peruviana 

4. Sepals consistently, both outer and inner, acute or 
acuminate at the apex. 5 

4. Sepals obtuse, rounded or slightly emarginate at the apex. 13 

5. Flowers in dense capitate or umbellate cymes; individual 
flowers sessile, rarely shortly pedicellate. 6 

5. Flowers solitary or loose cymes of 3-7; individual 

flowers pedicellate. 7 

6. Leaves elliptic, oblong-elliptic, 4 cm. or shorter, 2 cm. 
or narrower, obtuse-mucronate or acute-mucronate at the 
apex, lanate at least on the lower surface; corolla 
shorter than 1.5 cm.; inflorescence a dense terminal 

head. 23. B. sphaerocephala 

6. Leaves ovate or ovate-lanceolate, longer than 4 cm. or 
wider than 2.5 cm.; acuminate or acute at the apex, glabrous 
or puberulous ; corolla 1.8 cm. or longer; inflorescence 
axillary. 18. B. holtii 

7. Leaves linear, linear-lanceolate, narrowly elliptic or 
narrowly oval-elliptic , with length-width ratio of 2 or 
more, usually narrower than 1.5 cm., rarely slightly 

wider. 8 

7. Leaves ovate, broadly elliptic or ovate-elliptic, with 
length-width ratio of less than 2; if 2 or more, stem long- 
er than 1 m. , usually wider than 1.5 cm. ; if narrower then 
ovate or orbicular. 9 

8. Corolla blue or purplish blue, 3 cm. or longer; sepals 7 
mm. or wider, densely villous, velutinous or tomentose; 
leaves densely velutinous. 15. B. multicaulis 



1968 L^int (Sc Ward, Revision of Bonaniia li;7 

8. Corolla white, shorter than 3 cm. ; sepals 6 mm. or nar- 
rower, finely or densely sericeous; leaves sericeous. 

16. B. sericea 

9. Sepals ovate or ovate-lanceolate, with length-width 
ratio of 1.5 or less, usually 11 mm. or shorter, 

densely tomentose, velutinous or villous. 10 

9. Sepals lanceolate or oblong-lanceolate, with length- 
width ratio of more than 1.5, glabrous, puberulous or 
finely sericeous. 11 

10. Corolla blue, 3 cm. or longer; leaves ovate or orbicu- 
lar, 2.5 cm. or shorter; stem 1 m. or shorter, pro- 
cumbent or suberect. 14. B. ovalifolia 

10. Corolla white or yellowish, 2.5 cm. or shorter; leaves 
elliptic, oblong-elliptic, oblong-lanceolate or rarely 
ovate or orbicular; stem longer than 1 m. , scandent 

or twining. 10. B. menziesii 

11. Corolla shorter than 2.5 cm., 2.3 cm. or narrower at 

the limb. 13. B. sulphurea 

11. Corolla 3 cm. or longer, wider than 2.5 cm at the limb. 12 

12. Flowers in simple or compound cymes; pedicels 1 cm. or 
longer; leaves elliptic or ovate-elliptic, 3.5 cm. or 
longer. 12. B. elliptica 

12. Flowers solitary or rarely in simple cymes; pedicels 
shorter than 1 cm. ; leaves ovate or ovate-orbicular, 

3 cm. or shorter. 11. B. grand if lora 

13. Sepals glabrous or merely ciliate at the margin, rarely 
sparsely pubescent; leaves glabrous or sparsely pubescent. 14 

13. Sepals, at least inner sepals, tomentose, sericeous or 

f errugineous , rarely glabrescent; leaves tomentose, sericeous 
or villous at least on the lower surface, rarely glabrate. 16 

14. Leaves oblong or oblong-elliptic; inflorescences um- 
bellate cymes of 5 or more flowers, rarely 3 flowers. 

22. B. umbellata 

14. Leaves ovate or ovate-subcordate; flowers solitary or 

in simple cymes of 2 or 3. 15 

15. Stem longer than 1 m. , twining or scandent; styles 
fused to the middle or higher; leaves glabrous, 

attenuate or acute at the apex. 34. B. balansae 



Ili8 P H y T L G I A Vol. 17, no. 3 

15. Stnm 70 cm. or shorter, oroct or suberpct; stylos free 
nearly to the base; leaves with scattered hairs, obtuse 

or truncate and mucronato at the apex. 35. B. corumbaensis 

16. Individual flowers sessile or subsessile, occasionally 
with pedicels up to 1-2 (3) mm. long; peduncles of 
individual cymes absent. 17 

16. Individual flowers pedicellate or solitary and pedun- 
culate, with pedicels 3 mm. or longer; if shorter, indivi- 
dual cymes or flowers pedunculate. 19 

17. Corolla 1.2 cm. or shorter; leaves glabrous above. 

20. B. brevipedicellata 

17. Corolla 1.5 cm. or longer; leaves sericeous or tomen- 

tose above. 18 

18. Corolla white. 39. B. subs ess ilis 

18. Corolla purple, violet or with purple eye. 

40. B. mattogrossens is 

19. Stem 1 m. or shorter; if longer, leaves 3.5 cm. 

or shorter, rounded or obtuse at the apex. 20 

19. Stem mostly longer than 1.5 m. ; if shorter, leaves 

acute or acuminate at the apex. 21 

20. Stem erect or suberect, 30-60 cm. long; leaves sparsely 
pubescent or becoming glabrous. 35. B. corumbaensis 

20. Stem slender, prostrate or climbing, usually longer 
than 70 cm. ; leaves densely sericeous or villous. 

17. B. boliviana 

21. Flowers solitary, rarely in simple cymes. 

43. B. langsdorffii 

21. Flowers in simple or compound cymes, pseudo- 
panicles or racemose panicles. 22 

22. Leaves glabrous on the upper surface, rarely with 
scattered hairs. 23 

22. Leaves sericeous or tomentose on the upper surface. 26 

23. Corolla 1.2 cm. or longer; flowers in dense capitate 
clusters; styles shorter than 1 cm. 20. B. brevipedicellata 

23. Corolla 1.5 cm. or longer; flowers in loose cjmies , 

panicles or pseudopanicles. 24 



1968 Iifyint & Ward, Revisicm of Bonamia lli9 

24. Pedicels 1 cm. or longer; inner sepals rounded at 
the apex; outer sepals uniformly appressed 
sericeous. 19. B. maripoides 

24. Pedicels shorter than 1 cm. ; inner sepals truncate 
or slightly emarginate, rarely rounded at the apex; 

outer sepals toraentose or glabrate. 25 

25. Veins distinctly impressed on the upper surface; 
intercostal veins prominent at least on the lower 
surface; leaves mostly elliptic, 9 cm. or longer, 

acute or obtuse at the base. 36. B. agrostopolis 

25. Veins not impressed on the upper surface; inter- 
costal veins obscure; leaves mostly ovate; if oblong or 
elliptic, blades 8 cm. or shorter, rounded, obtuse 

or subcordate at the base. 33. B. trichantha 

26. Leaves subcordate or rarely truncate at the base; 
obtuse, rounded or slightly emarginate at the apex. 

38. B. tomentosa 

26. Leaves obtuse, rounded or truncate at the base; 

acuminate or acute, rarely obtuse at the apex. 27 

27. Leaves elliptic or oblong-elliptic, 9 cm. or longer, 
sparsely pubescent or glabrescent above; ovary 

glabrous. 36. B. agrostopolis 

27. Leaves ovate or ovate-elliptic, shorter than 8 cm. , 
densely toraentose or sericeous, rarely sparsely so 
on the upper surface; ovary pilose at least at the 
apex. 37. B. burchellii 



I. Section: Bonamia 

Stems woody or becoming woody, occasionally slender, twining, 
scandent, prostrate, procumbent or suberect, commonly long, rarely 
slightly shorter than 1 m. , occasionally attaining several meters 
long. Leaves petiolate or shortly petiolate, soft, herbaceous, 
subcoriaceous or somewhat leathery, rarely thin; blades mostly 
elliptic, elliptic-oblong, ovate, cordate or rarely lanceolate 
or oblong, usually large, 2.5 cm. or longer, 1 cm. or wider, rare- 
ly slightly short or slightly narrower, rounded, truncate, sub- 
cordate or cordate, rarely slightly attenuate at the base, acumi- 
nate, obtuse or acute at the apex. Flowers solitary or in cj^es 
of few to several flowers, in cj^mose panicles or in capitate or 
dense clusters, usually pedunculate; individual flowers distinctly 
pedicellate; pedicels short or up to 2 cm. long; bracts small or 
foliaceous. Sepals subcoriaceous or coriaceous, equal or unequal, 
often large, ovate, lanceolate, oblong-ovate or ovate-acuminate, 



150 P H y T L G I A Vol. 17, no. 3 

acute or obtuse, occasionally short -acuminate at the apex. Corolla 
blue, purplish blue, white or red, usually large, 2 cm. or long- 
er, sometimes slightly shorter, subentire, lobulate or lobed at 
the margin. Stamens included or partially exserted; filaments 
sparsely or densely long-villous or pilose, often glabrous or near- 
ly glabrous; anthers 2 mm. or longer, sagittate or cordate at the 
base. Ovary ovoid or ovoid-conical, long pilose or glabrous; 
styles free nearly to the base or fused to the middle or higher 
with a single vascular bundle fup to stigma) in each stylar branch; 
stigmas small or large, globose, subglobose, conical or capitate. 
Fruits 4- to 8-valvular capsules, thin-walled, rarely 0.5 mm. 
thick; seeds glabrous, smooth or punctate, brown or black, 3-6 
mm. long, rarely smaller. Cotyledons ovate, ovate-cordate, obo- 
vate or linear-bifid, corrugate, multiplicate or slightly folded. 

Type: B. alternifolia J. St. Hilaire. 

Tropics, subtropics and warm temperate of both hemispheres, 
covering the whole range of the genus, occurring on all continents. 

This section is less homogeneous than the other two sections 
because of inclusion of several species whose morphology is in- 
completely known. Future studies may lead to separating it into 
more than one section or subsection. 

1. Bonamia alternifolia J. St. Hilaire, Expos. Fam. 2:349. 1805. 
Bonamia madagascariensis Poir. Encycl. Meth. Bot. Suppl. 

1:677. 1810. 
Bonamia thouarsii Elliot, Journ. Linn. Soc. Bot. 29:35. 

1891. 

Perennial shrubs or woody vines. Stems erect or suberect, 
with weak branches, terete, finely pubescent or villous while 
young, becoming sparsely pubescent or glabrescent in age, 1.5- 
1.8 m. high. Leaves shortly petiolate, coriaceous or subcoriaceous , 
sparsely appressed-pubescent, more densely so while young; 
petioles 2-7 mm. long, brown-villous ; blades elliptic, ovate-acumi- 
nate or obovate, 3-7 cm. long, 2-4 cm. wide, undulate at the mar- 
gin, acute or obtuse at the base, obtuse-acuminate, obtuse or 
acute-mucronate at the apex; midrib prominent, with 3-5 pairs 
of prominent lateral veins; finer veins distinct, clearly visible 
with naked eye, especially on the lower surface. Inflorescence 
commonly congested terminal panicles of numerous flowers or few- 
flowered cymes in axils of upper leaves, shortly pedunculate; 
peduncles short, 5-10 mm. long, villous or finely pubescent; 
pedicels as long as peduncles or slightly longer, brown-villous; 
bracts subulate, 2-4 mm. long or sometimes smaller, deciduous 
or persistent. Sepals orbicular or ovate, coriaceous, villous, 
unequal or subequal; the two exterior smaller, mostly 4.5-5.5 
mm. long, orbicular, slightly emarginate or rounded at the apex; 
the interior longer, 6-7 mm. long, rounded or rarely obtuse at 
the apex. Corolla white, tubular campanulate or funnel-shaped, 
with cylindrical narrow tube and distinctly lobed limb, indup- 



1968 Ltyint & Ward, Revision of Bonamia l5l 

licate in the bud, 1.4-2 cm. long, pilose on interplicae and 
upper part of tube; corolla lobes 2-3 mm. long, mostly 3 mm. wide, 
rounded or obtuse at the apex. Stamens exserted; filaments ad- 
nate to the corolla tube, glabrous above, with scattered long 
hairs below, distinctly longer than styles; anthers dorsifixed, 
versatile, introrse, oblong, 2.5-3 mm. long. Ovary conical, 
glabrous; styles free to the base, glabrous; stigmas capitate, 
warted or rugose. Capsules with tightly appressed sepals, 
glabrous, slightly woody, ovoid, cuspidate or apiculate, 2- to 
4-seeded, apparently two-valved (really four-valved) , somewhat 
woody; seeds glabrous. Cotyledons ovate, folded. 

Type: Madagascar. Type specimen not seen, presemably at 
Paris. 

This species is endemic to Madagascar, where it grows at 
low altitude (Map 1). One collector ( Mocquerys 176 ) noted forest 
as the habitat of this species. It has been collected in flower 
in November. The flowering period otherwise is not definitely 
known. 

Bonamia alterni folia is somewhat related to B. spectabilis 
but differs from it in many features. It is a very recognizable 
species of the genus with the following distinctive characteris- 
tics: (1) strongly nerved leaves with undulate margin, (2) ter- 
minal congested panicles, (3) closely appressed sepals, (4) dis- 
tinctly lobed corolla with narrow limb and long tube, (5) exser- 
ted stamens with versatile anthers, (6) styles free to the 
base, (7) warted or rugose stigmas, (8) somewhat woody capsules 
with slightly appressed sepals, and (9) glabrous seeds. Of 
these several differences from other species in the genus, 
lobed corolla and exserted stamens are most outstanding and were 
paid serious attention by earlier botanists in maintaining it as 
a monotypic genus. Its slightly woody capsules (which dehisce 
into two valves) indicate a slight affinity to the members of 
section Trichantha, but it differs from the latter by its 
glabrous seeds. 

This species is the type species of the genus, first des- 
cribed by Thouars (1805) without a binomial name. In the following 
year J. St. Hilaire published Bonamia alternifolia for the plant 
Thouars had described. Although this binomial was published 
earlier for the species later described as B. madagascariensis 
Poiret (1810) , the latter name has replaced it in most recent 
literature. This adoption of the later name is probably owing 
to the fact that St. Hilaire 's publication was rare, and since 
the name was included only in the appendix of Index Kewensis , 
it was overlooked. House (1907) first found this overlooked 
name and designated it as the type species of the genus. But 
the compiler of conserved generic names did not note House's desig- 
nation of the type, thus resulting in the citation of B. madagas - 
cariensis as the type species of Bonamia in the International Code. 



PIIYTOLOGIA 



Vol. 17, no. 3 




Distribution of 
® B. alternifolia * B. mossarabicensis 



• B. spectabilis 



•Jf- B. velutina 



Map 1 



1968 Ityint & '(Yard, Revision of Bonania 153 

B. alternifolia must be reinstated as the correct name for the 
species, and the improper choice of name in the Code should be 
corrected in the future. 

Specimens examined: 

MADAGASCAR: Soanierana, Rivieroever, liana, bl. wit, welrie- 
kend, Lam en Meeuse 5548 , 30.11.1938 (L) ; Maroa , Forets a I'inter- 
ieur de la baie d'Antongil, Arbre mince, elance, fleurs blanches, 
A. Mocquerys 176 , 1897 (G) ; M. Richard de Bourbon 1966 (G). Un- 
known collector: N. de Madagascar, #367 (L). 

2. Bonamia spectabilis (Choisy) Hall. f. Bot. Jahrb. 16:529. 1893. 
Breweria spectabilis Choisy. Mem. Soc. Phys. Geneve 8:68 

1839. 
Breweria hildebrandtii Vatke, Linnaea 43:523. 1882. 

Type: Hildebrandt 2903 
Bonamia hildebrandtii (Vatke) Hall. f. Bot. Jahrb. 18:91 

1893. 
Bonamia minor Hall, f. Bot. Jahrb. 18:91. 1893. 

Type: Pogge 1214 
Bonamia minor var. argentea Fries, Wiss. Ergebn. Schwed. 

Rhod.— Kongo— Exped. 1:268. 1916. Type: Fries 827 

Perennial, woody, climbing or twining, rarely trailing vines, 
growing all year around. Root thick, woody; stems terete, fre- 
quently ridged, sparsely appressed-pubescent , becoming glabrous in 
age, about 1.5-2.5 mm. in diameter. Leaves petiolate, membranous 
to subcoriaceous , glabrescent and green or silvery pubescent above; 
moderately or densely silky pubescent below; petioles mostly 5-12 
mm. long, slightly winged and canaliculate above, minutely appressed- 
pubescent or becoming glabrous in age; blades elliptic, 1.8-5.5 cm. 
long, occasionally shorter, 6-20 mm. wide, sometimes slightly wider, 
entire at the margin, attenuate or cuneate at the base, acute or 
obtuse and apiculate at the apex, with about 5-8 pairs of lateral 
veins. Inflorescences shortly pedunculate cymes of 2-3 or few 
flowers , aggregated towards the end of branchlets ; peduncles and 
pedicels mostly 5-10 mm. long, pubescent or glabrescent; bracts 
small. Sepals oblong-elliptic, 7-8 mm. long, abruptly acute, rare- 
ly obtuse, silky pubescent outside. Corolla blue, funnel-shaped, 
(1.5) 2-2.5 cm. long, silky pilose on interplicae, entire or sub- 
entire. Stamens included; filaments dilated and hairy below; an- 
thers oblong with cordate base. Ovary glabrous; style bifid from 
about the middle or lower; stigmas ellipsoidal, rugulose. Capsule 
globose, about 7 mm. in diameter, shortly apiculate, glabrous. 
Seeds ovate-oblong, compressed on the inner side, 3-4 mm. long, 
brownish or blackish, with hyaline golden wings on edges. Cotyle- 
dons oblong, deeply bifid; cotyledonary petioles fused. 

Type: Madagascar, Bombatok, Bo j er (K - lectotype, not avail- 
able; W - isotype.') 

Republic of Congo, Northern Rhodesia, Tanganyika and Madagascar 
(Map 1). 



l^U P H Y T L G I X Vol. 17, no. 3 

Colloctors recorded i-dgo of donso forest, savannah forost on 
stoep rockhills, mixed woodlands on sandy slopr.-s, and degraded 
thickets on Kalahari sands at edge of river flats as habitats of 
this species. It has been collected in flower from March through 
December and in fruit from July through Dfcrmiber. 

Hallier (1897), after careful study of the Madagascarian species, 
treated B. hildebrandtii as conspecific with B. spectabilis , al- 
though he had accepted them previously (1893) as distinct species. 
He retained B. minor which he described in his earlier work. The 
type specimen of B. minor has not been seen in the present study, 
but a number of sheets which Hallier annotated have been examined. 
Hallier used the glabrous stem as a principal feature to distinguish 
it from B. spectabilis . Examination of sheets annotated by him 
shows pubescence to be consistent only if they are compared with 
specimens annotated by him as B. spectabilis . If one considers 
the additional collections now available it seems apparent that 
the characteristic which Hallier mentioned falls well within the 
total range of variation of a single species. Verdcourt (1963), 
realizing this fact, remarked, "Hallier unites B. hildebrandtii 
and B. spectabilis but retains B. minor as distinct. B. hilde- 
brandtii is, however, undoubtedly identical with B. minor "' 

This species is highly polymorphic in several features, 
particularly leaf shape and size (Figure 1) , and villosity on 
stem and leaves. Future collections might reveal consistent 
features to account for the infraspecific groups in it. 

Specimens examined: 

REPUBLIC OF CONGO: KATANGA: Kasenga, W. Robyns 1845 , 3. IV. 
1926 (K). 

MADAGASCAR: Env. de Majunga, CD. Alleizette , 30. XI. 1906 
(L) ; Central Madagascar, R. Baron 4906 (BM); Seandit in sylvis 
juxta Mazangay in Bombatok ora occidentalis ins. Madagas: Flor. 
Aug. 5. Flores in spicam longam congregati Cyanei , Bo j er , II. q. 
1830 (W); Nosse-be, J.M. Hildebrandt 2903 , April 1879 (BM, L, W) ; 
Beravi interior: Gebrige, fl. albi , Hildebrandt 3093 , July 1879 
(BM, W) ; Nosifaly 8 (L) ; lies Maurice, de Madagascar et Coraorres , 
Mac William , Aout-Octobre 1838 (G). 

NORTHERN RHODESIA: Barotse: Sesheke, climbing over small 
shrubs on edge of dense Baikiaea "mutemwa", Longe Forest, N. of 
Machile, climber with bright blue flowers, A. Angus 956 , 19. 12. 
1952 (EA); on Kalahari sand in open degraded Baikiaea ''mutemwa" 
on edge of Kazu Forest near Machile, suffrutex with woody root- 
stock and numerous small shoots together, flower blue, Angus 983, 
20. 12. 1952 (EA); Abercorn Dist. 2700 ft. B.D. Burtt 6325 , 20. 5. 
1936 (BM, EA) ; Barotseland, Nangweshi, 3400 ft., semiwoody climber 
in mixed woodlands on sandy slopes, blue flowers, L.E. Codd 7156 , 
23. 7. 1952 (L); between Pemba and Mazabuka, I.B. Pole Evans 2807 , 
11. 7. 30 (K) ; L. Mweru Dist. common vine scrambling over ever- 
green thicket, blue, showy with paler or white guide lines, D.B. 
Fanshawe F-4653 , 6. 8. 1958 (EA); Abercorn Dist., A.H. Gamwell 68 
(BM); Abercorn Dist. Alt. 4800 ft. , A.H. Gamwell 94, August 1935 



1968 



^lJ'lnt & 'i7ard. Revision of Bonamia 



155 



15 




Figure 1 19 

Variations in sizes and shapes of leaves in Bonamia spectabilis 



156 P H y T L G I A Vol. 17, no. 3 

(BM); Kafuo 35 mi. s. of Lusaka, noar Kafuc Mr-thodist Mission, 
roadside, climbing over shrubs and on red soils, A.C. King 55 , 
11 July 1955 (K); Mpulunga Road ]0 mi. from Abercorn, 4000-5000 
ft., herbaceous climber, flowers delicate sky blue, leaves dull 
green, pubescent, R.M. Lawton 208 , June 1955 (LA); roadside, 
Kafue River, flower blue, plant mat forming, I/?ach and Brunton 
9997 , 12. 6. 60 (EA); J.D. Martin 196 (K); Abercorn Mpulunga Road, 
not far from Mukoma turning, alt. 4000 ft. on bank and trailing 
over low bushes, in a tangled mass, Mrs. H.M. Richards 5307 , 5. 
4. 1955 (EA); Mporokoso Dist. , Sumba Malango Road, 900 m. in 
sandy soil on side of road, trailing plant, Mrs. Richards 6264 , 
24. 9. 1956 (EA); 26 mi. north of Choma, dry bush by roadside, 
climber rising on shrubs to height of 12 ft. , E.A. Robinson 767 , 
17. V. 1954 (K) ; southern prov. , Mazabuka Dist., Choma to Lusaka, 
Gt. North Road, mile 34, growing in Pterocarpus angolensis — Com - 
bretum mechowianum woodland, F. White 2285 , 19. III. 1952 (LA); 
Namwala Dist. , 20 mi. west of Mamwala Boma, growing in degraded 
thicket on Kalahari sands at edge of Kafue Flats, White 2962 , 22. 
VI. 1952 (EA). 

TANGANYIKA: Mpanda Dist., Masigo, Mulele Hills Forest Reserve, 
alt. 4500 ft. , blue flowered creepers in thicket, J. Proctor 2086 , 
July 1962 (TA)', Ufipa Dist., Kasanga, alt. 840 m. , side of road 
in very gritty soils, climbing over low bushes, Mrs. Richards 10093 , 
13. 6. 1957 (EA, K) ; Escarpment above Kasanga, alt. 900 m. , climb- 
ing over dense vegetation by twining stems, Mrs. Richards 11001 , 
30. 3. 1959 (EA). 

3. Bonamia d ens i flora (Baker) Hall. f. Bull. Herb. Boiss. 5:999. 
1879. 

Breweria densiflora Baker, Journ. Linn. Soc. Bot. 25:336. 
1890. 

Perennial twining vines. Stems slender, woody or becoming woody, 
terete, finely striated or smooth, finely pubescent and glabrescent; 
internodes mostly 2-5 cm. Leaves shortly petiolate, soft-subcoria- 
ceous or herbaceous, sericeous with very fine, soft, appressed hairs 
when young, becoming sparsely sericeous or nearly glabrous (except 
on veins) in age; petioles 2-4 mm. long, 0.5 mm. thick, sparsely 
sericeous with soft hairs; blades oblong-elliptic or lanceolate- 
elliptic, 3-4.8 cm. long, 1-1.4 cm. broad, rounded or subtruncate 
at the base, obtuse-mucronate or acute-mucronate at the apex; midrib 
slightly impressed above, distinct beneath, with 6-8 pairs of thin 
lateral veins. Inflorescences axillary, pedunculate, umbellate 
cymes of three to seven flowers or terminal panicles; peduncles 
slender, short, 1-2 cm. long, finely pubescent or becoming sparsely 
so; pedicels slender, 3-6 mm. long, becoming slightly longer in 
fruit-bearing stage, sparsely soft-sericeous; bracts minute, linear, 
0.5-1.5 mm. long. Sepals ovate or ovate-orbicular, rounded or 
slightly emarginate at the apex, coriaceous or subcoriaceous , soft- 
sericeous outside, slightly unequal; outer two ovate, mostly 3 mm. 
long, 2 mm. broad, and rounded at the apex; inner three orbicular 
or ovate-orbicular, 3-4 mm. long, mostly 3 mm. broad and slightly 
emarginate at the apex, scarious at the margin. Corolla white. 



1968 Ityint & Vfard, Revision of Bonania 157 

shortly tubular-campanulate or funnel-shaped, 1.2-1.8 cm. long, 
densely soft -pilose on interplicae. Stamens inserted; filaments 
filiform and glabrous above, dilated and villous below; anthers 
oblong, 1.5-2 mm. long, sagittate at the base. Ovary ovoid-conical, 
with distinct circular disc at the base, glabrous; styles filiform, 
glabrous, bifid for upper one-third or one-fourth; stigmas small, 
capitate. Capsules globose-subacute, 4-6 mm. long, apiculate, 
glabrous, 2- to 4-seeded, 4 valvular; seeds ovate-oblong, 3 mm. 
long, black or dark brown. Cotyledons deeply bifid; cotyledonary 
petioles short. 

Type: Madagascar, R. Baron 5868 (K-not seen). 

Known only from Madagascar. 

Collectors give no definite location nor habitat. It was 
collected only a few times during the last century, and no recent 
collection is available for the present study. The species is 
poorly known, and the above description is mainly based on a 
single specimen and the original description by Baker. 

The outstanding features of this species are (1) slender and 
sparsely sericeous stem, (2) softly sericeous and glabrescent 
leaves, thin in texture, (3) ovate-orbicular or orbicular and 
small sepals and (4) smaller corolla. It is a close relative of 
B. spectabilis from which it can be distinguished by its finely 
sericeous and glabrescent leaves, smaller sepals and shorter 
corolla. ^Vhen a larger number of specimens of this species, show- 
ing more completely the variations to be found in it, is avail- 
able in the future, its separation from B. spectabilis might be 
reconsidered. 

Specimen examined: 

MADAGASCAR: "N. de Madagascar, No. 213" unknown collector (L). 

4. Bonamia thunbergiana (Roem. et Schult. ) Williams, Bull. Herb. 
Boiss. (ser. II) 7:371. 1907, 
Convolvulus Thunbergianus Roem. et Schult. , Syst. Veg. IV: 

884. 1819. 
Convolvulus C3miosus Thunberg ex Roem. et Schult. , Syst. Veg. 

IV: 303. 1819; not C. cymosus Desr. in Lamarck Encycl. 

Meth. Ill: 556. 1792. 
Bonamia cymosa (Roem. et Schult.) Hall. f. Bot. Jahrb. 18: 

91. 1893. 
Convolvulus senegambiae Spreng. , Syst. Veg. 1:610. 1825. 
Ipomoea senegambiae Choisy, in DC Prodr. 9:351. 1845. 
Ipomoea secunda Don, Gen. Syst. IV:282. 1838. 
Breweria secunda Benth. ^D Hook. Niger Fl. 470. 1849. 

Perennial, woody climber reaching 4 m. long. Stems twining, 
terete, 1.5-4 mm. thick, pubescent with brown hairs, densely so 
while young, becoming glabrous in age. Leaves petiolate, sub- 
coriaceous or membranous , green and glabrous or rarely thinly 



158 P H Y T L G I A Vol. 17, no. 3 

puberulous above, donsoly pubescent with golden brown hairs be- 
neath; petioles mostly 5-13 mm. long, 1 (-1.5) ram. thick, pubes- 
cent; blades oblong to oblong-lanceolate, about 2.5-8.5 cm. long, 
1.5-3.5 cm. broad, entire at the margin, rounded at the base, ob- 
tuse-mucronulate, rarely acute or acuminate at the apex; nerves 
sunken above, prominent beneath; lateral nerves about 6-10 pairs. 
Inflorescences dense cymes of many flowers, usually secund on 
short peduncles or congested into a terminal panicle; peduncles 
5-15 mm. , tomentose; pedicels 5-10 mm, tomentose; bracts minute, 
lanceolate. Sepals oblong-lanceolate to ovate, acuminate or 
acute at the apex, about 6-8 mm. long, the inner slightly shorter, 
coriaceous to glumaceous , densely silky tomentose on the back. 
Corolla white, 1.6-2 cm. long, obscurely lobed or subentire; out- 
side pilose or hirsute on the interplicae, glabrous on the plicae. 
Stamens included; filaments unequal, filiform, widening toward 
the base, glabrous above, pilose along the edge near the base; 
anthers oblong, cordate at the base. Ovary ovoid, with a disc 
at the base, pilose near the apex; style bifid above the middle, 
with scattered long hairs; stigmas conical, rugose. Capsule 
ovoid, 8-valved , 4-seeded, rarely less, apiculate, glabrous, 
about 5-8 mm. long; seeds black, ovate-oblong, glabrous. 

Type: Sierra Leone. 

Coastal districts of tropical west Africa, from Gambia, French 
Guinea, Sierra Leone, Liberia, Ivory Coast, Gold Coast, Nigeria, 
Cameroun and the western part of Congo Republic (?) (Map 2). 

It has been collected in flower from November to May and in 
fruit from December to April. 

Hallier in his Convolvulaceae of Africa and monograph of the 
genus, used the specific epithet cymosa , since he had overlooked 
Roemer and Schultes' correction of c3^osus to thunbergianus in 
the errata at the end of the volume. As pointed out by Willieims 
(1907) , the epithet cymosus is illegitimate since it is pre- 
occupied by C. C3mos us of Desrousseaux. The species was first 
collected by Thunberg in Sierra Leone and was named in his honor. 

Specimens examined : 

CAMEROUN: Gross-Batanga, M. Dinklage 684 , Ende Juni 1890 (HBG); 
Dinklage 684 , 16. XI. 1890 (HBG); Bipende, Urwaldgebiet, G. Zenker 
4112 , IQiriBM). 

FRENCH GUINEA: Kouakry, Boue 38 , II. 1910 (G) ; Maraou, G. 
Roberty 10600 , 9. 2. 1948 (G). 

FRENCH WEST AFRICA: Adiopodoume N (B. 30. 8. Df . ) , Roberty 
15655, 24. 11. 1954 (G) ; U Zo, NW. (B. 29. 22. Bd.), Roberty 16076 , 
17, 12. 1954 (G). 

IVORY COAST: Mau, Roberty 6737 , 26. 12. 1946 (G) ; Basse cote 
d'lvoire, secteur cotier, boiseraents perilagunaires , Dabou, Roberty 
13626 , 1950-51 (G). 

LIBERIA: Along Dukwia R. , vine with white flowers, common name 
Doo, G.P. Cooper 220 , 1929 (F, NY, US); Monrovia, Max Dinklage 2148 , 



1968 



l^int & V/ard, Revision of Bonamia 



159 




Distribution of B. thunbergiana 

Based on specimens seen in the 
present study 

O Based on records in Hutchinson 
and Dalziel's Flora of West 
Tropical Africa (1931) 



160 P H Y T L G I A Vol. 17, no. 3 

1903-04 and 2399 , 9 Jaunar. 1909 CB) ; 3 mi. north-east of Suacoco, 
Gbarnga, central prov. , thicket, gravelly soil, vine 9 ft. tall, 
V.P. Konneh 125 , Feb. 17, 1951 CMO) ; within a radius of 20 mi. 
from Kakatown, A. Whyte , April 1904 CBM, W). 

NIGCKIA: F3aikie's Nigeria Expedition CLagos), C. B arter 
2227 , 1857-59 f GH) ; main road from Oron to Eket— 28 mi., mostly 
farm clearing, Hket Dist. , southern Nigeria, P. A. Talbot, 1912- 
13 (BM); white, tinted palest mauve at edge of corolla, Oban, 
P. A. Talbot 88A f=1535), 1909 TBM). 

SIERRA LEONE: Afzelius CBM); R.H. Bunting , 1913-14 (BM); 
Njala, Allison v. Armour Expedition 1926-27, J.M. Dalzi el 8048 , 
20. 1. 27 (US); Makump, roadside and old farms, creeper with 
white flowers, Temne~"Rakil," R.R. Glanville 108 , 9. 12. 1928 
(K); Mamansu, V. Marmo 145 , 30. 11. 1958 (K) ; bush near Regent, 
Mahera, Kitchom, G.F. Scott-Elliot 3930 , Doc. 6. (BM, GH) ; Scott - 
Elliot 5835 , March 4, 1892 (BM); climbing over bushes near Madina, 
Scott-Elliot 5865 (BM); Smeatman (BM); Rornks , alt. 200 ft. , 
N.W. Thomas 5780 , 24. 11. 1914 (EA); Magvile, alt. 100 ft., Thomas 
6385 , Dec. 8, 1915 (BM); Thomas 6604 , 1915 (B), 6818 , Dec. 23 - 
Jan. 2, 1914-15 (B) , 6860 , 1915 (B) , 7051, Dec. 31, 1914 (B) , 
8513 , 1915 (W). 

(?) CONGO REPUBLIC: oubangui, Reg. de Tanga, Herb, de C. 
d'Alleizette, May 1920 (L). 

5. Bonamia mossambicensis (Klotzsch) Hall. f. Bot. Jahrb. 18:91. 
1893. 
Prevostea mossambicensis Klotzsch J_n Peters, Reise Mossamb. 

Bot. 1:244, t. 39. 1861. 
Breweria buddleoides Baker, Kew Bull. 1894:69. 1894. 

Perennial, shrubby climbers to 4-5 m. Stems terete, velvety 
with patent and tangled hairs, white or grey in life, golden-brown 
when dry. Leaves shortly petiolate, coriaceous or subcoriaceous , 
velvety pubescent, more densely so beneath; petioles 4-10 mm. long, 
velvety pubescent with hairs similar to those on the stems, blades 
elliptic lanceolate to oblong-ovate, 2.5-8.5 cm. long, 1-4.8 cm. 
wide, rounded or slightly cordate at the base; acute, acute-mucron- 
ulate, acuminate or apiculate at the apex; veins distinctly de- 
pressed above, prominent below; about 6-10 pairs of lateral veins. 
Inflorescences capitate, bracteate, densely hirsute, shortly pedun- 
culate or nearly sessile; peduncles up to 3.2 cm. long; pedicels 
almost absent; bracts elliptic or oblong-elliptic, 1-1.2 cm. long, 
5-8 mm. wide, hirsute outside, almost glabrous inside. Sepals 
unequal, coriaceous except near the apices of outer ones; the two 
exterior larger and hirsute near the apices. Corolla blue, funnel- 
shaped, 2 cm. long, subentire or almost entire; outside pilose on 
interplicae, glabrous on plicae. Stamens included; filaments 
short, unequal, adnate to corolla tube, glabrous or with scattered 
hairs; anthers oblong, about 2 mm. long and 1 mm. broad. Ovary 
hairy at the apex; style bifid above the middle; stylar branches 
unequal; stigmas ovoid or globose, rugose. Capsule 4-valved , 2 
(-4)-seeded, thin-walled, hairy outside, at least at the apex; 
seeds ovate-oblong, 2-3 mm. long, dark brown with narrow hyaline 
wings on edge. Cotyledons ovate-cordate. 



1968 LJyint & V/ard, Revision of Bonamia 161 

Type: Mozambique, Sena, Peters ( B-holotype-not available). 

Restricted to thickets and secondary evergreen forests on loam 
and sand in coastal districts at the altitude of 120-450 ra. in 
Mozambique and Tanganyika (Map 1). 

The outstanding characteristics of this species are: (1) 
densely velvety pubescent leaves and stems, (2) foliaceous bracts, 
(3) capitate, densely hirsute inflorescences, (4) absence of pedi- 
cels and (5) short peduncles. 

Specimens examined: 

MOZAMBIQUE: Port Amelia, white, R. Dummer 64 , July 1913 (Bh); 
Niasa Dist. , Port Amelia, 150 ft. "fl. sky blue," J. Gerster 7172, 
24. 6. 1949 (L, K) ; 4 mi. west of Lumbo; pale mauve blue, L.C . 
Leach and Rutherford-Smith 10944 , 21. 5. 1961 (EA); 11 km. on 
the road to Monapo on light sandy soil, Pedro-Pedrogar 3139 , May 
5, 1948 (EA) ; between Femad Veloso and Nacala on red sandy soil 
in dense secondary bush-thicket, Pedro-Pedrogar 4813 , Aug. 15, 
1948 (EA); Trepadeira de flores azul-purpures , Mocimboa da Praia, 
entre Diaca e Meuda, Pedro-Pedrogar 5216 , Sept. 15, 1948 (EA). 

TANGANYIKA: Orero-Kilwa Kivindje, Braun 1304 , 4. 11. 1906 (EA); 
Daressalaeun Dist., Puguhills ; exposed banks and railway cuttings; 
trailing or rambling habit to 4 ft. ; fl. pretty sky blue, leaves 
silver green, B.D. Burtt 4470 , 25. 4. 1933 (K) ; W. Busse 2565 (1903) 
(EA) and 2467a (EA) ; Kisarawe, Karonzurir (Kizararao), a scandent 
shrub with clusters of pale blue flowers , very common with Dicha - 
petalum spp. cuid Acacia pennata in Antidesma , Xylopia , Trema, 
Diospyros , Enclea ; secondary evergreen forest on red sandy soils, 
1000 ft. alt., P.J. Greenway 4993 , 1. 8. 1937 (EA); Lindi, Mkae 
Plantation; blue creeper which affects badly most of the land 
on the estate. Manager 14 , 5. 1932 (EA) ; Tandagura to Lindi, foot 
of Notoplateau, alt. 900 ft., old farm land, climber furry stalk 
and leaves, fl. terminal buff bracts, blue, monopetal, F.W.H. Migeod 
812 and 813 , 22. 8. 30 (BM); Usaramo, Puguberge, bem. 21. 5-24, 5, 
entland der Bahnstrecke, blace, A. Peter 31316 , 24. X. 1926 (B); 
Daressalam — Mbagara — see, P. Schlingt, blau, Peter 44927 , 5. IX. 
1926 (B); Bagamoyo — Mapinga, Meist Verblicht, tila, Peter 51646 , 

5. XII. 1915 (B); Usaramo bei Toga, Peter 51649, 13. XII. 1915 (B); 
Usambara, Bwiti Urwald bei Maramba , Blau, ca. 280 m. , Peter 51705 , 

6. VI. 1917 (B); Mahenge, Sali, ca. 35 km. sudlich Station Mahenge 
Savanne and Bushland , 900-1000 m. scclinger, vereinzelt, Blute 
blau, H.J. Schlieben 2242 , 24. 5. 1932 (B, BM, G, HBG) ; 40 km. 
west of Lindi, 240 m. u. M. Lutamba-see, schlinger in gr. Gruppen 
uber Stranchern sehr haufig, Schlieben 5193 . 29. 8. 1934 (B, BM, 
G, HBG); Usaramo, Stuhlmann 105 , 18. VIII. (18)88 (HBG); Pugu 
Hills, powder blue flowers, very common all up the road, J.H. 
Vangham 2340 , April 13, 1936 (EA); 41 mi. from Daressalaam on 
main road to Morogoro, climbing over trees and shrubs in hillside 
thicket margins; also trailing plant on roadsides; climber, 10-12 
ft.; stem covered with greyish white hairs, sage green, paler on 
backs; corolla tube cream, lobes pale blue, very common, J.R . 
Welch 303, July 4, 1955 (K). 



162 PHYTOLOGIA Vol. 17, no. 3 

6. Bonamia cordata fHall. f . ) Hall. f. Bull. Horh. Boiss. 7:4.'i 
1899. 

Prpvostea cordata Hall. f. Bot. Jahrb. 18:93. 1894. 
Not Brfworia cordata Blumo. Bydr. Fl. Nederl. Ind. 722. 1825. 

Ppmnnial twining vines. Stents woody, terete, twining, long, 
finely villous, more densely villous when young; intemodes 6-10 
cm. long. Leaves petiolate, soft, herbaceous, thin or sometimes 
submembranous , moderately or densely villous or scabrous on the 
upper surface, densely villous of ferrugineous on the lower sur- 
face; petioles 1.5-3 cm. long, villous; blades ovate-cordate to 
ovate-acuminate, 4.5-7.5 cm. long, 3-5 cm. wide, cordate at the 
base, acuminate or acute at the apex; midrib impressed above, 
prominent beneath, with 6-10 pairs of lateral veins. Inflores- 
cences axillary, pedunculate C3mies of few to several flowers; pe- 
duncles long, usually 3-8 cm., 1.5-2 mm. thick, rigid, densely 
villous or tomentose; secondary peduncles 4-6 mm. or rarely longer; 
pedicels short or nearly absent; bracts foliaceous, petiolate, 
ovate-lanceolate, 1-2 cm. long, 6-10 mm. wide, indumentum as on 
leaves; bracteoles lanceolate or ovate-lanceolate, 5-8 mm. long, 
sepals coriaceous or soft-coriaceous, unequal; outer sepals orbicular- 
mucronate or ovate-orbicular, 11-14 mm. in diameter, villous or 
ferrugineous outside, moderately or densely ferrugineous inside, 
abruptly acuminate or obtuse-acuminate at the apex; inner sepals 
small, ovate or ovate-lanceolate, 7-9 tran. long, 4-6 mm. wide, 
densely ferrugineous or villous outside, glabrous inside, acute 
at the apex. Corolla and stamens not known. Ovary globose or 
subglobose, sparsely shoii:-pilose or glabrous; styles bifid to the 
middle or higher, glabrous; stigmas small, globose. Fruits and 
seeds not known. 

Type: Madagascar (Cote orientale) , Boivin 2184 , 1846-1852 
(G-lectotype) . 

Endemic to east coast of Madagascar, and known only by type 
collection, which is incomplete, lacking corolla, stamens, fruits 
and seeds. 

Although the only available material is the type which is frag- 
mentary, this is a very distinct species and can easily be recog- 
nized by its unequal sepals, outer enlarged sepals being pubescent 
on both surfaces, foliaceous bracts and large cordate leaves. It 
superficially resembles B. semidigyna , to which it seems to be more 
closely related than to any other species, because of similarity 
in their indumentum, large cordate leaves, long pedunculate cymes 
and long intemodes. It differs from B. semidigyna in its unequal 
sepals and foliaceous bracts. 

Hallier (1894) hesitantly described this species under the 
genus Prevostea, presumably because of its unequal sepals. He later 
transferred it to Bonamia because the sepals are merely unequal and 
not accrescent as in the genus Calycobolus ( =Prevostea ). 



1968 Uyint & Ward, Revision of Bonamia I63 

7. Bonamia semidigyna (Roxb.) Hall. f. Bot. Jahrb. 16:528. 1893. 

Convolvulous semidigynus Roxb. Fl. Ind. (ed. Carey et Wall.) 

2:47. 1824. 
Breweria cordata Blume, Byrd. Fl. Nederl. Ind. 722. 1825; 

not Bonamia cordata (Hall. f. ) Hall, f . Bull. Herb. 

Boiss. 7:43. 1899. 
Breweria roxburghii Choisy, Mem. Sec. Phys. Geneve 6:493. 

1833. 
Breweria madagascariensis Choisy, 1. c. 1933; not Bonamia 

madagascariensis Poir, Encycl. Meth. Bot. Suppl. 1:677. 

1810. 

Perennial, woody vines. Stems twining to a height of 15 m. , 
terete, densely brown or reddish brown tomentose. Leaves petiolate, 
subcoriaceous , soft, leathery or membranous, densely or sparsely 
tomentose underneath; petioles 18-35 (-60) mm. long, tomentose like 
stems, canaliculate above; blades broadly to narrowly ovate, 6.5- 
15 cm. long, 4-10 cm. wide, broadly cordate or occasionally truncate 
at the base, shortly acuminate or cuspidate at the apex; veins im- 
pressed above, prominent underneath; lateral veins 5-7 pairs. In- 
florescences axillary, pedunculate, umbelliform cymes of 2-5 (-7) 
flowers , rarely solitary by abortion of lateral flowers ; peduncles 
long, variable in length, mostly 2-12 cm. long, 1.5-2.5 mm. thick, 
tomentose as the stems; pedicels variable in length, 4-15 mm. long, 
1.5-2 mm. thick, densely tomentose; bracts linear or lanceolate, 
mostly small, rarely foliaceous , 5-10 mm. long, rarely 20 mm. long. 
Sepals ovate, or ovate-oblong, or ovate-acuminate, equal, subequal 
or the inner slightly shorter, 7-14 mm. long, densely tomentose, 
acuminate or acute at the apex. Corolla white, campanulate to 
funnel-shaped, 3-5 cm. long, lobulate or subentire at the margin, 
long-pilose on interplicae, glabrous on plicae. Stamens included; 
filaments glabrous above, sparsely pilose near the base; anthers 
oblong, 3-4 mm. long, cordate at the base. Ovary ovoid, with 
dense long hairs; styles bifid to the middle or lower, glabrous 
above with scattered hairs near the base; stigmas globose-peltate. 
Capsules broad-ovoid to subglobose, apiculate, hairy at the apex, 
about 10-14 mm. in diameter, 2 celled, 4- to 8-valved , 4-seeded, 
rarely less seeded by abortion; seeds glabrous, black, oval in 
outline, convex on one side, and plane on two other sides, 5-6 
mm. long. Cotyledons ovate, broadly cordate at the base, folded; 
cotyledonary petioles fused. 

Type: India; type specimen not available. 

From Madagascar through Ceylon, India, Pakistan, Burma, Thai- 
land, South Vietnam, Malaya, Borneo, Sumatra, Java, Celebes, Culion 
and Luzon to Moluccas and New Guinea (Map 3). 

This is the only species of the genus so widely distributed, 
commonly collected and represented in most large herbaria of the 
world. Hallier recognized three varieties under this species 
mainly on account of differences in indumentum. However, Van 
Ooststroom (1954) skeptically treated var. ambigua , stating, 



16U 



PIIYTOLOGIA 



Vol. 17, no. 3 




1968 Ityint & Ward, Revision of Bonamia 165 

"It is... more difficult to draw a satisfactory line between var. 
semidigyna and this variety, than with var. farinacea . It is not 
impossible that Hallicr is right that we have here a hybrid before 
us." Specimens (bearing the varietal epithet ambigua and annotated 
by Hallier) are somewhat variable in several features, but pre- 
sumably all were collected by Hallier from a single plant in the 
Botanical Garden of Bogor. It had best be retained as an aberrant 
form of var. semidigyna from which it differs mainly by its very 
large leaves. 

Key to Varieties 

1. Stems and lower leaves with a dense brown or reddish brown to- 
mentum; finer nervations of leaves indistinct or rarely trans- 
verse veins between adjacent lateral veins (or intercostal 

veins) barely visible; outer sepals acute to acuminate 

var. semidigyna . 

1. Stems and lower leaves sparsely tomentose or covered with 
short, closely appressed paler hairs; finer nervations of 
the leaves often more visible by the absence of a dense hair 
coating; outer sepals acute var. farinacea . 

7a. B. semidigyna (Roxb. ) Hall, f . var. semidigyna . 

B. semidigyna (Roxb.) Hall. f. var. ambigua Hall. f. Bull. 
Herb. Boiss. 5:817. 1897. 

Stems tomentose with brown or reddish brown hairs. Leaves 
densely tomentose underneath, with barely visible finer nerves (be- 
cause of dense coating of hairs). Sepals 10-14 mm. long, rarely 
slightly shorter, densely brown tomentose, acute or acuminate at 
the apex, thick, rarely with slightly visible nerves. 

Edges of secondary forests, thickets, hedges, waysides and 
river banks, from sea-level to about 250 m. , rarely to higher 
altitudes. 

B. semidigyna var. ambigua of Hallier is treated here as a form 
of the typical variety. 

Specimens examined: 

ANDAMAN ISLANDS: S. Andaman, Port Monat-hill jungle. King's 
Collector, 19. 12. 1891 (US); Prain's collector 21 , 8. 3. 1901 
(BM, G); Prain's collector 95 , March 1901 (G); (doubtful) "Tenas- 
serim and Andamans," Heifer 5874 (W). 

BURMA: Maulmein, Sammlugen 0. Kuntze's Weltreise 6289 , X. 75 
(NY); Sandoway, Arracan, Capt. Margrave (L) ; Tenasserim, J.D.V. 
Packman (BM); Pegu, Scott (L) ; without loc. J.H.B.C. 1405 (L). 

CEYLON: James Macrae 533 (BM); H.K. Thwaites 2853 , 1855 (BM, 
G, GH, W); Col. Walker (G). 

COCHINCHINA (SOUTH VIETNAM): M. Germain 78 , 1879 (G), Recule, 
1 Avril 1880 (F, L, UC) ; fl. albi. Hab. ad Um dzan mot in Oust. 
Coch. L. Pierre, 1. 1864 (BM); ad Um dzan mot. in Ouest. Coch. fl. 



166 P H Y T L G I A Vol. 17, no. 3 

albi. , L. Piorro 22 , 1. 1867 (A ,r,G,GH .NfY) ; ad Bien hoa in Oufst. 
Coch. fl. albi. , Pierro , 12. 1869 (F. L. UC) [many of Pierre's 
label notes not deciphered]; M. le Dr. Thorel 612 , 1862-1866 (^). 

INDIA: Travancore, Madaras, J.S. Gamble 14778 , Sept. 1884 
(HBG, K) ; Prope Mercara, Terr. Canara, R.F. Hohenacker 563 , Jan., 
Febr. , M. 1847 (BM, G) ; Tidal Creek, Naiti, N. Kanara Dist. , 
W.A. Talbot 2868 , 10. ]. 93 (G); "India," Wallich 1405 . 1, 1405. 
2, 1405. 3 (RM). 

INDONESIA: JAVA: Batavia, Heurdterrein z. van D.jassinga, 
250 m. , Backer 26030 (L); Blume 1851 (L); Buitenzorg, Boerlage (L); 
Java Res. Batavia Barengkok. W. v. Leuviliang, alt. 250 m. , Bak - 
huizen von den Brink Jr. 770 , 16. 6. 1921 (L, W) ; corolla lactea, 
cult, in Hort. Bog., Hallier C. 18. a., 24. V. 1893 (G, L) , C. 18. 
b. , 5. V. 1893 (L), C. 18. c. , 13 V. 1895 (L) ; Hallier 104d , ]4. 
8. 1896 (L); Cult, in Hort. Bog., Hallier (L) ; Korthals 226 (L); 
Kandang Japi , Korthals (L) ; Zollingero 1339 , 1844 CG). Location 
indefinite. Blume (L); Reinwardt 362 CL). Unknown collector: 
"Java" (1). SUMATRA: Korthals 48 (L), 1711 (L); Pasier Cantang, 
lat. 2°S, sea level, fl. white, H.C. Robinson and C.B. Kloss 2 , 
18. VI. 1914 (BM). SUNDA: Straights of Sunda , Macartney and 
Staunton (BM). 

MADAGASCAR: Envir. Tamatave, C.D'Alleizette , Nov. 1906 (L); 
central Madagascar, Rev. R. Baron 2773 , Dec. 1883 (BM); M. Goudot 
222 (?), 1833 (G); Hab. ad Tamatave, flores albi, Helsemberg (BM); 
Hunblet 211 (W) ; Tamatave, D. Paulay , June 1887 (W) . 

MALAYA: Kuband Ulu, Province Wellesley, C. Curtis , July 1890 
(BM); Pulo-Pinang, A. Delessert 632 , 1835 (G) ; Selangor, C.W. 
Franck 1013 , 16. 9. 1937 (A); Perlis , Kangas , alt. low, M.R. Hen- 
derson 22858 , Nov. 16, 1829 (.BM, BRI) ; "2" specim. lect. in Ins. 
Penang, G. Porter , in 1822 (NY); Penang, unknown collector, with 
Wallich Herbarium No. 1405.2, 1832 (G). 

PAKISTAN: Chittagong, Regio tropl. alt. 1000 ped. J.D. Hooker 
and T. Thomson , 1861 (BM, G, GH, L, W) ; The Chittagong Hill Tracts, 
Dr. King's collector 206, 1885 (K) , 615, 1887 (L). 

PHILLIPPINES: Central Luzon, A. Loher 4155 (US); Culion Island, 
E.D. Merrill 538 , Dec. 18, 1902 (NY, US),_618, Jan. 1, 1903 (NY, 
US). 

SARAWAK: Baram Mouth, Baram Dist., C. Hose 27 , Dec. 1894 (BM). 

THAILAND: Kao Saming (Krat) , under 50 m. , A. E.G. Kerr 9399 , 
25. 11. 1924 (A, L); Hat Yai (near Songkhla) , under 50 m. , Kerr 
13535 , 22. 12. 1927 (A, L) . 

Locations not determined: Anamallays , R.H. Beddome 5627 (BM); 
Sillet (Indes Or.), Wallich 1405 . 1, 1832 (G) ; Peninsula Indiae 
Orientalis, Wight 1999 (GH, L, NY). Locations unknown: Wallich 
1405, 1832 (G); Wallich 1405.2 (L). Collector unknown: (G). 

7aO;. B. semidigyna var. semidigyna forma ajjibigua (Hall, f . ) Myint 
and Ward, comb. nov. 

B. semidigyna (Roxb.) Hall. f. var. ambigua Hall. f. Bull. 
Herb. Boiss. 5:817. 1897. 



1963 Ifyint & Ward, Revision of Bonami a 16? 

Differs from the typical form by its greyish and thinner indu- 
mentum, larger and wider leaves with broadly rotund -cordate bases, 
longer petioles, larger bracts and bracteoles , and larger corolla. 

Type: Bangka (culta in horto Bogor.)j Hallier C. 17. a . , 23. 
V. 1893 (L-lectotype.'). 

Known only from type location. 

All specimens deposited at the Rijksherbarium were collected 
from a single plant grown in the Botanical Garden at Bogor, ac- 
cording to Van Ooststroom (1954). Since no further collection has 
been made, Van Ooststroom questioned it as a distinct variety. 
IVhen it was described, Hallier suggested it as a hybrid between 
var. semidigyna and var. farinacea, and Van Ooststroom remarked 
that Hallier may well be right. Future collections are much de- 
sired. 

Specimens examined : 

INDONESIA: Bangka, culta in horto Bogor., H. Hallier C. 17. a. , 

23. V. 1893 (L); C. 17. b . , 7. IV. 1893 (L) ; C. 17. c. , 29. 111. 

1893 (L); C. 17. d . , 5. V. 1893 (L) ; Hallier X. F. 75 (L). 

7b. B. semidigyna (Roxb.) Hall. f. var. farinacea Hall, f. Versl. 's 
Lands Pl.-tuin Btzg. 125. 1895 (1896). 
Letts omia bancana Miq. Fl. Ind. Bat. Suppl. 561. 1861. 

Differs from var. semidigyna in stems and lower leaves covered 
with short, closely appressed hairs of a paler color, grey or light 
brown; finer nerves of leaves distinctly visible (because of thin 
coating of soft hairs); sepals 7-12 mm. long, sparsely or moderately 
tomentose (not densely tomentose except in young buds), mostly a- 
cute or shortly acuminate, frequently with distinct nerves. 

Type: Celebes, cult, in horto Bogor., H. Hallier C. 16. a. , 

23. V. 1893 (L-lectotype.'). 

Thickets on beaches and rocks , both in marshy and dry locali- 
ties, from sea-level to 75 m. , in Malaysia from Banka and Celebes 
to Moluccas (Ceram) and New Guinea. Van Ooststroom notes that Ceram 
and Celebes specimens possess aberrant characters (longer pedicels 
and elliptic-obtuse sepals) which with future collections might 
prove to be taxonomically important. 

Specimens examined : 

CELEBES: Sudwest Celebes, Bau-Bau, Gestrupp, 0-75 m. , sehr 
trocken, korallenkalk, J. Elbert 2641 , Sept. 5, 1909 (L) ; Padan- 
goma, 0-10 ra. , Mangrovenwald , Strandbusch, Sumpf, lehmig, Elbert 
3250 , Oktober, 1909 (L) ; Cult, in horto Bogor., H. Hallier C. 16 . 
a7^3. V. 1893 (L), C. 16. b , 17. II. 1893 (L) , C. 16. c , 13. V. 
1895 (L) , C. 18. c , 13. V. 1895 (G) ; Provincia Minahassa, S.H. 
Koorders 16559B (L) ; Bonto Parang, Rachmat 4 (exp. van Vuuren) , 

24. 6. 1913 (L). 



168 P H Y T L G I A Vol. 17, no. 3 

Si;i{AM fCLKAM): l'o<aoo Tikof^s, + m. , Kornassi 1274 Cexp. 
Rutton), 10. 5. 1918 (L). 

NEW GUINCA: Papua: Lower Fly Kiver, east bank, opposite 
Sturt Island, robust climber in second growth, rain forest, flower 
white, L.J. Brass 8180 , Oct. 1936 (A). 

8. Bonamia elegans CWall. ) Hall. f. Bot. Jahrb. 16:529. 1893. 
Convolvulus elogans Wall. Cata. p. 38, no. 1392. 1828. 
Brcworia elegans (Wall. ) Choisy, Mem. Soc. Geneve 6:193. 

1833. 
Brewer iops is elegans (Wall.) Roberty, Candollea 14:31. 1952. 

Perennial, woody vines. Stems terete, twining, pilose or be- 
coming glabrous in age, thin and wiry or becoming 2-3 mm. thick. 
Leaves shortly petiolate, subcoriaceous or soft-coriaceous, thinly 
pilose above, densely strigose beneath; petioles 2-8 mm. long, den- 
sely sericeous or pilose; blades variable in shape and size, older 
being ovate-elliptic; younger leaves lanceolate or sublinear, 2-4 
cm. long, rounded or slightly cordate at the base, obtuse or 
obtuse-mucronate at the apex; veins impressed above, prominent 
beneath; lateral veins about 3-4 pairs. Flowers axillary, soli- 
tary or rarely in shortly pedunculate cymes of 2-3 flowers ; pe- 
duncles 5-18 mm. long, mostly 1 mm. thick, finely pubescent; 
pedicels 2-5 mm. long, frequently slightly thicker than peduncles, 
brown-pubescent; bracts two, opposite, linear or linear-lanceolate, 
3-6 mm. long. Sepals coriaceous or subcoriaceous, ovate-lanceolate, 
12-15 mm. long, 4-5 mm. wide, acuminate or acute at the apex, equal 
or slightly unequal, finely pubescent, more densely so near the 
base. Corolla blue, campanulate-infundibuliform, 4-5 cm. long, 
slightly lobulate or subentire, pilose on interplicae, glabrous 
on plicae; tube cylindrical, wide, not distinct from the limb. 
Stamens included; filaments glabrous above, with scattered hairs 
below, at least along the edges; anthers oblong or oblong-lanceo- 
late, 3-4 mm. long, cordate at the base. Ovary conical, with 
long hairs at the apex, glabrous near the base; styles fused near 
to the stigma, with scattered hairs on the lower part; short sty- 
lar branches unequal; stigmas depressed -capitate. Fruits not 
known. Cotyledons oval, not folded in young stage, unknown in 
mature stage; cotyledonary petioles fused. 

Type: Burma: Prome, Wallich (G-lectotypeJ , BM-isotypel ) 

Known only from the type locality (Map 3). 

This species is poorly known and rarely collected. It is rep- 
sented in a few large herbaria of Europe only by an old collection 
made by the author of the species more than a century ago. Ap- 
parently no further collections have been made since that time. 
The type collection is fragmentary, since it is only of a flower- 
ing branch and is without leaves of the vegetative parts. Ac- 
cording to Clarke (1885), the juvenile leaves are much larger, 
attaining 7.5 by 3.1 cm, whereas the leaves on flowering branches 
are 3.8 by 0.8 cm. Since the vegetative branches and fruiting 



1968 Myint &: Ward, Revision of Bonania 16? 

material are not available in the present study, a more complete 
description of the species has to await future collections. 

Roberty (1952) , in erecting his new genus Breweriopsis typi- 
fied by this species, lumped B. elegans , B. grand i flora and B. 
minor as constituting a single species. These three species of 
different continents differ in several important features. The 
most obvious common feature is their blue flowers , by which Rob- 
erty characterized his new genus. His treatment is quite artifi- 
cial in many respects. He excluded several species possessing 
blue flowers, two of which were treated under an entirely dis- 
tinct genus , Stylisma . But perhaps the most unacceptable part 
of his classification was his treating B. spectabilis and B. minor 
under two different genera, whereas these two, as pointed out 
by Verdcourt, are so similar in all features that they are con- 
specific. 

9. Bonamia dietrichiana Hall. f. Bull. Herb. Boiss. 5:1012. 1897. 
Bonamia pannosa sensu Hall. f. 1. c. 5:810, as to the 
description and quoted specimen, not Brcweria pannosa 
R. Br. Prodr. 488. 1810. 

Perennial twining vines. Stems slender, terete, becoming woody, 
mostly 1-2 m. long, 1-2 mm. thick, fulvous -tomentose or subseri- 
ceous; internodes variable in length, 1.5-5 cm. long. Leaves shortly 
petiolate, soft and thick or slightly leathery, sericeous and ap- 
parently dark green above, densely sericeous and pale green beneath; 
petioles 4-8 mm. long, sometimes slightly shorter, sericeous; blades 
ovate or ovate-subcordate, 2.5-4 cm. long, 1.7-2.7 cm. broad (smal- 
ler on upper leaves), subcordate or truncate at the base, obtuse- 
mucronate or emarginate-mucronate at the apex; mucro 1 mm. long; 
nerves indistinct above, prominent beneath; lateral nerves 5-7 
pairs. Flowers axillary, solitary or in simple cymes of 2-3, 
shortly pedunculate; peduncles 3-7 mm. long, terete, slender, ser- 
iceous; pedicels 2-4 mm. long; bracts opposite, ovate, shortly pet- 
iolate, 7-10 mm. long, 4-6 mm. broad, finely tomentose or sericeous. 
Sepals lanceolate or ovate-lanceolate, 12-13 mm. long, subcoria- 
ceous or coriaceous; outer sepals densely villous or sericeous out- 
side, acuminate at the apex, slightly longer than inner ones; inner 
sepals sparsely pubescent or nearly glabrous outside, acute or 
shortly acuminate at the apex. Corolla white (?), funnel-shaped, 
3.5-4.5 cm. long, 3-3.5 m. broad, subentire or slightly lobulate 
at the limb, long-pilose on interplicae. Stamens inserted; fila- 
form, glabrous above, pilose near the base; anthers linear-oblong, 
2-3.5 mm. long, cordate at the base. Ovary ovoid, long-pilose at 
the apex, glabrous below; styles bifid to the middle or nearly to 
the base, filiform, glabrous or with scattered hairs near the base; 
stigmas large, globose-capitate. Fruits and seeds not known. 

Type: Queensland, Australia, A. Dietrich 19 (HBG) . 

This species, endemic to Queensland, Australia (Map 3), is dis- 
tinct from all other Australian species in its longer and lanceo- 



170 P H Y T L G I A Vol. 17, no. 3 

late sepals, larger corolla and foliaceous bracts. It is some- 
what related to B. pannosa because of similarity in its indumentum, 
slightly unequal sepals and larger subcordate leaves. It is pro- 
bably more closely related to B. elegans of the Orient because of 
similarity in their sepals and corolla. 

Hallier, in his earlier treatment, included this species under 
B. pannosa and later realized its distinction from the latter. It 
differs from B. pannosa in its larger and white corolla, nearly 
equal, closely appressed and lanceolate sepals, outer sepals being 
glabrous inside, and leaves being obtuse-mucronate or emarginate- 
mucronate at the apex (Figure 5). 

Specimens examined: 

AUSTRALIA: Queensland: A. Dietrich 19 (HBG-lectotype and iso- 
type) ; Stony Creek, Stuart, Townsville, " K. K." 8 , April 4, 1954 
(BRI). 

10. Bonamia menziesii Gray, Proc. Am. Acad. 5:336. 1862. 

Breweria menziesii (Gray) Bentham and Hooker, Gen. PI. 

2:877. 1876. 
Bonamia Herminieri Hall. f. Bot. Jahrb. 16:529. 1893. 
Peris permum albiflorum 0. Degener, Fl. Hawaiiensis, Fam. 

307. 1932. Type: Degener, Park and Nitta 4111 .' 
Perispermum menziesii (Gray) 0. Degener, Fl. Hawaiiensis, 

K6. 1934. 

Perennial, woody, slow-growing, coarse vines, up to 10 m. 
Stems twining, terete, long, without milky juice, glabrescent and 
with pale yellowish bark, fulvous-tomentose when young, distantly 
leafy, soon becoming woody, later bearing short leafy spurs. Leaves 
petiolate, soft-coriaceous, tomentulose and hoary to glabrous above, 
densely fulvous-tomentose below; petioles 8-25 mm. long, sulcate, 
fulvous-tomentose or becoming less tomentose in age; blades oblong- 
elliptic, ovate or rarely orbicular, 3.2-9 cm. long, 1.5-3.5 cm. 
broad, rounded at the base; truncate, emarginate, obtuse or acute 
at the apex. Flowers axillary, solitary, rarely in cymes of two 
to few flowers; peduncles short, 2-5 mm. long, demarcated from 
pedicels by two inconspicuous bracts, fulvous-tomentose, mostly 
thicker than pedicels; pedicels longer than peduncles, commonly 
1.2-2 cm. long; floral buds mostly erect. Sepals ovate, densely 
fulvous-tomentose outside, glabrous inside, soft -coriaceous , per- 
sistent and brittle in the fruit, subequal; the two exterior about 
10 mm. long, 8 mm. wide; the three interior mostly 7 mm. long. 
7 mm. wide, thinner, less densely fulvous-tomentose and less acute 
at the apex. Corolla white, yellowish brown or greenish, funnel- 
shaped with narrow spreading limb of 5 subtruncate lobes (or lo- 
bules), 22 mm. long, 16 mm. wide; outside glabrous on plicae, hir- 
sute with pale tawny silky hairs on interplicae. Stamens inserted; 
filaments slightly lower than the corolla, filiform, stiff, adnate 
to corolla for about 6 mm.; anthers white, oblong, about 3-3.5 mm. 
long. Ovary with narrow dixc, ovoid-conical; styles connate for 
about 2 ram. near the base, about 15 mm. long; stigmas rugose-capitate, 
1 mm. wide. Capsules pendent on stiff pedicels, hardly dehiscent. 



1968 



Ityint & V/'ard, Revision of Bonamia 



171 




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172 PHYTOLOGI/l Vol. 17, no. 3 

glabrous with chartaccous wall, about 10-15 mm. long and 8-10 mm. 
wide, ovoid-conical, straw colored, 4- to 2- (-1) seeded, with 
very thin and soft septum; persistent sepals spreading and brittle. 
Seeds covered with black perisperm, glabrous, about 6-8 mm. long, 
and almost 5 mm. wide, ovoid-angular-convex, with yellov/ish brown 
or crimson testa. Cotyledons corrugate and folded, bilobed, cor- 
date at the base, with fused cotyledonary petiole. 

Type: lies Sandwich— Maui , M.J.Remy 420 , 1851-1855 CGH). 

On rocky slopes and valleys on Hawaii, Lanai , Maui, Molokai 
and Oahu Islands (Map 4) . 

Hallier (1893) gave the name B. herminieri , in honor of the 
collector, Herminier, to a specimen presumably from Guadeloupe, 
West Indies. Since this range dis.juction has not been supported 
by further collections, it seems reasonable to suppose the orig- 
inal label in error. The type specimen has not been examined, and, 
in fact, cannot now be located at the herbarium of Boissier, Geneva, 
where it was presumably deposited, but the name is placed in syn- 
onymy under B. menzicsii on the authority of Hallier (1897) who, 
on further study, considered B. herminieri to be no more than a 
somewhat aberrant form of B. menziesii. 

0. Degener (1932) erected a new genus for this species because 
of its hardfy dehiscent capsules, perispermous seeds and supposed 
lack of septa. Since the fruits (capsules) remain closed for a 
long period after ripeness, he assumed them as completely inde- 
hiscent, which does not seem to be true. Further, he missed the 
fact that the presence of perisperm on the seeds is common through- 
out the genus Bonamia . Degener definitely overlooked the presence 
of thin septa, thus characterizing his new genus with nonseptate 
capsules. He also proposed a new species, which he treated under 
this genus. The specimens collected on the island of Hawaii differ 
appreciably from the specimens collected on Lanai, Maui, Molokai 
and Oahu, and thus are treated here as belonging to a distinct 
variety (Figure 2). 

Specimens examined: 

LANAI: Dry forests, west end, C.N. Forbes 152. L , June 1913 
(A, F, MO, NY, UC, US); W. Hillobrand , 1874 (GH); Hillebrand , 1890 
(BM); A.S. Hitchcock 14712, Sept. 22, 1916 (US); Paomaio, G.C. 
Munro , 3. 19. 1914 (BM), 4. 18. 1914 (NY, UC, US). 

MAUI: Pakiloi, Forbes 2067 M , Mar. 23, 1920 (NY, UC , US); 
Komauu, Forbes 2067 MO , Mar. 23, 1920 (UC) ; M.T. Remy 420 , 1851- 
1855 (GH, lectotype), 421 (L). 

MOLOKAI: Kalapamoa, Forbes 430 MO , Aug. 1912 (MO, UC, US); 
west end, Kokio Gulch, J.F. Rock 14015 , May 21, 1918 (NY, UC , US); 
west end. Rock , Feb. 1920. 

OAHU: Small, arid, rocky gully two-thirds of a mile from the 
sea on the south slope of Keaau Valley, 0. Degener, K.K. Park and 
Y. Nitta 4111, Feb. 7, 1932 (MO); middle ridge of Niu Valley, on 



1968 



l^rint & Ward, Revision of Bonamia 



173 



Figure 2 
Leaf shapes and sizes in the varieties of Bonamia menziesii 




var. menziesii 



17li P H Y T L G I A Vol. 1?, ao. 3 

partly woodod, sunny slope, 50 ft. abovo stream, IX'^/tncr , \'ark 
and Nitta 5975 , Juno 4, 1932 (MO); small east-central ridf^e near 
head of Wailupe Valley, over bushes and low trees at 1500 ft., 
DoRener 21186 and W. Hatheway , Dec. 19, 1950 CBM, MO, UC) ; south- 
west side of Poamoho Gulch; south-west side of Brodie Camp, on 
rocky, grassy, sparingly shrubby, precipitous slope at 1500 ft. 
elevation, Degener 21257 and Hatheway , Jan. 25, 1951 (L'C, US); 
Kaala Mountains, H. Mann and W.T. Brigham 618 (BM, F, G, GH , MO, 
NY, US); U.S. Exploring Expedition under the command of Captain 
Wilkes (US). Unknown location: Hillebrand 1889 CUS). 

Bonamia menziesii var. rockii Myint and Ward, var. nov. 

Differt a varietate typica foliis orbicularibus , orbiculari- 
ovatis , vel ovatis , 2.5-4. cm. longis , l-1.2plo longioribus quam 
latioribus, raro l.Splo, apice emarginato, truncato, obtuso vel 
raro abrupte acuto. 

Differs from the typical variety in possessing orbicular, 
orbicular-ovate or ovate leaves, 2.5-4 cm. long, with a length- 
width ratio of 1-1.2, rarely 1.5, emarginate, truncate, obtuse 
or rarely abruptly acute at the apex. 

Type: Kona: Puu Waawaa, J.F. Rock , March, 1912 (GH). 

All three specimens cited here were collected by J.F. Rock 
between Kona and Puu Waawaa at varying dates ; this variety is 
named in his honor. 

Specimens examined : 

HAWAII: Kona: Lava beds between Huehue and Puu Waawaa, J.F . 
Rock 3541 , June 4, 1909 (GH) ; Puu Waawaa, Rock , March, 1912 (GH) ; 
Puu Waawaa, Kamanomano, Rock (NY). 

11. Bonamia grand i flora (Gray) Hall. f. Bull. Herb. Boiss. 5:810. 
1897. 
Breweria grand i flora A. Gray, Proc. Am. Acad. 15:49. 1880. 

Perennial, trailing, herbaceous or suffrutescent pseudoliana, 
growing annually from lower nodes of previous shoots or from 
slightly enlarged roots. Stems terete, rarely ridged or subterete, 
glabrous to finely puberulous , 2-4 mm. thick, becoming 3-5 m. long, 
with frequent branching. Leaves sessile or subsessile, subcoria- 
ceous to membranous, glabrous or finely puberulous and glabrescent; 
petioles almost absent or 1-3 mm. long, and curved; blades ovate, 
2.2-3 cm. long, 2-2.5 cm. broad, rounded or slightly cordate at 
the base; obtuse, acute or rarely retuse at the apex; lateral veins 
mostly 4-7 pairs. Flowers axillary, solitary; tA'^o lateral abor- 
tive buds sometimes present in the axils of the bracts; peduncles 
1-4 cm. long, sometimes becoming longer in age, grey pubescent or 
puberulous; pedicels short, slightly thicker than peduncles, dense- 
ly pubescent while young; bracts small, linear or scale-like, 1-2 



1968 Ifylnt & Ward, Revision of Bonamia 175 

mm. long. Sepals broadly lanceolate or oblong-ovate-lanceolate, 
acute or acuminate at the apex, equal, or unequal, outer being 
slightly shorter, 1.5-2.6 (2.8) cm. long, 4-10 mm. wide, membran- 
ous or subcoriaceous , outside finely puberulous with grey or 
silvery hairs. Corolla deep blue or purplish blue, lighter to- 
wards tubular base, tubular-campanulate or funnel-form, 7-8.5 cm. 
long, 5-7 cm. wide, shallowly lobulate, silky pilose with long 
hairs on interplicae, glabrous on plicae. Stamens included; fila- 
ments epipetalous , shorter than styles , as high as half the length 
of the corolla, slightly unequal, glabrous above, villous below; 
anthers 4-5 mm. long, oblong-lanceolate, cordate at the base, 
introrse by longitudinal slits. Ovary conical, glabrous with four 
vertical ridges, with a circular disc near the base; styles inser- 
ted, terminal, mostly 5 cm. long or sometimes longer, bifid above 
the middle (rarely trifid , then ovary trilocular and six-ovulate) ; 
stigmas globose-peltate. Capsules conical, apiculate, 4- or 8- 
valved, rarely 6-valved, 4-seeded, rarely 6-seeded. Seeds oval, 
brownish, glabrous, rarely with scattered hairs on dorsal sides. 
Cotyledons oval or oboval with emarginate apices, folded against 
the radicle, with free cotyledonary petioles. 

Type: Manatee and Sarasota, Florida, A. P. Garber, June, 1878 
(GH-Lectotype; F, FLAS , MO, PH, US - Isotypes.'). 

Dry, deep sandy areas in scrubs or edge of scrubs, more com- 
monly in open ground and disturbed areas, occasionally on ancient 
sand dunes , ranging from south to central Florida (Map 5) . 

Bonamia grand i flora is geographically completely isolated from 
related species of Mexico and Central America, and it is the only 
species entirely restricted to the continental United States. It 
has been included in B. elegans of the Orient by Roberty (1952) 
who treated it under his new genus Breweriopsis , which he charac- 
terized by the large corolla. However, B. grand i flora shows sev- 
eral distinct features from B. elegans , particularly leaf shape 
and size, fusion of the stylar branches and stigmas. In B. grandi - 
flora the leaves are orbicular or ovate, 2.2-3 cm. long and 2.2-5 
cm. wide, stylar branches are connate only for the lower half, and 
the stigmas are small and globose, whereas in B. elegans , the leaves 
are oblong or oblong-ovate, 3-4.5 cm. long and 8-15 mm. wide (ex- 
cept the lower ones which are slightly wider), the stylar branches 
are connate for three-fourths of the total length or higher, and 
the stigmas are larger and depressed-capitate. 

The derivation of the specific name is quite appropriate for 
its large, conspicuous and beautiful flowers, purplish blue in 
color. 

Specimens examined : 

FLORIDA: Highland County: Open, dry, sandy slope among the 
■'inland sand dunes" near Sebring, D.S. Correll and J.B. McFarlin 
6227 , August 3, 1936 (DUKE); Scrub, south end of Lake Jackson, 



176 



PHYTOLOGIA 



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1968 Myint & Ward, Revision of Bonamia 177 

Sebring, Ray Garrett , April 21, 1948 (GA) ; sand dunes, Sebring, 
F.W. Hunnewell 1049 , May 15, 1927 (GH) ; sand hills, Sebring, 
J.B. McFarlin 9414 , September 6, 1934 (FLAS), .5701, 6. 9. 1931 
(MICH); sandy scrub. Lake Placid, F.H. Sargent 7180 , May 23, 
1955 (SMU); scrub, Sebring, J.K. Small and E. West , September 
5, 1934 (FLAS). Lake County: In scrub near Mt. Dora, J.J . 
Fennel 463 , July 18, 1937 (UC); in vicinity of Eustis, A.S. 
Hitchcock , June-July, 1894 (F, FLAS, MO); in vicinity of Eustis, 
G.y. Nash 1326 , July 16-31, 1894 (F, G, GH , MICH, MO, PH, UC, 
US) ; near Lake Dora, Tavares , P.H. Rolfs 511 , June 29, 1893 
(F, FLAS, MO). Manatee County: Sand ridge, Manatee River, 
Bradenton, A. Cuthbcrt 1358 , June 23, 1916 (FLAS); A. P. Garber , 
June, 1878 (F, FLAS, GH , MO, PH , US); J.H. Simpson, 1889 (US); 
Palma Sola, S.M. Tracy 6431 , September 10, 1889 (NCU). Marion 
County: Sandy roadsides, Nat. Forest, A.V. Cleet , August 2, 
1937 (NSU) ; near observation tower on highway, Ocala National 
Forest, Bailey and Hume , August 19, 1935 (FLAS); in a scrub, 
Ocala National Forest, Hugh O'Neill , September 12, 1929 (FLAS, US); 
frequent; 1 mi. east and 4.8 mi. south of Central Tower, Ocala 
National Forest, D.B. Ward and T. Myint 2126 , July 28, 1960, 
(FLAS, FSU). Orange County: In sand scrub, Orlando, Hugh O'Neill 
8^, August, 1924 (US); sandy scrub near Windemere, P.O. Scallert 
20849, October 4, 1947 (SMU); sand hill among scrub oak, Orlando, 
E. West , May 24, 1929 (FLAS); Clarocona, C.S. Williamson , July, 
1895 (PH). Osceola County: Sandy soil, near swamp, Tampa high- 
way, Mary L. Singletary 370 , May 24, 1938 (DUKE, NSC). Polk 
County: Vicinity of Crooked Lake, J.B. McFarlin 3365 , October 
28, 1930 (FLAS, MICH). Sarasota County: Sarasota, A. P. Garber 
46, June, 1878 (F, FLAS, PH, US). Volusia County: Dry scrub near 
Seville, A.H. Curtiss 6687 , July 16, 1900 (G, GA, GH, L, MO, UC , 
US). County unknown: F. Rugel , 1842-1849 (MO, US). 

12. Bonamia elli^tica (Smith & Schubert) Myint & Ward, comb. nov. 
Breweria elliptica Smith & Schubert, Contr. Gray Herb. n. 
s. No. CXXVII: 31, tab. II, fig. 31 & 32. 1939. 

Perennial, shrubby, twining vines. Stems woody, terete, white, 
densely pilose, mostly 2-4 mm. thick. Leaves petiolate, soft, 
leathery or subcoriaceous , long-strigose above, densely strigose 
beneath; petioles 7-10 mm. long, densely strigose; blades elliptic, 
3-5.5 cm. long, mostly 2.5-4 cm. wide, rounded and slightly oblique 
at the base, obtuse-mucronate at the apex; veins distinct, with 
about 6-8 pairs of lateral veins. Inflorescences axillary, cymose, 
of 5-12 (-15) flowers, pedunculate; peduncles 2.5-5.5 cm. long, 
1-1.5 mm. thick, densely strigose; secondary peduncles as long as 
pedicels; pedicels 7-10 mm. long, densely strigose as peduncles, 
mostly 1 mm. thick; bracts linear or lanceolate, pilose, 3-10 mm. 
long, acuminate. Sepals ovate-acuminate, equal or slightly un- 
equal, densely strigose or becoming less so, with ciliate, thin 
margins, 12-15 mm. long, 4-7 mm. wide. Corolla blue or pale blue, 
infundibulifoiTii, mostly 4-5 cm. long, slightly lobulate or sub- 
entire, long-strigose on interplicae, glabrous on plicae. Stamens 



170 PHYTOLOGIA Vol. 17, no. 3 

inserted; filaments glabrous or only with scatt<.'recJ short hairs, 
unequal (two short, two lonp, and one medium, which are as high 
as stylos); anthers oblong, 3-4 mm. long, cordatr" at the base. 
Ovary conical, glabrous; styles bifid three-fourths the length, 
glabrous, as long as the medium stamen; stigma capitate. Fruit 
unknown. 

Type: Chorrera, Temascaltepec Dist. Mexico, G.B. Hinton 2176 , 
10. 14. 1932 (GH-holotype). 

Known only from Temascaltepec Dist., Chihuahua, Mexico CMap 5). 

Further collections of this species are to be desired, since 
it is known only from two collections from the same district, and 
fruit and seed are not yet known. The second collection has been 
associated with the vernacular name "manto."' 

This species is related to B. sulphurea of southern Mexico and 
Central America, from which it is different by its long pedunculate, 
much branched cymes of large numerous flowers, borne in the axils 
of the leaves of the primary branches. 

Specimens examined: 

MEXICO: Chihuahua: Chorrera, 1230 m. , Temascaltepec Dist., 
vine, flower blue, G.B. Hinton 2176 , 10. 14. 1932 (F, GH, MO, US); 
Volcan, Temascaltepec Dist. , flower blue, Hinton et al. 8487 , 9. 
24. 1935 (MO, US). 

13. Bonamia sun^ghurea (Brandg.) Myint & Ward, comb. nov. 

Breweria sulphurea Brand egee, Univ. Calif. Publ. Bot. 
4:384. 1913. 

Perennial, shrubby climber. Stems terete, mostly twining, 
pubescent and becoming glabrous in age, about 2-5 mm. in diameter. 
Leaves petiolate, coriaceous or subcoriaceous , greenish pubescent 
and glabrescent above, densely brown tomentose below; petioles 
3-20 mm. long, 1-1.5 mm. thick, sulcate above, finely pubescent or 
becoming glabrous; blades broadly ovate or elliptic, 3.5-7.5 cm. 
long, 2-4.5 cm. broad, entire at the margin, rounded or slightly 
cordate at the base, and acuminate, obtuse-mucronate or rarely 
acute-mucronate at the apex; veins inconspicuous above, prominent 
beneath; about 4-7 pairs of lateral veins. Inflorescences loose 
cymes of few flowers, rarely solitary in the axils of small leaves, 
frequently pseudopanicles composed of numerous cjmies on short 
leafy twigs; peduncles variable in length, sometimes hardly present; 
pedicels distinctly elongate, 1-2 (2.5) cm. long, 1-1.5 mm. wide, 
pubescent; bracts small, lanceolate, mostly inconspicuous. Sepals 
ovate-lancGolate, acute or obtuse at the apex, equal or subequal, 
densely brown-tomentose outside, inner two less densely so, 8-13 
mm. long, 3-7 mm. wide, coriaceous or membranous. Corolla white, 
mostly 1.5-2 cm. long, tubular-campanulate with narrow limb, lobu- 
late at the margin; outside surface pilose on the interplicae, 



1968 l^int & V/ard, Revision of Eonamia 179 

glabrous on the plicae. Stamens included; filaments adnate to the 
corolla tube, glabrous above, pilose near the base, shorter than 
styles; anthers oblong, about 2 mm. long, 1 mm. broad. Ovary hir- 
sute; styles bifid above the middle; stylar branches unequal; stig- 
mas globose-capitate. Fruit subconical capsule, valvular, shorter 
than persistent sepals, brown hirsute at the apex. Seeds ovate, 
glabrous, black. Cotyledons ovate-orbicular. 

Type: Mexico: Vera Cruz, Banos de Carizal, C.A. Purpus 5995 
(UC, 155241). 

Southern Mexico, Gautemala and Honduras at the altitude of 
200-1000 m. (Map 5). 

Most collectors reported damp bushy slopes, damp thickets, 
and rocky slopes as the habitat of this species. More collections 
will be needed to determine its general distributions, type of 
habitat and flowering periods. Specimens, mostly in flower, have 
been collected in August, September and October. 

This species is undoubtedly related to B. elliptica of north- 
ern Mexico. From the latter it is different in possessing flowers 
with longer pedicels and smaller corolla and less branched cymose 
inflorescences which are borne in the axils of leaves of the 
secondary leafy branches, rather than in the axils of leaves of 
the primary branches. 

Both this genus and this species are reported here from Guate- 
mala and Honduras for the first time. Previously this species has 
been known only from southern Mexico. In this general area B. 
brevipedicellata is also reported from British Honduras for the 
first time. 

Specimens examined : 

GUATEMALA: Chiquimula: Divide on the railway above El Ricon, 
alt. 870 m. , damp bushy slope, woody vine, Paul C. Stand ley 74730 , 
October 17, 1940 (F); damp bushy slope, vine over trees, corolla 
white, Standley 74755 , October 17, 1940 (F). Zacapa: Vicinity of 
Zacapa, alt. ca. 200 m. , damp thicket, large woody vine, buds only, 
Standley 74201 , October 7-16, 1940 (F, US); rocky slopes between 
San Pablo and Pepezca, alt. 200-250 m. , climbing in thickets, cor- 
olla white, leaves olive-dull green, yellow-green beneath, Julian 
A. Steyermark 29337 , October 8, 1939 (A, F). 

HOhfDURAS : Morazan: La Granja, along Rio Choluteca near Tequci- 
galpa, alt. 936 m. , fls. white, vine, Antonio Molina R. 10493 , 
September 8, 1946 (BM, MO, US). 

MEXICO: Vera Cruz: Banos del Carizal, C.A. Purpus 5998 , Aug- 
ust, 1912 (UC-holotypeJ BM, F, MO, NY, US-isotype). 

14. Bonamia ovali folia (Torr. ) Hall. f. Bot. Jahrb. 16:528. 1893. 
Evolvulus ovalifolius Torr. Bot. Mex. Bound. 150. 1859. 
Breweria ovalifolia (Torr.) A. Gray, Syn. Fl. N. Am. 2(1): 
217. 1878. 



lOO P H Y T L G I A Vol. 17, no. 3 

Ppmnmal, procumbont or subercct, occasionally prostrate, 
shrubby vinos, growing from lower nodes of old shoot or root. 
Root thick, mostly 5-12 mm. near the base, with pulpy bark. Old 
stems 3-10 mm. or thicker, woody; new branches 3-6 dm. tall, 
wiry or slightly woody, light green, densely sericeous or velu- 
tinous. Leaves sessile or subsessile, rarely with short petioles 
of 1-3 mm. long, soft and leathery or subcoriaceous , densely 
velutinous on both upper and lower surfaces; blades ovate, ob- 
long-ovate or rounded, 1.4-2.6 (3.0) cm. long, mostly 1-2 cm. 
wide, occasionally smaller, rounded or obtuse or slightly cordate 
at the base, obtuse or abruptly acute at the apex; lateral veins 
2-5 pairs, most commonly 3 or 4 pairs. Flowers axillary, solitary, 
shortly pedicellate or almost sessile, bracteate; peduncles al- 
most absent; pedicels 2-8 mm. long, densely villous; bracts two, 
opposite, close to leaf axils, linear or linear-lanceolate, 3-6 
mm. long. Sepals ovate or ovate-lanceolate, acute or acuminate 
at the apex, 9-14 mm. long, 7-12 mm. wide, slightly unequal, or 
subequal, partially united at the extreme base, densely villous 
outside, soft-coriaceous or subcoriaceous. Corolla blue or bluish 
purple, paler on interplicae and lower part, 3.5-5 cm. long, 
2.8-4 cm. wide, funnel-form or tubular campanulate, 5-10 lobu- 
late or subentire, hirsute with scattered long hairs on inter- 
plicae, glabrous on plicae. Stamens included; filaments with 
long, frequently interwoven, white hairs, unequal; anthers 3-4 mm. 
long, cordate at the base. Ovary conical, with long hairs near 
the apex; styles bifid about one-fourth the length, slightly to 
distinctly longer than filaments, with scattered and long hairs 
to nearly glabrous, with unequal stylar branches; stigmas minute. 
Capsules globose, apiculate, villous near the apex, usually 2- 
to 4-seeded, rarely 1-seeded by abortion; seeds globose, com- 
pressed on the inner side, glabrous, brown. Cotyledons bilobed 
(being emarginate at the apex and cordate at the base), flat while 
young, folded when mature. 

Type: Mexico: On the Rio Grande below San Carlos, C.C . 
Parry , October (GH). 

Limited to the valley of the Rio Grande River, on deep, sandy, 
arid plains in Mexico and Texas (Map 5). 

The distribution of this species has been extended to New Mex- 
ico by House (1907) , but no specimen has been seen to authenticate 
such extension. Further collections of this species are needed, 
as it is known only from two locations and representative specimens 
are very rare, even in the larger herbaria. 

This species is more closely related to B. multicaulis than 
to any other known species of the genus. It can, however, be 
distinguished from the latter by its oval leaves, wiry or thin 
procumbent stems , filaments with long hairs and longer stylar 
fusion. 



1968 l/yint Sc V/ard, Revision of Bonamia l8l 

Specimens examined: 

MEXICO: Coahuila: Rio Grande, below San Carlos, C.C. Parry , 
October, Mexican Boundary Survey under the direction of Major W.H. 
Emory (GH). 

UNITED STATES: Texas: Brewster County: Big Bend National 
Park, arid desert plains and hills, Boquillas Canyon, locally 
common along edge of slope of deep sand, G.L. Webster 4482 , July 
22, 1952 (SMU, W) . 

15. Bonamia multicaulis (Brandg.) House, N.Y. State Mus. Bull. 
233-234: 61. 1922. 

Br ewer i a multicaulis Brandegee, Univ. Calif. Pub. Bot. 
4:185. 1911. 

Perennial, woody subshrubs. Roots woody, thick, with pulpy 
bark. Stems woody, terete, densely sericeous with silvery hairs, 
numerous annual culms from thick stumps of previous-year shoots, 
5-6 mm. near the base, occasionally thicker, 2-6 dm. high. Leaves 
sessile, rarely with short petioles of 1-2 mm. , soft and leathery 
or subcoriaceous , densely sericeous on both surfaces; blades lanceo- 
late, 1.5-3.5 cm. long, 4-10 mm. wide, mostly cuneate or rarely 
acute or obtuse at the base, acuminate or acute at the apex; veins 
inconspicuous except the midrib, rarely lower pair of lateral 
veins barely visible. Flowers axillary, solitary, shortly pedun- 
culate or shortly pedicellate or almost sessile; bracts two, linear 
or linear-lanceolate, 4-6 mm. long, 1-1.5 mm. wide, densely seri- 
ceous. Sepals ovate or ovate-lanceolate, acuminate at the apex, 
10-13 mm. long, 6-8 mm. wide, equal or slightly unequal, densely 
sericeous or villous, soft-coriaceous or subcoriaceous. Corolla 
blue, paler on interplicae and lower part, 3-4 cm. long, with 
limb of about 2.5-3.5 cm. in diameter, tubular-campanulate, entire 
or subentire, hirsute on interplicae, glabrous on plicae; tube 
short, about 1 cm. long. Stamens included; filaments glabrous 
with short, scattered hairs above, pilose on the basal parts adnate 
to corolla tube; anthers oblong or oblong-ovate, 3-5 mm. long, 
slightly cordate at the base, rounded at the apex. Ovary long- 
hirsute or sericeous, conical; styles bifid toward the middle or 
higher, glabrous above, with scattered hairs near the base, longer 
than filaments, slightly shorter than corolla; stigmas minute. 
Fruits valvular capsules, 2- to 4-seeded, or one-seeded due to 
aborted condition, conical, sericeous, becoming less sericeous 
in age; seeds glabrous, black or dark brown. Cotyledons oboval or 
bilobed with emarginate apices; cotyledonary petioles free. 

Type: On sand dunes near Sierra del Rey, Coahuila, Mexico, 
C.A. Purpus 4457 , June, 1910 (UC). 

From the material examined , it appears that this species is 
localized in Coahuila in northern Mexico (Map 5). 

Brandegee, in describing this species, correctly stated that 
it is nearest to B. ovalifolia. The two species are similar in 



182 PHYTOLOGIA Vol. 17, no. 3 

their habit, indumr'ntum, Sf'pa]s and corolla. Howr/vfr, H. multi - 
caulis can roadily bo distinguished from M. ovalifol ia by its 
narrow, lanceolate leaves, thicker and erect stf-ms , and nearly 
glabrous filaments. 

Specimens examined: 

MEXICO: Coahuila: On sand dunes, near Sierra del Rey, C.A . 
Purpus 4457 , June, 1910 f UC-holotype; BM, F, GH , MO, US-isotypes) ; 
21 mi. west of Kl Oro, road to Guimbalete, flowers blue, S.S. White 
2013, July 24, 1939 (GM, MDXU, MICH). 

16. Bonamia sericea (Griseb.) Hall. f. Bot. Jahrb. 16:528. 1893. 
Breweria sericea Griseb. PI. Lorentz. 181. 1874. 
Convolvulus Breweraceus 0. Ktze. Rev. Gen. 3C2):212. 1898. 

Perennial shrubby or herbaceous -suffrutescent, erect or procum- 
bent plants. Roots woody, thick near the base. Stems woody at 
the base, 5-15 mm. thick; new stems herbaceous above, somewhat woody 
at the base, 15-40 cm. high, 1-3 mm. thick, densely to finely seri- 
ceous with soft-appressed short hairs; intemodes 5-40 mm. long. 
Leaves shortly petiolate, coriaceous, subcoriaceous or membranous, 
sericeous on both upper and lower surfaces; petioles 1-7 mm. long; 
blades elliptic, elliptic-lanceolate or elliptic-ovate, 8-35 mm. 
long, 3-20 mm. wide, obtuse, acute or attenuate at the base, obtuse 
or abruptly acute and mucronate at the apex; midrib prominent with 
4-6 pairs of lateral veins. Flowers shortly pedunculate, solitary 
or in cymes of 2-3, axillary or frequently terminal; peduncles 
short, 3-10 ram. long, sericeous; pedicels 2-5 mm. long, sericeous; 
bracts linear or linear-lanceolate, minute, 2-3 mm. long. Sepals 
ovate or ovate-acuminate, acute or acuminate at the apex, concave, 
7-10 mm. long, 3-4.5 mm. wide, pubescent. Corolla white, infundi- 
buliform-campanulate, 15-30 mm. long, ferrugineous -pubescent on 
interplicao. Stamens included; filaments glandular villous near 
the base; anthers oblong-sagittate, 3.5-4.5 mm. long. Ovary conical, 
densely pilose-hirsute near the apex; styles 12-14 mm. long, bifid 
above the middle; stigmas subglobose-capitate, papilose. Capsules 
subglobose or conical, 5-6 mm. in diameter, pilose-hirsute at the 
apex; seeds black, 3-5 mm. long, glabrous. Cotyledons oval, cor- 
date at the base; cotyledonary petioles fused. 

Type: Argentina: Cordoba; not available. 

Known only from northern Argentina, where it seems to be fairly 
abundant at an altitude of about 400-500 m. at a few localities 
(Map 6). Although it has been collected more than other species 
from southern South America, its habitat is poorly known. One 
collector ( Venturi 2074 ) noted its habitat as a railroad embankment. 
It has been collected in flower in October, November and December 
and in fruit in December. 

This species is rather variable in leaf shape and size, and in 
indumentum of stems and leaves. Mainly on account of these features, 
O'Donnell separated it into two varieties. 



1968 



L^yint & "vVard, Revision of Bonamia 



183 




• 


var. sericea 


• 


var. lati folia 


•X- B. 


boliviana 


^>^ B. 


holtii 



Map 6 



18U P II Y T L G I A Vol. 17, no. 3 

Kfy to Varieties 

1. Plants densely sericeous; leaves 3-10 mm. wide, 8-20 mm. long, 
narrowly elliptic to lanceolate var. sericea 

1. Plants less densely or sparsely sericeous or puberulous , with 
softer and shorter hairs; leaves 4-20 mm. wide, 15-35 rm. long, 
elliptic to olliptic-ovate var. lati folia 

B. sericea (Griseb. ) Hall. f. var. sericea . 

Stems 15-30 cm. long, densely pubescent; old stems woody, 5-15 
mm. thick; internodes 5-15 mm. long. Leaves shortly petiolate, 
densely sericeous on both upper and lower surfaces; petioles 1-4 
mm. long; blades narrowly elliptic to lanceolate, 8-20 mm. long, 
3-10 mm. wide, acute or obtuse and mucronate at the apex. Pedicels 
5-14 long; bracts 2-3 mm. long. Sepals 8-10 mm. long, 3-4.5 mm. 
wide. Corolla 15-30 mm. long. 

Specimens examined : 

ARGENTINA: Cordoba: Althos del S. Y. 0. B.W. Bodenbender 
8823 (NY, R) ; E. Fielding (BM); Ischilin, Quilino al km. 855, 
T. Meyer 13543 , 16. XII. 1947 (W) ; Ischilin, La Florida, Meyer 
13730 , 16. XII. 1947 (W) ; Barrio S. Martin, C.A. O'Donncl y. J.M . 
Rodriguez 329 , 17. III. 1944 (F, UC). Chaco (rA) : en los campos , 
flor. bianco, alt. 250, S. Venturi 9794 , November 19, 1929 (BM). 

B. sericea (Griseb.) Hall. f. var. latifolia O'Donell, Lilloa 
29:31. 1959. 

Stems 20-40 cm. long, sparsely sericeous; old stems woody, 
5-30 mm. thick; internodes 1-4 cm. long. Leaves shortly petiolate, 
sparsely sericeous; petioles 2-7 mm. long; blades elliptic to 
elliptic-ovate, 15-35 mm. long, 4-20 mm. wide, obtuse or occa- 
sionally acute and mucronate at the apex. Pedicels 5-8 mm. long; 
bracts 2-4 mm. long. Sepals 7-10 mm. long, 3-4.5 mm. wide. Co- 
rolla 30 mm. long. 

Type: Argentina: type specimen not available. 

Specimens examined : 

ARGENTINA: Sgo del Estero, Ojo de Agua ( aired ed ores ) , B. 
Baleguo 1379 , 17. XII. 1947 (W) ; Tucuman, Cruz Alta, saliendo 
de Las Cejas por el ramal que va a Antilla, C.A. O'Donell 5413 , 

14. XI. 1947 (W) ; Las Cejas, Tucuman, 450 m. , Schreiter 3956 
("1799") , 18. XL. 1923 (GH, NY, US); Santiago del Estero, El 
Palomara Pampa Pozo , 400 m. , f 1. blanca , Schreiter 6706 ("4046" ) , 

15. XI. 1931 (NY); Las Cejas, Cruz Alta, 400 m. , blanca, 0.40 
ra. , en campos abiertos, S. Venturi 1525 , December 3, 1921 (US); 
Las Cejas, Cruz Alta, 400 ra. , flor blanca (Terraplen del F.C. a 
Antillas), Venturi 2074, October 21, 1923 (A, GH, NY, US). 



1968 Ityint & V/ard, Revision of Bonamia 185 

17. Bonamia boliviana O'Donell, Lilloa 23:458, tab. 1. 1950. 

Perennial woody climbers. Stems twining, terete, 1-2.5 mm. in 
diameter, toraentose, with internodes of 1-4 cm. long. Leaves pet- 
iolate, subcoriaceous or soft-coriaceous, tomentose and glabres- 
cent above, more densely tomentose underneath; petioles 2-9 mm. 
long, tomentose; blades elliptic to ovate, 1-4 cm. long, mostly 
7-23 ram. wide, rounded, subcordate or truncate at the base, ob- 
tuse-mucronate or acute-mucronate at the apex; midrib impressed 
above, prominent beneath, with 5-7 pairs of lateral veins. Flowers 
solitary, in axils of normal or reduced leaves or in axillary 
racemose inflorescences of few flowers on short branches; pedun- 
cles short, 1-4 mm. or rarely longer, tomentose; pedicels 3-9 
mm. long, tomentose; bracts linear, alternate or occasionally op- 
posite, 2-4 mm. long, tomentose. Sepals slightly unequal or equal, 
coriaceous or subcoriaceous; outer ovate to subovate, 5-6 mm. long, 
4.5-5 mm. wide, concave, obtuse, toraentose; inner suborbicular , 
4.5-5.5 mm. long. 4.5-5 mm. wide, obtuse, toraentose above, glabrous 
along lateral hyaline margins. Corolla pale yellow, campanulate, 
17-18 mm. long, with entire or subentire limb, ferrugineous with 
long hairs on interplicae. Stamens included, 12-13 mm. long; fila- 
ments pilose with long hairs near the base; anthers oblong, 2.5-3 
mm. long. Ovary fusiform and attenuate to the stylar base, gla- 
brous; styles free nearly to the base, glabrous, unequal; stigmas 
reniform. Fruits and seeds unknown. 

Type: Bolivia: Cordillera, La Cuesta, 386 m. , flor amaril- 
lenta, I. Peredo , 8. II. 1946 (F, NY, US, W-isotypes). 

Known only by the type collection from Bolivia (Map 6) . 

This species is poorly known. Since furit and seed are not 
known, its placement in the section Bonamia is tentative until 
future collections are available. 

In superficial appearance, this species resembles some Bra- 
zilian species, particularly B. subsessilis and B. burchellii . 
However, it is well distinguished from these by its smaller leaves 
with indistinct intercostal veins, shorter stem and solitary or 
few-flowered cymes. It is also distinct from B. subsessilis by 
its pedicelled flowers. Future collections might show that this 
species should properly be placed in the section Trichantha to- 
gether with these Brazilian species. 

18. Bonamia holtii O'Donell, Lilloa 30:59. 1960. 

Perennial twining vines. Stems becoming woody, terete or 
slightly angular and striated, 1-2.5 mm. thick, sparsely and mi- 
nutely pubescent; internodes mostly 4-8 cm. long. Leaves shortly 
petiolate, subcoriaceous or leathery, sparsely pilose or becoming 
glabrous; petioles 8-17 ram. long, sparsely pilose; blades elliptic 
or elliptic-ovate, 5-9 cm. long, 3-5 cm. wide, rounded and slight- 
ly asymmetrical at the base, acute-mucronate or acuminate at the 



186 P I! Y T L G I A Vol. 17, no. 3 

apex; midrib prominont b^noath, with 4-6 pairs of lateral vr-ins. 
Inflorescences shortly pedunculate, axillary, dense-capitate cymes 
of few to several flowers; peduncles short, 2-9 mm. long, 1-1.5 mm. 
thick, minutely pubescent or sericeous; bracts foliaceous, ellip- 
tic or elliptic-lanceolate, 6-12 mm. long, 1.5-5 mm. broad; bract- 
eoles lanceolate, 3-5 mm. long, sericeous. S«-'palfi coriaceous or 
subcoriaceous , slightly unequal; outer sepals lanceolate or ovate- 
lanceolate, 8-10 mm. long, 3-5 mm. wide, acute or acuminate at 
the apex, appressed-tomentose or finely pubescent outside; inner 
sepals elliptic, ovate or ovate-elliptic, 6-7 mm. long, 4-5 mm. 
wide, obtuse or obtuse-acute at the apex, nearly glabrous or some- 
what pubescent on the median lines. Corolla white, infundibuli- 
form, 1.2-1.9 cm. long, with 5-lobulate or subentire limb, long- 
pilose on interplicae. Stamens inserted; filaments 7-14 mm. long, 
glabrous; anthers oblong, 3 mm. long, doi-si fixed, sagittate at the 
base. Ovary ovoid, long-pilose at the apex, glabrous below; styles 
fused for lower one-fourth and free above, filiform, glabrous; 
stigmas capitate. Capsules subglobose, 4 mm. in diameter, long- 
pilose at the apex, 4-valvular, 4-seeded; seeds black, 3 mm. 
long, glabrous, rugose. Cotyledons cordate, slightly asymmetrical, 
rounded at the apex; cotyledonary petioles fused. 

Type: Colombia: Rio Orinoco, Boca del Vichada, alt. about 
100 m. , E.G. Holt and 'W. Gehriger 223 , January 12-24, 1930 (IS- 
holotype; isotype at Caracas, Venezuela, not seen). The labels 
of the type carried the data, "Venezuela, Amazonas Territory: 
Rio Orinoco; Boca del Vichada;" thus specifying a location in 
Colombia at the mouth of the Rio Vichada. The species is endemic 
to this region and known only by the type collection (Map 6) . 

This species is different from all other South American species 
by its capitate inflorescence, acute or acuminate sepals, foliaceous 
bracts and bracteoles, and slightly twining and somewhat striated 
stem. It is, however, distantly related to B. umbellata because 
of the similarity in their inflorescence and indumentum. Bonamia 
holtii is poorly known and is described from two sheets of the same 
collection. The specific name is derived from the name of its 
collector and was first used as an unpublished name under the genus 
Prevostea by Dr. H. Pittier, a field botanist of Venezuela. 

19. Bonamia maripoides Hall. f. Bot. Jahrb. 16:529. 1893. 

Maripa spectabilis Choisy, in D. C. Prodr. 9:327. 1845. 
Prevostea spectabilis Meissner, in Mart. Fl. Bras. 7:325. 

1869. 
Calycobolus spectabilis (Choisy) House, Bull. Torr. Bot. 

Club. 34: 146. 1907. 

Perennial, twining liana growing all year around. Stems woody, 
terete, yellowish or brownish tomentose, glabrescent, climbing to 
20 m. or higher. Leaves petiolate, subcoriaceous, glabrous and 
shing above, golden or yellowish sericeous (with unidirectional 
hairs) beneath; petioles 8-15 ram. long, sericeous and glabrescent; 



1968 



Ifyint & Ward, Revision of Bonamia 



18? 




Distribution of 

• B. maripoides 

* B. ferruginea 



Map 7 



188 P H Y T L G I A Vol. 17, no. 3 

blades broad-ovato or flliptic, 6-J4 min. long, mostly 3-7.5 cm. 
broad, shortly acuminato or obtuse at the apex, rounded at the 
base, with 6-10 pairs of lateral veins. Inflorescences axillary, 
shortly peduncled, compound, umbelliform cymns , often secund; 
pedicels distinctly elongate, usually 1-2 cm. long, densely seri- 
ceous and glabrescent, ridged; bracts small, lanceolate, often 
inconspicuous. Sepals coriaceous, mostly equal or slightly un- 
equal in length, the two exterior orbicular or broad-elliptic, 
acutish and tomentose, the three interior orbicular, obtuse and 
nearly glabrous. Corolla white, funnel-shaped, mostly 2-2.5 cm. 
long, yellow-villous on interplicae, glabrous on plicae, entire 
or subentire. Stamens included; filaments, short, adnate to 
corolla tube, glabrous; anthers oblong, about 2 mm. long. Ovary 
hairy; styles bifid or free nearly to the base, longer than fila- 
ments; stigmas globose. Capsule ovate, acute, hairy at the apex, 
8-valved, 4- (2-) seeded, about 6 mm. long and 5 mm. in diameter. 
Seeds ovate, compressed on the inner side, 3-4 mm. long, black, 
glabrous. Cotyledons ovate or obovate. 

Type: Brazil: type specimen not available. 

Northern Brazil, British Guiana and Surinam fMap 7). 

According to the collectors, this species is a high-climbing 
woody vine to 20 m. on lofty shrubs or in virgin forest. No 
definite habitat has been recorded by any collector. It has been 
collected in flower in March, April, August and October, and in 
fruit in February, March, May, June and July. 

Specimens examined: 

BRAZIL: Amazonas : Borba, Rio Madeira, R.L. Froes 26109 , 28. 
11. 1950 (US). Para: Belem do Para, A. Ducke 3304, 5. 3. 1903 
(RB, US); Belem do Para, M. Guedes 1602 , 28. 5. 1898 (US). Peram- 
buco: Estrada de aldeia, flores alvas , trepadeira, C. Leal e Otavis 
Silva , 19. 7. 1950 (RB) ; flowers very numerous, white, abundant at 
one spot in the matto of Berberibe, climbing over the tops of lofty 
shrubs, Ridley, Lea and Ramage , October 4, 1887 (BM). 

BRITISH GULANA: Bullet Tree Island: Ebini Experiment Station, 
Berbice River, margin of jungle, S.G. Harrison 1243 , 28. VI. 1958 
(K). 

SURINAM: M. Berthond-Coulon 219 , 1841 (BM); Forest Reserve Zan- 
deriz 1, sand virgin forest, bud brown, liana, J. Lanjouw 362 , July 
31, 1933 (NY); Saramacca River, liana climbing to 20 m. , leaves 
dark green above, tawny sericeous beneath, bush to rear of Jacob 
Kondre, Bassett Maguire 23761 , June 19, 1944 (BM, F, NY, US); Cop- 
pername River near Onobissi, '' B.W." 1103 , 4. 3. 1915 (L) ; Copper- 
name River, Raleighfalls , -' B.W.-' 6232 , 2. 8. 1923 (NY, US). Lo- 
cation indefinite: Berlyn, Scandens in sylvis Paraensis prope 
plant, flores albi , F.L. Splitgerber 743 (L); Splitgerber 362 (L). 

20. Bonamia brevipedicellata Myint and Ward, sp. nov. 



1963 



l,Ii''int & Y/ard, lievision of Bonaiaia 



189 




Bonamia brevipedicellata 




1.5 X 



3 X 




3 X 



Bonamia maripoides 




Figure 3 




Leaf shape, inflorescence and floral parts of 

B. brevipedicellata , and inflorescence and 

floral parts of B. maripoides 



190 P I! Y T L G I A V&l. 17, rto. 3 

Frutex alte scandens, usque 16 m. Folia petiolis 2-3 cm. 
longis ; laminae ellipticao vel ovato-ellipticae, 8-12 cm. longae, 
4-7 cm. latae, basi rotundata vr-l obtaisa, apice breviter acumi- 
nato, supra glabrae, infra dense tenui-pubescentes. Inflorescen- 
tiae cymae breviter pedunculatae, dense mu]ti florae. Pedunculi 
breves, plerumque 1-3 cm. longi. Sepalo ovato-orbicularia , suba- 
equalia, 4-5 mm. longa, dense tenui-pubescentia. Corolla campan- 
ulata» 1-1.2 cm. longa, limbo angusto, viridi-alba, prompta de- 
cidua. Stamina inclusa; fila glabra; antherae basi cordata. 
Styli libri ad basim, minores quam 1 cm. longi; stigmata peltato- 
subglobosa. Fructus et semina ignota. 

Perennial, woody vines. Stems terete, about 1-2.5 cm. in 
diameter and 16 m. tall, puberulous or glabrescent. Leaves petio- 
late, membranous to subcoriaceous , glabrous above, densely fine- 
pubescent with unidirectional hairs underneath; petioles 2-3 cm. 
long, 1.5-2.5 mm. thick, finely pubescent or puberulous and glabres- 
cent, canaliculate above; blades elliptic to ovate-elliptic, 8-12 
cm. long, 4-7 cm. wide, rounded or obtuse at the base, shortly 
acuminate or acute at the apex; midrib impressed above, prominent 
beneath, with 5-6 pairs of lateral veins. Inflorescences shortly 
pedunculate, dense multiflorous cymes in the axils of young or 
reduced leaves on short branches; peduncles 4-10 mm. long, finely 
pubescent; pedicels short, mostly 1-3 mm. long, pubescent; bracts 
minute, 1-2 mm. long, linear. Sepals ovate-orbicular, subequal, 
4-5 mm. long, 3-4 mm. wide, densely fine-pubescent with grey or 
silvery grey hairs. Corolla greenish tinged, readily dropping off, 
campanulate, 1-1.2 cm. long, with narrow limb, hirsute on inter- 
plicae; tube cylindrical, wide, about 4 mm. long. Stamens in- 
cluded; filaments glabrous; anthers oblong, 2.5-3.5 mm. long, cor- 
date at the base. Ovary conical, with long hairs at the apex, 
glabrous near the base; styles free to the base, glabrous, equal 
or slightly unequal, the longer less than 1 cm. long; stigmas 
peltate-subglobose. Fruits and seeds not known. 

Type: British Honduras: Machaca, alt. 50 ft., very tall 
vine growing in broken forest in swampy places ; flowers light 
green which easily drop off in the process of felling. '"Rare," 
50 ft. , 1 in. diameter, W.A. Schipp 1210 , September 11, 1933 (GH). 

Although the type material is incomplete, it is clearly dis- 
tinct from all other known species. The material, even though 
lacking fruits and seeds, is sufficient to permit a technical des- 
cription. The name B. breviped icellata is derived from the very 
brief pedicels. 

This species is superficially very suggestive of B. maripoides , 
to which it is definitely related because of similarity of leaves, 
indumentum, sepals, cordate anthers, glabrous filaments, free styles 
and subglobose stigmas. However, B. breviped icellata shows a series 
of distinctive features which appear to offer a sound basis for ad- 
judging it a separate species. The leaves in B. maripoides are 
densely pubescent with long appressed hairs, whereas the hairs are 



1968 l^nt & Ward, Revision of Bonania 191 

shorter and soft in the B. brevipedicellata . The inflorescences 
in _B. maripoides are loose compound cymes, while the cymes in 
_B. brevipedicellata are dense. Individual flowers are long-pedi- 
cellate in B. maripoides , while they are short-pedicellate or 
almost sessile in B. brevipedicellata . The corolla in this species 
is very small (with a narrow and short limb) compared to the 
large corolla in B. maripoides and, as described by the collector, 
is greenish and readily deciduous (Figure 3). 

This species probably is a large climber, reaching to a height 
of 16 m. , and is rare according to the collector. It has been 
misidentified as a species of the Solanaceous genus Lysianthes . 

21. Bonamia ferruginea (Choisy) Hall. f. Bot. Jahrb. 16:530. 1893. 
Prevostea ferruginea Choisy, Annal. Sci. Nat. 4:498. 1825. 
Breweria ferruginea Hook. f. & Jackson, Ind. Kew. 1:337. 

1893. 
Calycobolus ferruginea (Choisy) House, Bull. Torr. Bot. Club. 

34:146. 1907. 
Not Trichantha ferruginea Karst. & Triana, Linnaea 28:438. 

1856. 

Perennial, densely f errugineous , woody climbers. Stems twining 
or scandent, terete, densely tomentose-f errugineous with crisped 
and brownish hairs, frequently branching. Leaves shortly petiolate, 
herbaceous, subcoriaceous and soft or leathery, densely ferrugineous 
on both surfaces, more densely so underneath, with brown or reddish 
brown hairs; blades broad-ovate or elliptic-ovate, 5-13 cm. long, 
3-8 cm. wide (upper leaves subtending inflorescence and on young 
shoots smaller), abruptly acute or obtuse and mucronate at the apex, 
rounded or cordate at the base; midrib prominent, with about 6-10 
pairs of lateral veins. Inflorescences sessile or pedunculate, 
multiflorous, capitate cymes in the axils of upper or reduced leaves, 
frequently on short lateral branches; peduncles, when present, fer- 
rugineous like stems; pedicels absent or very short, frequently some- 
what elongate in fruiting stage; bracts linear, 5-10 mm. long, oc- 
casionally reduced, ferrugineous. Sepals coriaceous, unequal; the two 
exterior larger, broad-ovate, 8-10 mm. long, 5-7 mm. wide, occasionally 
smaller, densely ferrugineous outside, pubescent along recurved margin 
inside, obtuse and reflexed at the apex; the three interior smaller, 
orbicular or ovate-orbicular, 4-6 mm. long, 3-5 mm. wide, glabrous 
or sparsely minute-pubescent, rounded or truncate at the apex. Co- 
rolla white, tubular-campanulate , 1.2-1.8 mm. long, with sublobulate 
limb, pilose outside on interplicae, glabrous on plicae; tube dis- 
tinct, narrow. Stamens included; filaments glabrous; anthers oblong, 
3-4 mm. long, cordate at the base. Ovary globose or conical-globose, 
apiculate, 4-valved , 2- to 4-seeded; seeds triangular-ovate, brown, 
glabrous. Cotyledons oval-cordate, rounded or truncate at the apex, 
broadly cordate at the base, in hard cartilagenous endosperm; coty- 
ledonary petioles fused. 

Type: Brazil, Amazonas : type specimen not available. 



192 P H Y T L G I A Vol. 17, no. 3 

Judging from tho sppcimens rrxaminpd in this study, this species 
appears to bo localized in the states of Amazonas , northwestern 
Brazil, from Manaus and Borba to Tefe TMap 7), A few collectors 
recorded edge of forest and dry highland as the habitat of this 
species. It has been collected in flower from May to September 
and in fruit from August to October. 

This species was originally described under Prevostea r= Ca]y - 
cobolus ) by Choisy and was accepted by Meissner. Hallier, reali- 
zing that its sepals are not accrescent, transferred it to Bonamia . 
House later transferred the species to Calycobolus because of the 
unequal sepals which he erroneously thought characterized this 
latter genus. 

Specimens examined : 

BRAZIL: Amazonas: Manaos , loco Flores , silva secundaria non 
inundabili, Frutex scandens, flor. albis, Ducke 210 , 30. 5. 1936 
(A, R); Tefe, A. Ducke 18017 , 15. 6. 1906 (RB) ; municipality of 
Borba, near Urucurituba, basin of Tio Madeira, B.A. Kruoff 5952 , 
September 4-6, 1934 (BM), .5953 (G) ; Ega Amazonas, in Sylvan Margin, 
Poeppig 2589 , September, 1831 (F, GH , W) ; Manaos, Estrada da Raiz, 
vine, flower white, R.E. Schultes and G.A. Black 8085 , August 7-12, 
1946 (GH, MO, NY, US); Manaus, Schwacke 210 , 1882 (R) ; ad oram 
meridionalem Rio Negro, usque ad coneursum flum. Solimoes , R. Spruce 
1568 , Maio 1851 (BM, G, W) ; flowers white, J.W.H. Traill 558 , 12. 
6. 1874 (K) ; Rio Negro, Windent auf Gestraiich bei Flores, Manaus, 
Bluto Weiss, E. Ule 5195 , July 29, 1900 (G, HBG, L) . Unknown col- 
lector: Fragment ex Herbario Musei Paris iens is (F). 

22. Bonamia umbellata (Choisy) Hall. f. Bot. Jahrb. 16:530. 1893. 

Prevostea umbellata Choisy, Ann. Sci. Nat. 4:497. 1825. 

Calycobolus umbellata (Choisy) House, Bull. Torr. Bot. Club 
34:146. 1907. 

Perennial, ligneous or herbaceous and suffrutescent vines. 
Stems t;>?ining or scandent, 1.5-3 mm. thick, pilose while young, 
puberulous or becoming glabrous in age; intemodes usually 3-8 cm. 
long, occasionally shorter. Leaves petiolate, soft and herbaceous, 
rarely leathery, appressed-pilose on upper surface, more densely 
so underneath, becoming less pilose or nearly glabrous in age; 
petioles 4-17 mm. long, 1-1.5 mm. thick, pilose or glabrate; blades 
oblong-ovate, mostly 3.5-8 cm. long, 2-4.5 cm. wide, cordate, sub- 
cordate or occasionally rounded at the base, obtuse-mucronate at 
the apex, with about 5-7 pairs of lateral veins. Inflorescences 
pedunculate, axillary, subumbellate cymes of few to many flowers 
(mostly 5-15); peduncles variable in length, mostly 2-7 cm. long, 
occasionally much shorter, 1.5-2 mm. thick, slightly thicker when 
fruits mature, pilose as in stems; pedicels 5-15 mm. long, thinner 
than peduncles, pilose; bracts minute or foliaceous , lanceolate, 
2-17 mm. long. Sepals herbaceous or subcoriaceous , puberulous, 
glabrescent or with scattered hairs; exterior two larger, ovate, 
6-10 mm. long, 5-9 mm. wide, obtuse or rounded at the apex; in- 



1968 l^nt & Ward, Revision of Eonamia 193 

terior three narrower or shorter, usually glabrous (except at the 
base), ciliate. Corolla white, funnel-shaped, 2.5-3 cm. long, 
with tube slightly longer than sepals, subentire or lobulate at the 
limb, long-pilose on interplicae. Stamens included; filaments 
glabrous, shorter than styles; anthers oblong, about 3.3 mm. long, 
cordate at the base. Ovary conical, about 3-4 mm. long, glabrous; 
styles bifid for upper one-fourth or one-fifth, glabrous; stigmas 
globose-capitate. Capsules globose, 5-6 mm. in diameter, 4- 
valvular, 2- to 4-seeded; seeds triangular-ovate, 3-4 mm. long, 
dark brown, glabrous. Cotyledons ovate-cordate, rounded at the 
apex; cotyledonary petioles fused. 

Type: Brazil: type specimen not available. 

Kno\sm only from southern Brazil (Map 8). 

Meissner (1869), while treating this species under the genus 
Prevostea, proposed a new variety in addition to the typical one, 
mainly based on the length of petioles. This feature, however, 
is extremely variable and no satisfactory line can be drawn to 
account for infraspecific segregation in the species. As Hallier 
did not make a transfer of Meissner's new variety, it is evident 
that Hallier did not accept it. 

Specimens examined : 

BRAZIL: Burchell 775 (NY); Burchell 1858 (K) ; bushy places 
by Rio de Janeiro, Gardner 5560 , July, 1841 (BM) ; Santa Theresa 
(Rio de Janeiro) voluvel, flores blanca, r. 26 de Dezembro de 1869, 
G. Glaziou 4131 (R) ; Rio de Janeiro, Schott 5462 (W) ; Rio de 
Janeiro, Sellow 225 (NY); Rio de Janeiro, G. Staunton (BM, W) ; 
Estado de Rio, Morro da Nova, Cintra, Trepadeira, E. Ule 3849 , 
February 25, 1896 (HBG, R) . 

23. Bonamia sphaerocephala (Dammer) v. Ooststr. Rec. Trav. Bot. 

Neerl. 33:212. 1936. 

Prevostea sphaerocephala Dammer, Bot. Jahrb. 23 (Beibl. 57): 
37. 1897. 
Perennial shrubby herbs growing erect from the base of old shoot. 
Stems woody, erect or suberect, about 50-80 cm. long, densely tomen- 
tose or lanate with grey or silvery grey soft hairs; young stems 
3-4 mm. thick, single or occasionally with one or two lateral 
branches. Leaves sessile or subsessile, subcoriaceous , lanate or 
tomentose on upper surface, densely white woolly underneath; blades 
oblong-elliptic, ovate-elliptic or elliptic-lanceolate, 2-4.5 cm. 
long, 1-2 cm. wide, sometimes slightly narrower, subcordate or 
truncate at the base, obtuse-mucronate or acute-mucronate at the 
apex; revolute at the margin; veins distinctly impressed above, 
prominent underneath; lateral veins 3-5 (6) pairs. Inflorescence 
terminal, multiflorous, dense capitate, 2-3.5 cm. in diameter; 
flowers sessile or shortly pedicellate; bracts linear, as long as 
and long-pilose as the sepals. Sepals coriaceous or subcoriaceous, 
unequal; exterior two larger, thicker, lanceolate-acuminate, 6-8 
mm. long, 1.5-2 mm. wide, densely long-pilose outside; inner three 



191; 



P II Y T L G I A 



Vol. 17, no. 3 




Distribution of 
•Jf- B. umbellata 
^^ B. sphaerocephala 
(•) B. kuhlmannii 
Ml. B. peruviana 



1968 Ityint & Ward, Revision of Bonania 195 

smaller and submembranous , lanceolate and shortly acuminate or 
acute, 4-5 mm. long, 1.5-2 mm. wide, long-pilose outside. Corolla 
white or blue, tubular-campanulate or funnel-form, slightly longer 
than outer sepals, mostly 9-12 mm. long, densely long-pilose on 
interplicae, with narrow, entire or subentire limb. Stamens in- 
cluded; filaments glabrous, slightly shorter than styles; anthers 
oblong. Ovary conical, with dense, long hairs near the apex; 
styles bifid for upper half, hairy at the base; stylar branches 
glabrous; stigmas globose-capitate. Capsules subangular globose, 
apiculate, finely pubescent near the apex, glabrous below, with 
coriaceous wall, 2- to 4-seeded, breaking by anmilar scission at 
the base; seeds oval, glabrous, dark brown. Cotyledons oval. 

Type: Brazil, Haut de la Serra Dourada, a Olha d'Agua pres 
de Goyaz, M.A. Glaziou 21797 , August 13, 1895 (BM-lectotype, R- 
isotype). Van Ooststroom designated a specimen at Berlin as the 
type; this material, however, was not included in a recent loan 
and is presumed to have been destroyed during the war. A dupli- 
cate specimen at the British Museum is designated here as the 
lectotype. 

This species is known from southern Brazil (Map 8). Van 
Ooststroom reported it from Minas Geraes , but no specimen was seen 
in the present study. Further collections with detailed descrip- 
tion of habitat and flower color are to be desired, since its 
habitat is not recorded by the collectors and flower color is 
differently recorded by the same collector ( Glaziou 21797 , BM, R). 

The outstanding features of this species are the strongly 
nerved and lanate leaves with long mucros , the erect and single 
stem, terminal globose heads, lanceolate-acuminate sepals with 
long hairs and small corolla. It seems to be related to the 
African species, B. mossambicensis , because of the following com- 
mon features: inflorescence dense, unequal sepals with long hairs, 
and long bracts with long hairs. However, the two can be readily 
distinguished by their habit, length of stem, shape, size, apices 
and petioles of leaves and size of sepals and corolla. 

Specimens examined : 

BRAZIL: Haut de la Serra Dourada, Goyaz, fl. b]anc, M.A. 
Glaziou 21797 , August 13, 1895 (BM); Serra Dourada (Goyaz), frutes- 
cente, flores azulos , Glaziou 21797 , August 13, 18Q5 (R) ; Serra 
Dourada, Goias , subarbusto campestre, Agnes, A. Macedo 3730 , 30. 
VII. 1952 (MO, NY). 

24. Bonamia kuhlmannii Hoehnc, Anex. Mem. Inst. Butantan 1 (4): 
44, tab. 2. 1922. 

Perennial, high-climbing, shrubby vines. Stems woody, 2-4 mm. 
thick, densely short-ferrugineous or subvelutinous. Leaves petio- 
late, submembranous or soft and leathery, softly velutinous on 
both surfaces, lighter in color underneath; petioles 1-2.5 cm. long. 



196 PHYTOLOGIA Vol. 17, no. 3 

1-2 rnm. thick, shortly ferruginoous ; blades ovatf or ovatf-cordatf, 
5-12 cm. long, 3-8 cm. wide, cordate or truncate, rarely rounded 
at the base, obtuse-mucronate at the apex; midrib slightly impres- 
sed above, prominent beneath, with 5-7 pairs of lateral veins. 
Inflorescences shortly pedunculate, axillary, simple or compound 
cymes of few to several flowers; peduncles 1-2 cm. long, 1-2 mm. 
thick, short-ferrugineous as peduncles, bracts small, linear or 
triangular-acuminate, 1-2 mm. long, ferrugineous. Sepals subcoria- 
ceous or herbaceous, unequal; exterior two larger, ovate or ovate- 
subcordate, 1.2-2 cm. long, 10-17 mm. wide, occasionally smaller, 
adnate to pedicels at the base, obtuse at the apex, densely velu- 
tinous-ferrugineous outside, shortly ferrugineous inside (except 
at the glabrous center), interior three ovate-orbicular, 5-7 mm. 
long, glabrous or sparsely pubescent. Corolla white, narrowly 
campanulate or funnel-form, 2.5 cm. long, sparsely pilose on inter- 
plicae. Stamens included; filaments glabrous; anthers oblong, 
sagittate at the base. Ovary ovoid-conical, glabrous; styles bi- 
fid to the middle or nearly to the base; stigmas globose. Fruits 
unknown. 

Type: Brazil: Mato-Grosso: Comissao Rondon, entre Buriti 
e Formigueiro, liana do cerrado, fl. alva, J.G. Kuhlmann 2268 , 
6-1918 (R-isotype). 

This is a poorly known species, rarely collected and so far 
known only by the type collection from southwestern Brazil (Map 8). 
The type specimen was collected with flowers which mostly are only 
in bud. Its fruit is unknown and a complete description of the 
species has to wait future collections. 

This species is characterized by its completely ferrugineous 
parts, large leaves (cordate or subcordate at the base), unequal 
sepals, short peduncles and nearly free styles. It, however, is 
distinct in its densely ferrugineous leaves, which are cordate or 
subcordate at the base and obtuse-mucronate at the apex, larger 
sepals, shorter peduncles and pedicels, and deeper bifurcation of 
styles. 

25. Bonamia peruviana van Ooststroom, Recu. Trav. Bot. Neerl. 30: 
192. 1933. 

Perennial liana. Stems woody, scandent, 2-4 mm. thick, densely 
ferrugineous-tomentose; intemodes 2-6.5 cm. long. Leaves petio- 
late, subcoriaceous or soft and leathery, minutely tomentose a- 
bove, more densely so underneath; petioles 6-16 ram. long, mostly 
1 mm. thick, tomentose similar to stems; blades ovate or ovate- 
elliptic, 5-7 cm. long, 2.5-4 cm. wide, rounded at the base, shortly 
and acutely acuminate at the apex; midrib distinctly impressed 
above, prominent beneath, with 6-8 pairs of lateral nerves. In- 
florescences pedunculate, axillary, simple or compound cymes of 
few to several flowers; peduncles 1.5-4.5 cm. long, sometimes 
apparently dichotomous due to absence of central flowers ; pedicels 
slender, elongate 1.5-2 cm. long, occasionally longer, minutely 



1968 Myint & Ward, Revision of Bonamia 197 

tomentose, bi-acts linear-subulate, about 2 min. long. Sepals sub- 
coriaceous or herbaceous, unequal; exterior two larger, broadly 
ovate, 8-14 mm. long, 7-12 mm. wide, densely ferrugineous-tomen- 
tose on both surfaces except glabrous center inside, subcordate 
at the base, obtuse or subobtuse at the apex; interior three smal- 
ler, orbicular, 4-5 mm. long, glabrous. Corolla white, infundi- 
buliform, 2-2.5 cm. long, sparsely pilose on interplicae. Stamens 
included; filaments filiform, glabrous; anthers oblong, 2.5-3 mm. 
long, cordate at the base. Ovary ovoid-conical, glabrous; styles 
bifid for the upper half, with slightly unequal branches ; stigmas 
globose. Fruits glabrous, known only in immature stage; seeds 
glabrous. 

Type: Peru: Loreto: Michuyaeu, near Iquitos , at 100 m. , 
liana, fls. v;hite5 clearing (forest) G. Klug 232 , October-November, 
1929 (F-holotype; NY, US-isotypes ; G, L-fragments) . 

The type collection, the only material available for this 
study, was collected in flower and in young fruit, and is not 
sufficient for a description of the fruit. Cotyledons dissected 
from immature seeds appear to be ovate-cordate with fused cotyle- 
donary petioles. 

This species is closely related to B^. kuhlmannii of south- 
western Brazil; however, it is different from the Brazilian species 
by its leaf size, form, apex and base, indumentum and smaller se- 
pals. Macbride (1959), in his Convolvulaceae of Peru, added a 
comment that the differences between B. peruviana and B^, kuhlmannii 
may prove to be due to age and variability in a series of collec- 
tions. Since both species are known only from type collections, 
this statement cannot now be verified. From the materials avail- 
able at present they appear to differ in several features and thus 
are treated here as distinct species. 

II. Section: Breweria (R. Br.) Myint, Burma Jour. Life Sci. 1:31. 
1968. 
Breweria R. Br. Prodr. Fl. Nov. Holl. 487, 1910. 

Stems herbaceous, woody or becoming woody, prostrate, procum- 
bent, twining or erect, usually short, 1-2 m. long, rarely longer, 
mostly 1-2 mm. thick or slightly thicker. Leaves sessile or short- 
petiolate, soft, herbaceous, rarely subcoriaceous , mostly thin; 
blades elliptic, ovate-elliptic, oblong-elliptic, linear, lanceolate, 
oblong-ovate, ovate, ovate-subcordate or occasionally orbicular, 
3 cm. or shorter, rarely somewhat longer, mostly 2 cm. or narrower, 
rarely 2.5 cm. or slightly wider, with length-width ratio of 1.5 
or higher, occasionally lower, rounded, subcordate or truncate at 
the base; obtuse, rounded or slightly emarginate and mucronate at 
the apex. Flowers axillary, sessile or shortly pedunculate, soli- 
tary or rarely in simple cymes of two or three; bracts linear, small 
or as long as pedicels or longer, persistent. Sepals herbaceous, 
subcoriaceous or rarely somewhat coriaceous, equal or unequal, 
lanceolate, ovate-lanceolate, ovate or ovate-acuminate, acute, 



198 P H Y T L G I A Vol. 17, no. 3 

acuminate or rarely obtuse at the apirx, mostly 4-8 mm. long, outer 
sepals 12 mm. long (when unequal). Corolla white, blue, pink or 
red, 8-15 mm. long, occasionally longer, subentire or 5- to 10- 
lobulate at the margin. Stamens included; filaments glabrous or 
sparsely villous or rarely densely villous below and glabrous 
above; anthers 1-2 mm. long, rarely longer, slightly cordate at 
the base. Ovary ovoid or oblong-ovoid, sparsely or densely pilose 
or glabrous; styles free nearly to the base or fused to the middle 
and readily separable to the base; vascular traces single in the 
stylar branches, not branched; stigmas globose, subglobose or 
capitate, usually large. Fruits 4- to 8-valvular, 2- to 4-seeded, 
thin-walled; septum thin; seeds glabrous, smooth or punctate. 
Cotyledons ovate, obovate, ovate-cordate or orbicular, flat, fol- 
ded or somewhat corrugate. 

Type: Bonamia linearis (R. Br.) Hall. f. (as Breweria linearis 
R. Br. , 1810.) 

Tropical Australia and southern Africa. 

This section is very sharply defined from section Trichantha , 
but not very distinctly from section Bonamia , from which it differs 
by its smaller corolla, thin and mostly smaller leaves, smaller 
sepals, solitary flowers or simple cymes, and peduncles very short 
or absent. 

The circumscription of this section and section Bonamia may 
necessarily be modified when the morphology of the plants involved 
is kno\>m better. 

26. Bonamia linearis (R. Br.) Hall. f. 3ot. Jahrb. 16:530. 1893. 
Breweria linearis R. Br. Prodr. Fl. Nov. Holl. 488. 1310. 
Bonamia linearis (R. Br.) Hall. f. var. genuina Hall, f. 
Bull. Herb. Boiss. 5:1011. 1897. 

Perennial, herbaceous or suffrutescent vines. Stems prostrate 
or shortly twining, slender terete, becoming 3-9 dm. long, mostly 
1-2 mm. thick, soft-pilose or sparsely pilose, becoming almost 
glabrous in age; old stems 2-4 mm. thick; internodes 1-2.5 cm. 
long. Leaves shortly petiolate, soft, thin or submembranous , pi- 
lose with long and very fine hairs or becoming sparsely so or near- 
ly glabrous; petioles slender, 2-4 mm. long, pilose; blades linear 
or linear-lanceolate, 1.5-3.5 cm. long, 2-6 mm. broad, occasionally 
slightly broader, acute or attenuate, rarely rounded or subtruncate 
at the base, acute or acuminate at the apex; veins indistinct ex- 
cept thin midrib. Flowers axillary, solitary, shortly pedicellate 
or nearly sessile; peduncles absent; pedicels 0-3 mm., pilose; 
bracts small, linear or filiform, 1-3 mm. long. Sepals ovate-lanceo- 
late or ovate-acuminate, 5-7 mm. long, 2-3 mm. broad, herbaceous, 
subcoriaceous near the base, equal or subequal, pilose or sericeous 
outside, acuminate or acute at the apex. Corolla white, funnel- 
shaped, 1-1.5 cm. long, long-pilose on interplicae. Stamens includ- 
ed; filaments filiform, glabrous above, scattered-villous on the 



I960 



I.Iyint & Ward, Revision or Donamia 



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200 PHYTOLOGIA Vol. 17, no. 3 

lower parts adnato to corolla tubo; anthers oblong-oval, cordate 
at the base. Ovary oblong-oval, long-pilose; styles bifid to 
the middle or lower, filiform, glabrous; stigmas capitate or sub- 
globose-cap itato. Capsules ovate-apiculate, pilose at the apex, 
2- to 4-seeded, 4-valvular; seeds oblong-ellipsoid, glabrous, 
black or brown. Cotyledons orbicular. 

Type: Australia (Nov. Hollandia tropical), F. Bauer 318 , 
"1801-05" (W-isotype). 

Known from northern Queensland and islands of the Gulf of Car- 
pentaria, growing mostly on sandy grounds and coastal dunes (Map 9). 

This species has been collected in flower in April and June. 
Specimens with mature fruits were not available for this study. 

This species is closely related to B. media . Specimens (such 
as F.W. \'/hitehouse - BRI) with wider leaves are very similar to 
the typical variety of B. media and are vegetatively hardly distin- 
guishable from the latter. Hallier treated B. linearis and B. media 
as two distinct species in the first part of his synopsis, but later 
in the same paper he treated them as varieties of the same species. 
However, B. linearis seems to be distinct from B. media because 
of its larger corolla and generally narrower leaves (Figure 4). 

Specimens examined : 

AUSTRALIA: Northern Territory: On landward edge of coastal 
dune, Little Lagoon, Groote Eylandt, in the Gulf of Carpentaria, 
prostrate herb, corolla white, R.L. Specht 230 , April 13, 1948 
(L, US). Queensland: Gilvert River, Elgrey per N.A.R. Pollock 
(BRI); Doomadgee Mission, F.W. ^Vhitehouse (BRI). Location inde- 
finite: Stannary Hills, T.L. Bancroft , June, 1909 (BRI); Nova 
Hollandia tropica, Ferd. Bauer 318 (W). 

27. Bonamia oblongifolia Myint, Burma Jour. Life. Sci. 1:32. 1968. 

Perennial, herbaceous or suffrutescent, densely ferrugineous 
plants. Stems terete, erect or suberect, densely ferrugineous with 
brown hairs, 1.5-2 dm. tall, about 3 mm. thick at the base; branches 
1-1.5 mm. thick. Leaves shortly petiolate or subsessile, subcoria- 



Figure 4 

Variations in sizes and shapes of leaves in jB. media , 
brevifolia , B. oblongifolia and _B. linearis 



1-9 


B. 


media var. media x 2. 


10-13 


B. 


media var. villosa x 2. 


14-16 


B. 


brevifolia x 2. 


17-21 


B. 


oblongifolia x 2. 


22-26 


B. 


linearis x 2. 



1968 



Uiylnt & Ward, Revision of Donamia 



201 




In 



202 P H y T L G I A Vol. 17, no. 3 

ceous or leathery, densely ferrugineous on both surfaces; petioles 
1-2 mm. long or indistinct; blades oblong or rarely oblong-ellip- 
tic, 1-2 cm. long, 3-4 mm. wide, rounded at the base and apex; 
midrib slightly impressed above, distinct below; lateral veins in- 
visible (because of thick coating of hairs), 4-5 pairs. Flowers 
axillary, solitary, shortly pedicellate or almost sessile; pedun- 
cles absent; pedicels 1-1.5 mm. long, ferrugineous; bracts small, 
linear, 1-1.5 mm. long, frequently slightly exceeding pedicels in 
length. Sepals ovate, oblong-ovate, or ovate-acute, equal or slight- 
ly unequal, 3-4 mm. long, rarely slightly longer, coriaceous or 
subcoriaceous , acute or shortly acuminate at the apex, densely 
ferrugineous or sericeous with brownish hairs outside. Corolla 
blue, campanulate-funnelform, 6-8 mm. long, subentire or slightly 
lobulate at the margin, pilose on interplicae. Stamens included; 
filaments filiform, glabrous; anthers dorsifixed, oblong, 0.5-1 
mm. long. Ovary oblong with circular disc at the base, long- 
pilose near the apex; styles bifid to the middle, filiform, glab- 
rous; stigmas capitate. Capsules ovoid-conical, 5-6 mm. long, pi- 
lose at the apex, glabrous below, 2- to 4-seeded, 4-valvulcir; seeds 
oblong-ellipsoid, 2.5 mm. long, black or dark brown, glabrous. 
Cotyledons orbicular or orbicular-ovate. 

Type: Western Australia: Ville de Broome, dans le gazon sur 
le sable "rues" de Broome, alt. ca. 5-10 m. , herba, flos. bleue, 
B.P.G. Hochreutiner 2840 , 4. II. 1905 (G). 

Known only from the type location, growing on a sandy or gravelly 
meadow (Map 9). 

The collector mentioned that this plant was herbaceous, but 
gave no actual habit. From the material examined , it appears to 
be erect. His specimen has been erroneously referred to as B. 
pannosa , which is very distinct in possessing unequal sepals, 
long bracts , and broad leaves . 

This plant is undoubtedly related to B. media because of simi- 
larity of the indumentum and to _B. linearis because of narrow leaves 
(Figure 4). However, B. oblongifolia possesses a series of distinc- 
tive features which offers a sound basis for adjudging it a separate 
species. The stem and its branches in B. oblongifolia are short, 
stout and thick at the base, and possess short intemodes, while 
they are long, slender and weak in _B. media and B. linearis . Leaves 
in B. oblongifolia are oblong or oblong-elliptic, sessile or sub- 
sessile, and rounded at both ends. But leaves in B. media are 
ovate-lanceolate or ovate-subcordate, shortly petiolate, obtuse, 
truncate or subcordate at the base, and acute, obtuse or emar- 
ginate at the apex. Leaves in B. linearis are linear or linear- 
lanceolate, distinctly petiolate, attenuate, acute or obtuse at 
the base and acute at the apex. Sepals in _B. oblongifolia are 
smaller and acute or obtuse at the apex, while they are larger and 
mostly acuminate at the apex in B. linearis . 



1968 Vlylnt & TTard, Revision of Bonamia 203 

B. oblongifolia , as the name signifies, is characterized by 
oblong leaves. Other distinguishing features of this species are 
erect or suberect habit, short stems and branches, indumentum of 
dense bro\>ni hairs, smaller sepals, shorter corolla and filiform, 
glabrous stamens. 

28. Bonamia brevifolia (Benth.) Myint, Burma Jour. Life Sci. 1: 
33. 1968. 

Brewer ia brevifolia Benth. Fl. Austr. 4:436. 1869. 
Bonamia linearis (R. Br.) Hall. f. var. brevifolia (Benth.) 
Hall. f. Bull. Herb. Boiss. 5:1012. 1897. 

Perennial, herbaceous or suffrutescent vines, growing from 
thick and hard rootstock. Stems prostrate, slender, terete, 8-10 
dm. long, long-sericeous or pilose, with grey or silvery gre}' 
hairs, becoming less hairy in age; internodes 1-2.5 cm. long. 
Leaves shortly petiolate, soft, herbaceous or somewhat leathery, 
rarely subcoriaceous , with scattered long hairs or glabrous above, 
densely appressed-pilose with silvery grey or light brownish hairs 
below; petioles slender, 2-7 mm. long, pilose; blades ovate-cor- 
date, 1-1.7 cm. long, 9-14 mm. broad, cordate at the base, acute 
or acute-mucronate at the apex; veins impressed above, distinct 
below; lateral veins 3-5 pairs. Flowers axillary, solitary, short- 
ly pedicellate; peduncles very short or absent; pedicels 2-4 mm. 
long, sericeous or pilose; bracts linear or narrowly subulate, as 
long as pedicels or longer, mostly 3-4 mm. long. Sepals lanceo- 
late or ovate-lanceolate, 3.5-5 mm. long, 2-3 mm. broad, herbaceous 
or subcoriaceous, equal or slightly unequal, long-pilose or long- 
sericeous outside, acuminate or acute at the apex. Corolla blue, 
funnelform or shortly tubular-campanulate , 8-11 mm, long, pilose 
on the interplicae. Stamens included, filaments filiform, sparsely 
villous at the base or nearly glabrous; anthers oblong-ovate, cor- 
date at the base. Ovary ovoid-conical, long-pilose or sparsely 
long-pilose or nearly glabrous; styles connate for lower one-fourth, 
readily separable nearly to the base, filiform, glabrous; stylar 
branches unequal; stigmas globose, rarely subglobose. Capsules not 
available for study. 

Type: Australia, Port Essington, Armstrong (BM-isotypel ) . 

Known only from northern districts of Northern Territory, Aus- 
tralia (Map 9). 

Collectors have given no information about the habitat of the 
plant. It has been collected in flower in February. 

Hallier (1897), in the first part of his synopsis, treated this 
species as belonging to Bonamia media, but later both B. media and 
B. brevifolia were treated as two different varieties of B. linearis . 
_B. brevifolia is only distantly related to this latter species 
because of its shorter, broader and cordate leaves (Figure 4), 
longer bracts, smaller sepals, blue corolla and sparsely pilose 
ovary, whereas B. linearis possesses long, narrow and linear or 



20li P H Y T L G I A Vol. 17, no. 3 

linoar-lanceolatn loaves, minuto bracts, longf;r sf-paJs, white 
corolla and densely pilose ovary. H. brevi folia seems to be 
more closely related to B. mod i a , from which it differs by its 
distinctly cordate and acutf leaves fvndest near the base), long- 
er bracts, smaller sepals, sparsely villous filaments and sparse- 
ly pilose ovary. These characteristics, in addition to the dif- 
ferences in the general appearance of plant, length of stem and 
indumentum, appear to offer a sound basis for treating it as a 
distinct species. 

Specimens examined: 

AUSTRALIA: Northern Territory: Port Essington, A?-nstrong , 
1840 (BM); Humpty Doo, prostrate, leaves dark green above, light 
green below, flowers blue, U.S. McKee 8328 , February 10, 1961 (DRI). 

29. Bonamia media (R. Br.) Hall. f. Bot. Jahrb. 16:528. 1893. 
Breworia media R. Br. Prodr. Fl. Nov. Holl. 488. 1810. 
Bonamia linearis (R. Br.) Hall. f. var. media CR. Br.) Hall, 
f. Bull. Herb. Boiss. 5:1011. 1898. 

Perennial, herbaceous or suffrutescent vines. Stems prostrate, 
occasionally procumbent or suberect, slender or stout, terete, be- 
coming 2-15 dm. long, rarely longer, mostly 1-3 mm. thick, soft- 
sericeous with silvery grey or light bro^•mish hairs and becoming 
less sericeous in age or nearly glabrous; internodes 1-2 (-3) cm. 
long. Leaves shortly petiolate, soft and thin to subcoriaceous , 
sericeous, villous or ferrugineous or nearly glabrous or glabrate: 
petioles 2-7 (-11) mm. long, slender; blades highly variable, ovate, 
elliptic-ovate, ovate-subcordate, oblong-ovate, ovate-lanceolate, 
ovate-emarginate or ovate-cordate, mostly 1-2.5 cm. long, 6-15 mm. 
broad, usually with length-width ratio of 2 or less, truncate, sub- 
cordate or obtuse at the base; obtuse, abruptly acute, truncate, 
obtuse-mucronate or slightly emarginate at the apex; veins impressed 
above and distinct beneath to scarcely perceptible because of dense 
coating of hairs; lateral veins 3-5 pairs. Flowers shortly pedi- 
cellate or nearly sessile, axillary, solitary or occasionally in 
simple cjmies of two or three; peduncles very short or absent; pedi- 
cels 1-5 mm. long, sericeous; bracts small, linear, 1-2 mm. long. 
Sepals ovate-lanceolate or ovate-elliptic, mostly 5-7 mm. long, 
2.5-3.5 mm. broad, herbaceous, subcoriaceous near the base, equal 
or slightly unequal, sericeous, densely sericeous or villous out- 
side, acute or acuminate at the apex. Corolla blue, light blue, 
or often white, shortly tubular-campanulate or funnelform, 8-15 
mm. long, pilose on interplicae. Stamens included; filaments fili- 
form, glabrous; anthers oblong or oblong-oval, cordate at the base. 
Ovary ovoid, long-pilose near the apex, with scattered hairs or 
glabrous below; styles bifid to the middle or lower, filiform, 
glabrous; stigmas subglobose-capitate. Capsules ovoid-apiculate, 
pilose at the apex, glabrous or rarely with scattered, minute 
hairs below; seeds brown or black. Cotyledons orbicular or ovate- 
cordate, folded. 

Type: Australia (Nova Hollandia tropical), F. Bauer 321 , "1801- 
05" (W-isotype). 



1968 Ityint Sc Ward, Revision of Bonamia 20^ 

Red sand, yellow sand, sandy loam, lateritic or nonlateritic 
soils in open Eucalyptus forest, dry banks, gullies, timbered flats 
or desert from Northern Territory to Queensland and New South Wales 
(Map 10). 

This is the most wide-ranging species in Australia. It has 
been collected in flower in January, February, March and October, 
and in fruit in January, March, June and December. One collector 
noted the flowering period from summer to winter. 

This species has been treated as a variety of Bonamia linearis 
by Hallier, who treated it as a distinct species in his earlier 
work. However, B. media is distinct, characterized by elliptic, 
ovate-elliptic, ovate-subcordate or ovate-cordate leaves, mostly 
obtuse at the apex. 

Although Hallier (1897) has pointed out the similarity of leaf 
tissue of B. media with those of B. spectabilis and B. sericea , 
it is most closely related to B. linearis and B. brevifolia . 

This species is quite variable in leaf shape, size and pubes- 
cence (Figure 4). It is clearly separable into three varieties. 

29a. Bonamia media (R. Br.) Hall. f. var. media. 

Bonamia linearis (R. Br.) Hall. f. var. media (R. Br.) 
Hall. f. Bull. Herb. Boiss. 5:1011. 1897. 

Stems long, slender, finely soft-sericeous or pilose or oc- 
casionally sparsely so, rarely dense-sericeous. Leaves sericeous, 
pilose, with scattered hairs or almost glabrous on the upper sur- 
face, sparsely pilose, sericeous or rarely densely sericeous be- 
low, with clearly visible lateral veins, obtuse or obtuse-mucronate 
at the apex. 

This variety is inconsistent in several features and future 
studies may find it to be composed of more than one variety. Since 
the material available at present is scanty, it is treated here as 
a large, polymorphic group. The following specimens are worth 
describing briefly to show the variations within this variety: 

Bauer 321 (W) : Leaves narrower, with length-width ratio ex- 
ceeding 2, glabrous on the upper surface, sparsely long-pilose on 
the lower surface. 

Collector unknown; " 129 " Settlement Ck. , N.T. (BRI): Leaves 
larger, with leaf blades as long as 3 cm. , glabrous on the upper 
surface, sparsely long-pilose or glabrescent on the lower surface; 
bracts 2-3 mm. long. 

R.A. Perry 3433 (BRI, US): Leaves very small, 14 mm. or short- 
er and 9 mm. or narrower (thus erroneously annotated as B^ brevi- 
folia) , sericeous on both surfaces. 

S.L. Everist 2903 (BRI): Numerous slender stems from a sin- 
gle rootstock; leaves with longer petioles, somewhat attenuate at 
the base, truncate or slightly emarginate at the apex; upper flow- 
ers in cluster of two or three. 



206 



PHYTOLOGIA 



Vol. 17, no. 3 




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1968 I'iyxnt & Ward, Revision of Bonamia 207 

James Keys 637 (BRI): Slender and frequently branching stems; 
leaves smaller, dense long-sericeous or densely pilose on the lower 
surface. 

M.S. Clemens , October 16, 1945 (BRI, L) : Leaves densely seri- 
ceous ; corolla blue. 

S.L. Everist 1910 (GH) : Stems and leaves densely sericeous 

thus somewhat intermediate between the two varieties, media and 
villosa . 

Specimens examined : 

AUSTRALIA: Northern Territory: 31 mi. north of Devil's Mar- 
bles, prostrate herb, white flowers, rare in red sand, G. Chippen- 
dale 949 , 8. 3. 1955 (BRI); 21 mi. south of Elliott, prostrate 
spreading herb, flowers white, common in light red sand, Chippen- 
dale 1024 , 9. 3. 1955 (BRI); 6 mi. north of Katherine, in limes- 
tone pavement country with red soil, prostrate, leaves g^ey green, 
flowers white, H.S. McKee 8515 , February 17, 1961 (BRI); 15 mi. 
north of Victoria River Down Station, creeping greyish plant common 
on skeletal soils on cherts with E. brevifolia and Plectrachne sp. , 
R.A. Perry 2111 , 10. 6. 1949 (BRI, US); 30 mi. south -southwest of 
Wavehill Station, common near edge of truncated lateritic desert, 
prostrate creeping plant with runners several feet long. Perry 
2217, 21. 6. 1949 (BRI, US); 20 mi. northwest of Ooratippa Station, 
prostrate, grey plant, trailing for several feet with white flowers, 
common on red tertiary nonlateritic soil with Eucalyptus gamophylla , 
Perry 3433 , 14. 3. 1953 (BRI, US). Unknown collector: Settlement 
Ck. , 129, February, 1922 (BRI). QUEENSLAND: Sandy place, race 
course, Charleville, Warrego Dist. , M.S. Clemens , October 6, 1945 
(G) ; Cemetry, Charleville, Warrego Dist., flower blue, Clemens , 
October 16, 1945 (BRI, L) ; Yalleroi-Jericho and vicinity, Mitchell 
Dist., stems prostrate, Clemens , April 1, 1946 (F, UC) ; Nive River, 
about 30 mi. north of Augathella, prostrate plant, common in yellow 
sand, leaves silvery, silky tomentose, flowers white, S.L. Everist 
1910, October, 1939 (GH) ; Boatman Station, Maranoa Dist., in red 
sandy soils, herb with many slender prostrate stems, radiating 
from woody rootstock, flowers white, Everist 2903 , 24. 3. 1947 (BRI); 
"Curragh" Station near Cunnamulla, around bore in paddock in 
brown loam, prostrate herb, greyish green leaves, white corolla, 
alt. 620 ft., C.E. Hubbard and C.W. Windero 6220 , 4. 1. 1931 (BRI); 
Ad el's Grove, via Camooweal, trailing perennial herb, stems to 6 
ft. long, fls. white, summer to winter, dry banks, gullies and tim- 
bered flats, A. De Lestang 162 , 20. 1. 1946 (BRI); Ayr. Rev. N . 
Michael 1522 (BRI); Gilbert River, N.A.R. Pollock (US). Torrens 
Creek, common in sandy soil, open Eucalyptus forest, fls. white, 
C.T. White 8931 , 18. 3. 1933 (BRI, US); Carbean near Cunnamulla, 
Warrego Dist. ; numerous prostrate stems from a long taproot, flow- 
ers white. White 12014 , 26. 3. 1941 (A, BRI); Doomadgee Mission, 
W. Whitehouse (BRI); Charleville, " J.F.B ." March, 98 (BRI). Ix)- 
cations indefinite: Nova Hollandia tropica, Ferd. Bauer 321 (W) ; 
"Bustarst Hern, James Keys 637 " (BRI). 

29b. Bonamia media (R. Br.) Hall. f. var. villosa (Benth.) Myint, 
Burma Jour. Life Sci. 1:33. 1948. 



208 PHYTOLOGIA Vol. 17, no. 3 

Hrnworia mc(\ i a H. Mr. var. villosa iJonth. 11. Austr. 4: 

436. 1869. 
Honamia linfaris var. mod la subvar. villosa Olenth.) 

Mall. f. Dull. Herb. Boiss. 5:1011. 1897. 

Differs from the typical variety by densely sericeous and stouter 
stems frequently branching; densely sericeous and thicker leaves, 
truncate or slightly emarginate at the apex; the hairs on all parts 
turning to brown on drying, thus appearing to be f errugineous ; 
lateral veins indistinct. 

Type: Australia: Victoria River, F. Mueller (presumably at 
BM, not seen). 

Known from Northern Territory and New South Wales , growing on 
red sandy soil (Map 10). 

Tnis variety was described by Bentham (1869) , who questioned 
its validity, and later was treated as a subvariety by Hallier 
(1897). However, it appears to be a very distinct variety because 
of the dense coating of hairs over the entire plant. Future study 
of more materials may modify the circumscription of this variety. 

Specimen examined : 

AUSTRALIA: New South Wales: 30 mi. west of Uantabulla, red 
sandy soil, N.C.W. Beadle 36308 , 1. XII. 1944 (US). 

29c. Bo namia med ia (R. Br.) Hall. f. var. emarginata Myint & Ward, 
var. nov. 

Differt a var. media et var. villosa foliis apice emarginato, 
basi distincte cordata vel subcordata, sparse pilosis, 1.2plo 
longioribus quam latioribus vel paulo; venis lateralibus plerumque 
3 binis ; bractis 1.5-2.5 mm. longis ; stigmatibus depresso-capi- 
tatis vel peltatis. 

Differs from var. media and var. villosa by leaves emarginate 
at the apex, distinctly cordate or subcordate at the base, sparsely 
pilose, with length-width ratio of 1.2 or less; lateral veins most- 
ly 3 pairs; bracts 1.5-2.5 mm. long; stigmas depressed-capitate 
or peltate. 

Type: Australia: Queensland: Gladstone, unknown collector 
(BRI). 

Known only by a single collection from the east coast of Queens- 
land, Australia (Map 10). 

The collector gives no information on the habit, flower color 
and habitat of this variety. It is quite distinct from the other 
two varieties of the species and it may be found to be a separate 
species in future studies, when more specimens of better condition 
become available. Because the only available specimen is f ragmen- 



1968 L^int & V/ard, Revision of Eonamia 20? 

tary, it is treated here as a variety of _B. media . 

This variety is more closely related to var. media than to var. 
villosa because of its indumentum, and smaller leaves. 

30. Bonamia rosea (F. v. Muell. ) Hall. f. Bot. Jahrb. 16:528. 1893. 
Breweria rosea F. v. Muell. Fragon. Phytogr. Aust. 1:233. 
1839. 

Perennial, erect subshrubs or undershrubs of 3-9 (10) dm. tall. 
Roots thick, woody, deep-penetrating. Stems terete, densely tomen- 
tose, hirsute or ferrugineous with grey or brovmish hairs; main 
stems about 2-5 mm. thick, with pulpy bark near the base, readily 
branching, thus having numerous culms; older stems becoming less 
tomentose. Leaves shortly petiolate or frequently subs ess ile, 
thick, leathery, subcoriaceous or coriaceous, densely ferrugineous 
or hirsute on both upper and lower surface; petioles 1-5 mm. 
long or almost absent, 1-1.5 mm. thick, densely ferrugineous or 
hirsute; blades orbicular, ovate or obovate, mostly 7-15 mm. long, 
about 6-13 mm. wide, with length-width ratio of mostly one, entire 
at the margin, rounded or slightly cordate at the base, truncate, 
slightly emarginate, obtuse or obtuse-mucronate at the apex, with 
about 3-5 pairs of indistinct lateral veins. Flowers, axillary, 
sessile or shortly pedunculate, solitary or occasionally in cymes 
of two or three, frequently aggregated near the terminal ends of 
branches; peduncles, when present, up to 15 mm. long, mostly 1 mm. 
thick, densely ferrugineous; bracts linear or linear-lanceolate, as 
long as 6 mm. or very small and inconspicuous. Sepals ovate-lanceo- 
late, 5-9 mm. long, acute or abruptly acute, rarely obtuse, densely 
long-hirsute outside, the inner ones smaller or narrower. Corolla 
pink or white, tubular-campanulate, or broadly urceolate, 1-1.8 cm. 
long, with limb of 8-15 mm. broad, long-hirsute on interplicae, 
glabrous on plicae, 5-10-lobulate at the margin; tube broad, cylin- 
drical and distinct, stamens included; filaments adnate and hairy 
at the base, free and glabrous above; anthers broadly oblong. Ovary 
hirsute with long hairs near the apex, glabrous below, conical; 
styles bifid above two-thirds or lower, filiform, glabrous except 
near the base; stylar branches unequal; stigmas large globose. Cap- 
sule valvular, 2- to 4- seeded, conical, hirsute at the apex. Seeds 
ovate or ovate-oblong, glabrous. Cotyledons ovate or ovate-orbicu- 
lar. 

Type: West Australia; type specimen not available. 

Western and central Australia from Nichol Bay and Dampier Archi- 
pelago south to Lake Moore and east to southern districts of the 
Northern Territory (Map 11). 

Collectors note "coarse sandy desert or grassland," "spin- 
ifex sand plain," "bushes on sands," "deep red sands in area of 
burnt Triodia pungens " and "sand heath" as the habitat of this 
species. 



210 



PHYTOLOGIA 



Vol. 17, no. 




® * 



1968 ^ylnt & Ward, Revision of Eonamia 211 

Specimens recently collected from central Australia by Chippen- 
dale and Lazarides are distinct in certain morphological features, 
particularly in minute or inconspicuous bracts, smaller corolla, 
completely sessile flowers and numerous stems from a single shoot. 
These morphological characteristics (especially the minute vs. long 
bracts and small vs. large corolla) seem to support the supposi- 
tion that the central Australian plants deserve a distinct taxonomic 
status, at least at the varietal level. But, with just a handful 
of material available at this time, it is not described here. 

The leaves vary relatively little (Figure 5). 

Specimens examined : 

AUSTRALIA: Northern Territory: Near Ulambaura Spring, Haast 
Bluff, subshrub 1 ft. , infl. white, infrequent in Triodia pungens 
assoc. , G. Chippendale 2568 , 23. 8. 1956 (BRI); 31. 6 m. north- 
west of Mt. Patricia, grey perennial herb 1 ft. , common in small 
area in deep red sand, in area of burnt Triodia pungens , Chippen- 
dale 4297 , 5. 5. 1958 (BRI); 65 m. northwest of Willowra H.S., 
dwarf shrub 1 ft., buds brown green, common in deep red sand, in 
area of burnt Triodia pungens , Chippendale 4792 , 31. 7. 1958 (BRI); 
59 mi. northwest of Mt. Doreen Station, dominant in patches in 
coarse sandy desert dominated by Plectrachne schinzii grassland, 
low hairy grey subshrub to 12 in. high and as wide, flowers white, 
culms numerous, branching and spreading, M. Lazarides 6020 , 17. 9. 
1957 (BRI, US). Western Australia: Dampiers Archipelago, B.F . 
von Mueller (BM) ; Greenoughs River, Mueller (GH, US); Nichol Bay, 
Mueller ( BM) ; Murchison River, Oldfield (W) ; in fruticetis arenosis 
inter flumina Moore et Murchison, E. Pritzel 606 , IX, 1901 (HBG, 
W), 616, IX, 1901 (A, BM, GH , L, MO, US); northeast of Melrose, 
N.H. Speck 1388 , 8. 9. 1958 (MO); 13 mi. northwest of Albion Downs, 
woolshed, Eremean Province, spinifex sandplain. Speck 1477 , 17. 
9. 1958 (BRI, MO). 

31. Bonamia pannosa (R. Br.) Hall. f. Bot. Jahrb. 16:530. 1893. 
Breweria pannosa R. Br. Prodr. 488. 1810. 

Prostrate or twining vines growing from perennial rootstock. 
Stems terete, soft and herbaceous while young, soon becoming woody, 
mostly 2-3 mm. thick, densely hirsute with soft ferrugineous or 
silky hairs. Leaves shortly petiolate, herbaceous or soft-coriaceous, 
densely hirsute on both upper and lower surface; petioles 3-20 mm. 
long, densely hirsute; blade ovate or orbicular, rarely ovate-oblong, 
2-4 cm. long, mostly (1-) 1.2-3 cm. broad, truncate, subcordate or 
rounded at the base (lower leaves sometimes attenuate at the base), 
obtuse, rounded or obtuse-acute at the apex. Flowers axillary, 
solitary or in cymes of two to few flowers, rarely several flowers 
(forming dense cluster), sessile or shortly pedunculate and/or 
shortly pedicellate; bracts subulate 5-10 mm., becoming slightly 
longer in age, hirsute. Sepals soft and thick or subcoriaceous , 
unequal; outer two sepals large, ovate, 8-11 mm. long, 7-8 mm. 
broad, hirsute on the back, hirsute inside except glabrous center, 
acute at the apex; in-out sepal (third sepal) oblique-ovate, as 



212 PHYTOLOGIA Vol. 17, no. 3 

l(jng as outer two sfpal.s, 4-6 rnrn. broad, hirsute as outr-r sopals ; 
innpr two sepals smallfr, ovate-acuminate or broadly lanceolate, 
6-7 mm. long, 3-4 mm. broad, hirsute outside, glabrous inside, 
acuminata at the apex. Corolla blue or violet blue, rarely white, 
funnel-shaped, 1.2-1.5 cm. long, with spreading and sublobulate or 
nearly entire limb, hirsute or pilose on interplicae, glabrous on 
plicae. Stamens inserted; filaments villous, slightly broadened 
at the base; anthers oblong, 1-1.5 mm. long, dors i fixed, cordate 
at the base. Ovary ovoid-conical, long-hirsute at the apex; styles 
bifid to the middle, v/ith unequal stylar branches; stigmas globose- 
capitate. Capsules thin-walled, 4- to 8-valved, 4-seeded, ovoid, 
5-6 mm. long, 4-5 mm. in diameter, hirsute at the apex, glabrous 
bp]ow; seeds glabrous, ovate-triangular, 2-3 mm. long. Cotyledons 
ovate-cordate, rounded or slightly emarginate at the apex; 
cotyledonary petioles fused. 

Type: (Nova Hollandia tropica) Tropical Australia, Bauer 325 
( BM-lectotype? ; W-isotypel ) 

Sandy soil, gravelly sand, limestone, shale and dry ridges, 
rarely in wet ground in tropical regions of Australia from Queensland 
to the northeastern districts of western Australia (Map 11). 

This species has been collected in flower from February to 
June and in fruit from April to July. The isotype at Vienna has 
been mislabelled as Pol3mieria lunata presumably by R. Brown. 

Specimens examined : 

AUSTRALIA: Northern Territory: 28 mi. south of Elliott, 
prostrate spreading herbs, corolla blue, common on gravelly sand, 
C. Chippendale 1021 , 9. 3. 1955 (BRI); Spring Vale, Port Darwin, 
Alfred Giles (BRI); 5 mi. from Katherine, on Wyndham Road in wet 
ground, prostrate, leaves pale green, flowers bright blue, H.S. 
McKee 8536 , February 18, 1961 (BRI); 12 mi. southwest of Katherine 
Township, prostrate grey bush several feet long, common on sandy 
soil with E. miniata open forest, R. A. Perry 1978 , 2. 6. 1949 
(BRI, US); 6 mi. south of Limbunya Station, prostrate plant with 
grey foliage, common on limestone outcrop, Perry 2337 , 4. 7. 1949 
(BRI, US); Groote Eylandt, S.H. Wilkin 98 , February, 1929 (BM). 
Queensland: Ad el's Grove via Camooweal, trailing plant, young stems 
erect to 9 in. high, flower dark blue, velvety, rather pretty, grows 
on dry ridges, A. De Lestang 5 (BRI); Mt. Isa, Burke District, 



Figure 5 

Variations in sizes and shapes of leaves in 
pannosa , JB. dietrichiana and B. rosea 

1-5 B. pannosa x 2. 

6-8 B. dietrichiana x 2. 

9-13 B. rosea x 2. 



1963 



Hyint &C. Ward, Revision of Bonamia 



213 




2m PHYTOLOGIA Vol. 17, no. 3 

Mrs. M. Morris , May, 1952 CUKI). Westom Australia: Slatey Creek, 
16 mi. northwest of Glenroy Moatworks , prostrato, grey green, hairy 
plant with runners severa] feet long and blue flowers, common in 
shaDey bed of creek, M. L,izarides 5155 , 22. 4. 1955 CBRI). Location 
indefinite: Nova Ho]landia tropica, l^uer 325 fW-isotype). 

32. Bonamia velutina Verdcourt, Kirkia 1:27, tab. III. 1961. 

Perennial, erect or suberect subshrubs. Stems woody or becoming 
woody, terete, tomentose, or densely appressed-sericeous , as high 
as 9 dm., 1-3.5 mm. thick, frequently branching; intemodes 5-25 
mm. long. Leaves shortly petiolate, or subsessile, soft and her- 
baceous or leathery, rarely subcoriaceous , densely velutinous with 
silvery grey or brownish hairs on both surfaces; petioles 1.5-4 mm. 
long, densely velutinous; blades elliptic or elliptic-oblong, 1.2- 
4.5 (-6) cm. long, 4-15 mm. broad, upper leaves smaller and lower 
leaves somewhat larger, rounded or subtruncate at the base; acute- 
mucronate or rare]y obtuse-mucronate at the apex; veins impressed 
above, prominent underneath; lateral veins 4-6 pairs. Flowers 
shortly pedunculate or sessile, axillary, solitary or in cymes of 
two or three; peduncles 0-3.5 (-6) mm. long; pedicels 0-1.5 mm. 
long; bracts linear, 2-4 mm. long, velutinous. Sepals ovate- 
lanceolate, obovate or spathulate, subcoriaceous at the base, folia- 
ceous at the apex, slightly unequal; outer three spathulate with 
oblong base as long as 3 mm. and 1.5 mm. wide, v;ith dilated apex 
3.5-4 mm. long and 2.5-3.5 mm. wide, densely velutinous outside; 
inner two ovate-lanceolate, 6.5 mm. long, 2.5 mm. wide, acute at 
the apex, not dilated, velutinous outside. Corolla white, infundi- 
buliform, 9-12 mm. long, 13 mm. wide at the apex, slightly 5- 
lobulate, densely pilose on interplicae. Stamens inserted; fila- 
ments filiform, dilated at the base, glabrous; anthers 1 mm. long, 
cordate at the base. Ovary ovoid, densely long-pilose; styles bifid, 
shortly connate (1-2 mm.) at the base, glabrous; stigmas lobulate- 
peltate. Capsules ellipsoid or globose-ellipsoid, 6 mm. long, suba- 
cute, minutely appressed-pilose, 2- to 4-seeded, 4-valvular; seeds 
angular-ellipsoid, 2-3 mm. long, glabrous, minutely punctate. Coty- 
ledons orbicular or ovate-orbicular; cotyledonary petioles short 
or absent. 

Type: Southern Rhodesia: Nuanetsi District, 0.4 km. within 
Southern Rhodesian border opposite Malvernia, in Guibourtia - 
Mopano Woodland on Umkondo sands , 450 m. , K. Wild 4688 , November 
1, 1955 (K-holotype, SRGH-isotype-not seen). 

Known only from Southern Rhodesia, southeastern border in 
Nuanetsi District (Map 1). 

According to the collectors this is an erect herb or subshrub 
in mopane or mixed woodland on sandstone, Umkondo sand or sandstone 
plateau. It has been collected in flower in November and in fruit 
in April. 



1968 Ityint & Ward, Revision of Eonamia 21^ 

This species was originally annotated as Seddera sp. , presum- 
ably because of its smaller flowers and shrubby habit. Although 
this species possesses certain morphological features and the habit 
of that genus, it is definitely a species of Bonamia as pointed out 
by Verdcourt. It seems to be more closely related to some Austra- 
lian species of that genus than to the other African species , be- 
cause of its erect habit, shorter stems, sessile or very shortly 
pedunculate flowers (mostly solitary or in simple cymes), smaller 
sepals and shorter corolla. 

Specimens examined : 

SOUTHERN RHODESIA: Nuanetsi District: Combretum Mopane, sand- 
stone, 1650 ft., herb 2-3 ft., R. Davis 1629 , November, 1955 (EA- 
paratype) ; Clarendon Cliffs, mixed woodland on sandstone plateau, 
erect 2 ft. , perennial, corolla white, R.B. Drummond 7809 , April 
29, 1962 (EA). 

III. Section: Trichantha Myint, Burma Jour. Life Sci. 1:34, 1968. 

Trichantha Karst. et Triana, Limnaea 28:437. 1856, 

not Trichantha Hooker, Icon. PI. tt. 666, 667. 
1844. 

Stems woody, twining or scandent, usually long and high-climb- 
ing, rarely short and suberect or erect, mostly 3 mm. or thicker. 
Leaves distinctly petiolate, often long-petiolate, soft, herbaceous, 
subcoriaceous or leathery, not membranous; blades mostly ovate, 
ovate-acuminate, or ovate-subcordate, rarely broadly elliptic, 
usually large, 3.5 cm. or longer, 2.5 cm. or wider, with length- 
width ratio of 1-1.5, rarely slightly higher, rounded, truncate, 
subcordate or cordate at the base; acute, obtuse, acuminate, round- 
ed or emarginate and distinctly mucronate at the apex. Inflores- 
cences axillary compound or simple cjmies of few to numerous flowers 
or terminal panicles, rarely uniflorous, pedunculate or sessile; 
bracts small, never foliaceous , deciduous or persistent. Sepals 
coriaceous, rarely subcoriaceous, mostly equal or slightly unequal, 
ovate, oblong-ovate or orbicular, obtuse, rounded or emarginate, 
rarely acute at the apex, mostly 3-6 mm. long. Corolla white, 
yellowish white, yellow or purple, 2-2.5 cm. long, rarely slightly 
shorter, subentire or 5- to 10-lobulate at the margin. Stamens 
included; filaments mostly villous or sparsely villous below, glab- 
rous above; anthers 2 mm. or longer, sagittate or cordate at the 
base. Ovary conical, ovoid-conical or ovoid-oblong, pilose, sparse- 
ly pilose or glabrous; styles free nearly to the base, fused to the 
middle or slightly higher; vascular traces branched into two in 
the upper part of stylar branches ; stigmas reniform or obscurely 
bilobed , large. Fruits mostly 4- or rarely 2-valvular, thick- 
walled, (0.5-1 mm. or thicker); septum hard, not membranous. Seeds 
densely villous or woolly, with long, soft hairs; hairs 1-3 mm. 
or shorter on the dorsal and ventral surfaces , 5 mm. or longer 
along the edges of seeds. Cotyledons ovate, ovate-cordate or obo- 
vate, corrugate or multiplicate when mature; non-corrugate or 
flat when young. 



216 P H Y T L G I A Vol. 17, no. 3 

Type: Dcjnamia trlchanthn Ha]], f. Tas Trichantha f^-rru^infa 
Karst. & Triana, 1856). 

Tropical Amprica from Panama to Brazil and Paraguay. 

This section is very distinct from the other two sections and 
is characterized by ligneous fruits, villous seeds, a branched 
vascular strand in each stylar branch, reniform or obscurely bi- 
lobed stigmas, and orbicular or ovate-orbicular sepals. 

33. Bonamia trichantha Hall, f . , Bot, Jahrb. 16:528. 1893. 

Trichantha ferruginea Karst. & Triana, Linnaea 28:438. 

1856. 
Brcweria mollis Pittier, Jour. Wash. Acad. Sci. 17:284. 

1Q27. 
Brpweria longipaniculata Pittier, Jour. Wash. Acad. Sci. 

17:284. 1927. 

Perennial, woody climber, apparently growing all year around. 
Stems twining, terete, smooth or warty, tomentose while young and 
becoming glabrous in age; about 1 cm. in diameter, becoming 10 m. 
long or longer. Leaves petiolate, coriaceous or submembranous, 
glabrous or sparsely pubescent above and densely tomentose or glab- 
rate on the lower surface, more densely so on the veins (especially 
in the glabrate forms); petioles 1-3 cm. long, 1-2 mm. thick, round 
or canaliculate above, minutely appressed pubescent or becoming 
glabrous; blades ovate, oblong or ovate-oblong, entire at the margin, 
obtuse or cordate at the base and acute, obtuse, attenuate or obtuse- 
mucronate at the apex; nerves slightly impressed above, prominent 
below; about 5-8 pairs of lateral veins. Inflorescences axillary 
cjmies of few to many flowers or terminal pseudopanicles ; cymes pedun- 
culate; peduncles and pedicels minutely appressed pubescent or be- 
coming glabrous; bracts small or inconspicuous. Sepals orbicular 
or orbicular-ovate, subequal or slightly unequal^ outer two fer- 
rugineous or pubescent-glabrescent ; inner ones sparsely pubescent- 
glabrescent, with hyaline margins, 5-7 mm. long, 4-5 mm. wide. 
Corolla white, 1.2-2 cm. long, about 1-1.5 cm. wide, tubular-cam- 
panulate, entire; outside surface with long hairs on interplicae, 
glabrous on plicae; tube short, about 3-5 mm. Stamens inserted, 
epipetalous ; filaments glabrous or villous, incurved or straight; 
anthers oblong or ovate, dorsifixed, emarginate or slightly cordate 
at the base, narrow at the apex. Ovary ovoid, with circular disc 
at the base, glabrous or villous at the apex; styles shortly connate 
at the base, glabrous or with scattered hairs; stigmas globose or 
capitate, mostly sub-bilobed. Fruit capsule, valvular with thick 
and ligneous wall, dehiscing mostly into two or four valves; parti- 
tion wall thin, coriaceous. Seeds ovoid, piano-compressed, densely 
villous, with longer marginal hairs. Cotyledons obovate or orbi- 
cular, emarginate at the apex; cotyledonary petioles fused. 

Type: Colombia: Magdalena: Piedras, Vallee du Magdalena, 
Nouvelle-Grande, prov. de Mariquita, J. Triana 2146 , 1851-1857 
(G-lectotype, BM, W-isotypes). 



1968 



^yint & V/ard, Revision of Bonamia 



217 






CO 




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CO 


TO 


U 


+J 


X> 


CD 


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iH 


O 


oq 



•X- 



218 P H Y T L G I A Vol. 17, no. 3 

I'rom sea level to 600 m. in I'anama , Colombia and Venezuela 

(Map 12). 

The collectors recorded this species as occurring in light 
forests on rocky hills, borders of forests, thickets on open rocky 
slopes or hill sides, river banks, arid bushy slopes and valleys. 
This species is separable, though not very clearly, into three 
varieties of which one may be divided into two forms. 

33a. Bo nam in trichantha Mall. f. var. trichantha . 

Bonamia trichantha Hal], f. var. typica v. Ooststroom, 
F^ec. Trav. Bot. Neerlandais 33:213. 1936. 

This variety is characterized by broadly elliptic leaves, ob- 
tuse, rounded or slightly emarginate at the apex and densely fer- 
rugineous underneath. (Figure 6). 

Restricted to Colombia, from Magdalena south into Tolima and 
Huila departments (Map 12). 

Specimens examined: 

COLOMBIA: Huila: Natagaima, rocky hill at gorge above N. 
Altitude 450-500 m. , light forest, shrubby vine, H.H. Rusby and F.W . 
Pennell 1159 , August 12, 1917 (GH). Magdalena: Piedras, Vallee 
du Magdalena, Nouvelle-Grande, prov. de Mariquita, 500 m. J. Triana 
2146, 1851-1857 (BM, G, W) ; Nouvelle-Grande, prov. de Jequendema, 
Triana 3801 , 1853 (BM). Tolima: E.P. Arbelaez 2173 , XII. 1932 (US). 

33b. Bonamia trichantha Hall. f. var. oblonga v. Ooststroom, Rec. 
Trav. Bot. Neerlandais 33:213. 1936. 

This variety is characterized by its oblong or oblong-lanceo- 
late leaves, mostly 5-8 cm. long and 2-2.5 cm. wide (Figure 6), 
and loose cymes of fewer flowers which usually are in terminal 
panicles. 

Type: Colombia: Santa Marta, Herbert H. Smith 1871 , 1898-1899 
(US-holotype, F, GH-isotypes). 

This variety is apparently endemic to northern Colombia and is 
known only from the type collection (Map 12). Its habitat is not 
known and the collector gives no more than the location. A des- 
cription of fruit and seed has to await future collections. 

33c. Bonamia trichantha Hall. f. var. ovata v. Ooststroom, Rec. 
Trav. Bot. Neerlandais 33:213. 1936. 

This variety is characterized by its ovate or ovate-acuminate 
leaves, rarely emarginate at the apex, and much varied in size and 
pubescence (Figure 6). 

Type: Colombia: Tolima: Honda, open rocky slope, alt. 300- 
400 m. , F.W. Pennell 3575 , January 3-4, 1918 (US-holotype, GH- 
isotype) . 



1968 



Lfcrint & Yifard, Revision of Donamia 



219 




Leaf shapes and sizes in Bonamia trichantha 



220 P H Y T L G I A Vol. 17, no. 3 

This variety ranges from I'anama and Colombia east into eastern 
Venezuela. It is separable into two forms. 

li. trichantha var. ovata f. ovata . 

This form is characterized by its ovate leaves, densely villous 
underneath, and distinctly ferrugineous sepals. 

Known from Colombia and Venezuela. 

Specimens examined: 

COLOMBIA: Cundinamarca: Hillside east of Apulo, thickets 
along trail to Anapoima, alt. 460-600 m. , woody vine, r..P. Killip, 

A. Dugand and R. Jaramillo 38147 , May 4, 1944 Cl'S). Magdalena : 
Valle de Magdalena, R. Garsten (?) (W) ; Bond, Enredadera, Caliz 
Verde, Velloso, Anteras Verdes, dorsimelifijas , biloculares , Ovario 
bianco en la base y verde en el apice, Fruto globoso, verde, R. 
Romero-Castaneda 703 , February, 1948 (F) ; Santa Marta: alt. 500, 
Herbert H. Smith 876 , December, 1898-1899 (GH, K, US); Santa Marta, 
a twiner to 20 ft. , rare on border of forest below 1000 ft., fl. 
October-November, leaves subcordate at base, Smith 877 , November 
21, 1890-1901 (GH). Tolima: Honda, open rocky slope, alt. 300- 
400 m. , shrubby vine, corolla white, F.W. Pennell 3575 , January 
3-4, 1918 (GH, US). 

VENEZUELA: Lara: Entre Carora y Trentino, Jose Saer 724 , 
Enero, 1931 (F). Portuguesa: Vine on Calvario Hill, Guanare, 
H. Pittier 12046 , December 28, 1925 (NY, US). Trujillo: Loma 
de Moron near Valera, a vine, flower white, Pittier 10733 , Novem- 
ber 18, 1922 (GH, NY, US). Yaracuy: Iboa, 450 m. trailing on 
bushes, a woody vine, fl. white, Pittier 13074 , January 1, 1929 
(F, GH, US). 

B. trichantha var. ovata f. glabrata Myint & Ward, f. nov. 

Differt a forma typica var. ovatae foliis glabratis , sepala 
et pedicellis minus dense pubescentibus vel fere glabris , et 
pedicellis parum longioris. 

This form is characterized by its much varied, usually large, 
glabrate leaves, less densely pubescent or nearly glabrous sepals 
and pedicels, and slightly longer pedicels. 

Type: Colombia: Atlantico, entre Baranoa y Galapa, A. Dugand 
5643 , April 2, 1961 (US). 

This form is known from Panama, Colombia and eastern Venezuela, 
thus covering the whole range of the species. 

Specimens examined : 

COLOMBIA: Atlantico: Entre Baranoa y Galapa, 100 m. bos que 
marginal de un arroyo temporario, bejuco 10 m. long tallo lenoso 
delgad, 1 cm. diam. , A. Dugand 5643 , April 2, 1961 (US); entre 
Lena y Candelaria, alt. 30-50 m. , Dugand and R. Jaramillo 2789, 



1968 l^int & V/ard, Revision of Lonamia 221 

Enero 11, 1941 (US). Bolivar: Vicinity of Turbaco, Bro. Heriberto 
448 , November 1920 (US); north of Arjona, alt. 30-50 m. , thickets, 
vine, corolla white, E.P. Killip and A.C. Smith 14532 , November 15, 
1926 (GH, US). Cundinamarca: Bejuco trepador, frutos armarillos? 
Ferrocarril a Salgar, rio Guaduero, alt. 450 m. , H. Garcia-Barriga 
12296 , July 23, 1947 (US). Santander: Rio Surata valley near 
Ekjcaramanga , alt. 400-600 m. , woody vine, corolla white, thicket, 
Killip and Smith 16218 , December 28, 1926 (GH) ; river bank, upper 
Rio Lebrija valley, northwest of Bucaramanga, alt. 400-700 m. , 
vine somewhat woody, corolla white, Killip and Smith 16300 , Decem- 
ber 29, 1926 (GH). Doubtfxil specimen (no leaves nor flowers); 
Boyaca: Los Llanos, Rio Meta, Orocue, alt. 140 m. , Sabana, J. Cuat- 
recasas 4438 , November 3, 1938 (US). 

PANAMA: Toboquilla Island, vine, G.S. Miller, Jr. 2000 , March 
30, 1937 (US); Penonome and vicinity, 50-1000 ft. elevation, climb- 
ing over bushes, R.S. Williams 93 , February 23 - March 22, 1908 (NT). 

VENEZUELA: Delta Amacuro : Curiapo, alt. m. , enredadera, 
f lores blanca, Hermano Gines 4945 , December, 1952 (US). 

34. Bonamia balansae Hall, f. Bull. Herb. Boiss. 5:1002. 1897. 

Perennial, woody climbers. Stems twining, terete, 7 m. long, 
2-5 mm. thick, glabrous, minutely striated; older branches with 
whitish lenticels ; young branches sparsely punctate. Leaves 
petiolate, coriaceous or submembranous , glabrous, shining above, 
dull underneath; petioles, 5-14 mm. long, canaliculate above, glab- 
rous or sparsely pubescent when young; blades ovate-acuminate, 3-6 
cm. long, 2-3.5 cm. wide, rounded or subcordate at the base, sub- 
cuspidate-acuminate or acute mucronate at the apex; midrib impressed 
above, prominent underneath, with 5-7 pairs of lateral veins; finer 
veins distinct underneath. Flowers axillary, in few-flowered cymes 
or solitary, frequently in pseudo-racemes on short lateral branches; 
peduncles terete, rigid, 4-20 mm. long, glabrous or sparsely pubes- 
cent, pedicels 4-7 mm. long, turning to black when dry; bracts 
subulate, 1 mm. long, deciduous. Sepals coriaceous, equal or slightly 
unequal, orbicular, 5-6 mm. long, glabrous, turning to black when 
dry, subscarious along narrow margin, finely ciliate. Corolla 
yellow, campanulate-infrindibuliform, 2 cm. long, long-pilose on 
interplicae, glabrous on plicae, with entire or subentire limb. 
Stamens included; filaments puberulous or villous, adnate to corolla 
tube for 6 mm. ; anthers oval-oblong, 3 mm. long, sagittate at the 
base. Ovary with narrow annular disc, conical, 2 mm. long, glabrous; 
styles bifid for upper one-third, glabrous; stigmas orbicular- 
subbilobed. Capsules ovoid-quadrangular, 14-15 mm. long, 12-13 mm. 
in diameter, subacute or obtuse at the apex, glabrous, with ligneous 
wall of 0.5-1 mm. thick, 4-seeded, breaking into two valves (really 
4-valved) ; septum hard; seeds 5-6 mm. long, short-villous on ventral 
sides, densely fulvous-villous with white or yellowish white, long 
hairs on dorsal sides and along the edges. Cotyledons cordate- 
bilobed , emarginate at the apex; cotyledonary petioles fused. 

Type: Paraguay, San Salvador, dans les campos , Balansa 1078 , 
May 26, 1876 (GI). 



222 PHYTOLOGIA Vol. 17, no. 3 

Known only from a few collections made at a high elevation in 
Paraguay (Map 13). According to one collector, this species grows 
in forest on calcareous black soil. It has been collected in 
flower in January and February. 

Bo nam i a balansae is closely related to B. corumbaensis of 
southern Brazil, from which it differs by its longer stem, glab- 
rous leaves, longer petioles, acuminate or cuspidate leaf tip, 
ciliate sepals, and shorter stylar branches. 

Specimens examined: 

PARAGUAY: San Salvador, dans les campos , B. Balansa 1078 , 
May 26, 1876 (G) ; Zwischen Rio Apa und Rio Aquidaban, goldgelb, 7 m. 
hochsteigend , Waldparcelle, Kalkahaltiger, schwarzer Bod en, Hugel, 
K. Fiebrig 4531 , January 1908-1909 (BM, G, GH, L, W) ; San Salvador, 
F. Ro.las 3028 (1802), II, 1917 (GH). 

35. Bonamia corumbaensis Hoehne, Anex. Mem. Inst. Butantan 1 
(4):45, tab. 3. 1922. 

Perennial, suberect, shrubby plants, 30-60 cm. high; stems 
woody, suberect or scandent and slightly twining at the top; old 
branches rigid, with white lenticels , glabrous; young branches 
smooth, sparsely pubescent, becoming striated in age. Leaves short- 
ly petiolate, subcoriaceous or leathery and soft, sparsely pilose; 
becoming glabrous; petioles 3-10 mm. long; blades ovate-lanceolate 
or ovate-elliptic, 3-7 cm. long, 1.5-3 cm. broad, rounded or slight- 
ly cordate at the base, obtuse-mucronate or acute-mucronate at the 
apex; midrib impressed above, with 5-7 pairs of lateral veins. 
Inflorescences pedunculate, axillary cymes or racemes of two to 
few flowers, rarely solitary; peduncles short, 1-2 cm. long, sparsely 
pilose; pedicels 4-6 mm. long or sometimes longer; bracts small, 
scale-like, triangular, 1-1.5 mm. long, glabrous. Sepals coriaceous 
or subcoriaceous, slightly unequal glabrous; outer sepals fre- 
quently smaller, oblong or suborbicular-oblong, 5-7 mm. long, round- 
ed at the apex; inner sepals slightly larger, suborbicular, 6-8 mm. 
long, ciliate, rounded or emarginate at the apex. Corolla yellow 
or yellowish white, campanulate, 2 cm. long, 5-lobulate, densely 
ferrugineous-pilose on interplicae. Stamens inserted; filaments 
glabrous above, villous near the base; anthers oblong, 3.5-4.5 mm. 
long. Ovary ovoid-conical, glabrous; styles bifid to the middle 
or lower, filiform, glabrous; stigmas globose-capitate. Capsules 
conical-acuminate, glabrous, 4-seeded, slightly exceeding the 
length of sepals; seeds fulvous-villous (according to Hoehne). 
Cotyledons not known. 

Type: Brazil: Commissao Rondon: Corumba, Mato-Grosso, Campo 
seco, F.C. Hoehne 3042 , 2, 1911 (R-isotype) ; Corumba, Mato-Grosso, 
parto do paiol de polvora, F.C. Hoehne 3044 , 2, 1911 (R-paratype). 

Endemic to dry soil in southwestern Brazil near the borders of 
Bolivia and Paraguay (Map 13). It is represented by only two col- 
lections from the same location. The color of the corolla was not 



1968 



Myint & '.Yard, Revision of Bonamia 



223 




Map 13 



22li P H Y T L G I A Vol. 17, no. 3 

7-pcordpcl by tho collfctor and it is doubtful whother it is yellow 
or yollowish as it seems to be on the dry specimen. 

Although Hoehne stated that this species closely resembles 
B, burchp31i i , it seems to be more closely related to B. balansae 
of northern Paraguay, from which it differs in its erect habit, 
sparsely pubescent leaves, at least on the veins, obtuse or ob- 
tuse-mucronate leaves, and longer stylar branches (or shortly 
connate styles). 

36. Bonamia agrostopolis (Vp13.) Mall. f. Bot. Jahrb. 16:529. 1893. 
Convolvulus agrostopo3is Veil. Fl. Flum. 68, tab. 51. 1825. 
Browerin venulosa Meissn. in_ Martius , Fl. Bras. 7:326. 1869. 
Breweria agrostopolis (Veil.) Roberty, Candollea 14:30. 1952. 

Perennial, woody vines. Stems scandent or twining, terete, 
slender or 3-5 mm. thick, finely tomentose or puberulous while young, 
becoming glabrous in ago, longitudinally rugose, rugulose or ver- 
ruculose, rarely smooth; internodes variable in length, mostly 2-8 
cm. , occasionally longer. Loaves petiolate, herbaceous or sub- 
coriaceous , occasionally submembranous , thin; upper surface sparsely 
or rarely densely appressed-pilose while young, becoming glabrous 
in age; lower surface pilose or tomentose while young, becoming 
sparsely pilose in age; petioles 2.5-5 cm. long, finely tomentose 
or sericeous while young, becoming sparsely pilose or nearly glab- 
rous in age, distinctly canaliculate above; blades elliptic, elliptic- 
ovate or elliptic-acuminate, 10-16 cm. long, 5-9 cm. broad ('slightly 
smaller on the upper leaves), mostly entire, occasionally somewhat 
undulate or wavy at the margin, obtuse, acute or rounded at the base, 
acuminate-mucronate or acute-mucronate at the apex; veins distinc- 
tly impressed above, prominent below; lateral veins mostly 9-13 
pairs, with distinct intercostal veins, inflorescences axillary or 
terminal, long, multiflorus panicles, composed of numerous cymes; 
individual cymes 3- to 5-flowered , occasionally 1- to 2-flowered 
or rarely 7-flowered , shortly pedunculate ; peduncles variable in 
length, slender or stout; pedicels short, mostly 2-4 mm. long, tomen- 
tose; bracts small, scale-like, linear or linear-lanceolate, 2-3 
mm. long, deciduous; bracteoles similar to bracts, smaller. Sepals 
coriaceous, slightly unequal or subequal; outer two slightly short- 
er and narrower, ovate or ovate-orbicular, 5-6 mm. long, 3.5-4.5 
mm. broad, obtuse or rounded at the apex, tomentose and glabrescent; 
inner three larger, orbicular, orbicular-obovate, 5.5-7 mm. long, 
5.5-8 mm. broad, slightly emarginate or truncate at the apex, tomen- 
tose outside. Corolla purplish white (according to Velloso) or 
purple (according to Meissner) , funnelform or subcampanulate, mostly 
2 cm. long or slightly shorter, with entire limb, brown-sericeous 
or pilose on the interplicae. Stamens included; filaments filiform, 
short; anthers moderate in size, introrse, cordate at the base (ac- 
cording to Hallier). Ovary ellipsoid-conical or ovoid, glabrous, 
with obscure disc; styles bifid to the middle, filiform, unequal, 
glabrous; stigmas obscurely bilobed or globose. Capsules ovoid- 
conical or ellipsoid-conical, 1.5-2 cm. long, glabrous, brown or 
dark brown, hard-walled, 4-valvular, rarely 2-valvular, 4-seeded, 



1968 L^int & Ward, Revision of Donamia 225 

with thin or thick septum; seeds oval-oblong, or ellipsoid, 9-15 
mm. long, densely pilose with soft and bro\m hairs along the edge, 
brown-tomentose on the dorsal and ventral sides. Cotyledons ovate- 
cordate, multiplicate or corrugate and folded against radicle; 
cotyledonary petioles free for upper one-fourth. 

Type: Brazil: ''Habitat silvis arenosis maritimis ad Agrosto- 
polim;" type specimen not seen, presumably not extant. 

Known only from southeastern Brazil (Map 13). 

According to Meissner, this is the plant of highland regions. 
Flowering, according to Velloso, is in June and July. The only 
fruiting specimen examined in the present study was collected in 
late August. No flowering specimen was available for study. This 
was the first species of the genus described from South America, 
but under Convolvulus , and as such has been much confused with B. 
burchellii , to which it seems to be closely related. Large, glab- 
rate, elliptic or oblong-elliptic leaves with impressed veins are 
its most distinctive feature. 

Choisy (1845) included this species under his Breweria burchellii . 
Hallier realized its distinction from Bonamia burchellii , but its 
limits, as conceived by him, are somewhat doubtful because of his 
inclusion of Gaudichaud 567 as var. velutina, a specimen which should 
properly be assigned to Bj_ burchellii because of its pilose ovary 
and tomentose leaves. 

Specimen examined: 

BRAZIL: Minas Geraes : Dist. Ilheu, Fazenda da Tabunha, main 
road to northwest in cut-over woods, alt. 210 m. , woody vine climb- 
ing trees, green fruit, Ynes Mexia 4999 , August 24, 1930 (BM, F, G, 
GH, MO, NY, UC, US). 

37. Bonamia burchellii (Choisy) Hall. f. Bot. Jahrb. 16:529. 1893. 
Breweria burchellii Choisy, Mem. Soc. Phys. Hist. Nat. 

Geneve 6:4«?3. 1833. 
Ipomoea terminalis Choisy, Mem. Soc. Phys. Hist. Nat. Geneve 

8:54. 1838. 
Ipomoea lundii Choisy, Mem. Soc. Phys. Hist. Nat. Geneve 

8:56. 1838. 
Bonamia agrostopolis var. velutina Hall. f. Bull. Herb. 

Boiss. 5:1005, 1897. Type: Gaudichaud 567 (C). 
Convolvulus agrostopolis var. burchellii (Choisy) 0. Ktze. 

Rev. Gen. 3 (2):212. 1898. 

Perennial, woody climbing vines. Stems twining or scandent, 
terete, 1.5-1.9 m. long (according to Meissner, 1869), slender or 
as thick as 4-5 mm. , tomentose, villous or puberulous when young, 
glabrous or rarely puberulous in age, sparsely white-dotted, more 
densely so on older region; internodes mostly 4-7 (-10) cm. long, 
shorter on younger branches. Leaves petiolate, herbaceous, densely 
tomentose or vclutinous on both surfaces, dark green above, light 



226 P H Y T L G I A Vol. 17, no. 3 

groen below; potioles 8-25 mm. Jong, tomnntoso, canalicij]atf? abov; 
blades ovatn or ovate-acuminato, mostly 3-10 cm. long, 2.5-7 cm. 
broad, slightly shorter or narrower on tho uppr-r leaves, slightly 
undulate or subundulate at the margin, rounded, truncate or rarely 
subcordate at the base, acuminate, shortly acuminate-mucronate or 
acute-mucronate, rarely obtuse-mucronate at the apey ; veins obscure 
or rarely slightly impressed above, prominent below; lateral veins 
4-7 pairs; intercostal veins more distinct below, subparallel. In- 
florescences multiflorus, pedunculate panicles, axillary to the 
leaves or terminal on the lateral branch lets ; peduncles slender, 
variable in length, tomentose; pedicels 3-5 mm. long, slender, tomen- 
tose; bracts and bracteoles small, scale-like, linear or linear- 
lanceolate, 2-3 mm. long, deciduous. Sepals coriaceous, ovate, 
orbicular or ovate-orbicular, slightly unequal or subequal; outer 
two are smaller, mostly 3-4 mm. long, 2-3 mm. broad, sericeous or 
glabrate, obtuse, rounded or rarely broadly acute at the apex; inner 
sepals orbicular, 3.5-4.5 mm. long, densely sericeous outside at 
the center, glabrous at the margin, rounded or emarginate at the 
apex. Corolla white and vdth purplish or dark-colored eye or pur- 
plej infundibuliform, 2-2.5 cm. long, densely fulvous -sericeous 
fwhile young) and becoming fine-sericeous (in age) on interplicae. 
Stamens included; filaments filiform, shorter than styles, glabrous 
above, villous below; anthers oblong 2-3 mm. long, dorsifixed. 
Ovary ovoid-conical, pilose or sericeous at least near the apex; 
styles bifid to the middle or lov;er, filifonn, glabrous, rarely with 
soft, scattered hairs on the lower part, longer than filaments; 
stigmas reniform or subbilobed. Capsules ovate-conical, 1-1.5 cm. 
long, brown or dark-brovsm, with hard thick wall, opening into two 
or four valves, 4-seeded5 rarely 2-seeded; seeds ovoid or ellipsoid, 
5-7 mm. long, densely long-pilose along the edge, densely sericeous 
or tomentose on the dorsal and ventral sides. Cotyledons ovate- 
cordate, corrugate. 

Type: Brazil, Burchell 2778 (K-lectotype-not seen, GH-isotypeJ ) . 

From the specimens examined, this species seems to be localized 
in coastal regions of southeastern Brazil (Map 14). Its habitat 
is not knovm except for shallow sandbanks (restinga) recorded by 
one collector. It has been collected in flower from December to 
April; fruiting specimens bear no date of collection. 

This species is closely related to B. agrostopolis and 13. tonen- 
tosa , from both of which it is poorly defined. Future study may 
find these three merely as varieties of a single species. 

Specimens examined: 

BRAZIL: Rio de Janeiro, Burchell 2778 (F, GH, L, NT): Rio 
de Janeiro, Jacarepagua in fruticetis , P. Pus en 1985 , 22. 3. 1903 
(F, GH, US); Rio de Janeiro, M. Gaudichaud 567 , 1833 (G) ; Cosme 
Velho a' Laranjeiras (Rio de Jan.), cipo, flores rosadas , Glaziou 
4142 (BM, NY, R) ; Env. de Rio de Janeiro, Glaziou 13027 , 1882 (G); 
Flore des environs de Rio de Janeiro, Glaziou 13037 (G); Environs 
of Rio de Janeiro, Glaziou 14127 , 1882 (K) ; Morro da Babylonia 
(R. J.) cipo, fl. blancas, Glaziou 18381, March 29, 1891 (NY, R) ; 



1968 



Hyint fit Ward, Revision of Bonamia 



227 




Distribution of 
^ B. burchellii 
(•) B. tomentosa 
_ _. subsessilis 
^ B. mattogrossensis 



Map 14 



228 P H y T L G I A Vol. 17, no. 3 

Forest, Lincilpy , 1840 (K); Rcstinga da Ti.jura, Estado da Juanabara, 
Othon Machado , 21. 12. 1943 (RR); Restinga da Gavea , Lstado da 
Juanabara, Othon Machado (RB) ; Schling pflanzr- bfi Jacarepagua, 
E. Ule 4675 , May, ]898 (H BG ) . 

38. Bonamia tomentosa Hassler, Ropert. Sp. Nov. 9:148. 1911. 

Perennial, woody, high-climbing vines. Stems scandent or twin- 
ing, terete, 6-8 m. long (according to Hassler), slender or as 
thick as 4-6 mm. , densely fulvous-tomentose while young, sparsely 
puberulous or nearly glabrous in age; bark smooth or warty-plicate, 
with few or numerous lenticels ; internodes mostly 3-6 cm. long, 
occasionally shorter or reduced on the younger branches. Leaves 
petiolate, herbaceous, densely tomentose, fulvous-tomentose or 
velutinous on both surfaces, dark green and occasionally glabres- 
cent above, pale green below; petioles 10-30 (-40) mm. long, canal- 
iculate above, densely tomentose or rarely becoming sparsely tomen- 
tose; blades ovate, ovate-elliptic or occasionally suborbicular, 
or subcordate, mostly 5-11 cm. long, 4.5-10 cm. broad, entire or 
slightly undulate at the margin, subcordate or truncate at the 
base, obtuse-mucronate, emarginate, round ed-mucronate or rarely 
short-acuminate at the apex; veins very prominent below; lateral 
veins 6-8 pairs; intercostal veins obscure above, prominent below. 
Inflorescences axillary or terminal pseudopanicles or panicles, 
multiflorus ; peduncles short, variable in length, densely tomentose; 
pedicels short, 3-5 mm. long, tomentose, bracts linear or linear- 
lanceolate, 2-3.5 mm. long, deciduous; bracteoles similar to bracts, 
smaller, persistent or deciduous. Sepals coriaceous, ovate or 
ovate-orbicular, slightly unequal or subequal; outer two smaller 
4-6 mm. long, 3-4 mm. broad, tomentose or glabrescent near the 
margin, obtuse or rounded at the apex; inner three larger, sub- 
orbicular, 5-8 mm. long, densely sericeous or tomentose outside, 
emarginate at the apex, scarious at the margin. Corolla white, 
infundibuliform-campanulate, 2.3-2.8 cm. long, lobulate or sub- 
entire, soft-sericeous or short pilose on interplicae. Stamens 
included; filaments short, glabrous above, villous or puberulous 
below; anthers oblong, 2-3 mm. long, dorsifixed. Ovary conical 
or ellipsoid-conical, glabrous or sparsely pilose; styles bifid 
to the middle or lower, filiform, longer than filaments, glabrous; 
stigmas reniform or capitate-subbilobed, papillose-verrulose. Fruits 
not known. 

Type: Paraguay: In viciniis Caaguazu, E. Hassler 9038 , 1905 
(G-lectotype, BM, F, MO, NY, W-isotypes.' ) 

Known from southeast coastal region of Brazil to Paraguay 
(Map 14). 

According to Hassler, this is a plant of calcareous areas. It 
has been collected in flower in March but its fruiting period is 
not known. One is struck by the close relationship of this species 
to B. burchellii , and study of future collections may show it to 
be only a variety of the latter. From this species, it seems to 



1968 L^nt & Ward, Revision of Bonamia 229 

differ by its emarginate, obtxise-mucronate or rounded leaves (sub- 
cordate or truncate at the base), slightly larger sepals and dis- 
tinctly white corolla. Hassler, in describing B. tomentosa, noted 
a supposed close relationship to _B. agrostopolis . 

Specimens examined: 

BRAZIL: Tlha do Governador, Distrito Federal, G. Pabst (4.424 ) 
4908 , 30. 3. 1958 (F). 

PAMGUAY: Hernandarias , Sta. Teresa, Bertoni 4887 , 9. III. 
1950 (L, W) ; In viciniis Caaguazu, frutex scandens 6-8 m. , corolla 
alba, E. Hassler 9038 , 1905 (BM, F, G, MO, NY, W) ; Regio calcarea 
cursus superior is fluminis Apa, alt. 5-8 m. , petala blanca, Hassler 
11044 , 1912-13 (A, G, MO, NY, UC, US). 

39. Bonamia subsessilis Hassler, Repert. Sp. Nov. 9:149. 1911. 

Perennial, high climbing vines. Stems woody, twining, terete, 
2-4 mm. thick or slightly thicker, about 4 m. long, densely brown 
villous or tomentose when young, becoming glabrous or minutely 
puberulous in age; old branches vs^ith purplish bark, minutely ver- 
ruculose-punctate, longitudinally plicate-rugose; internodes 4-10 
cm. or slightly longer. Leaves petiolate, soft, subcoriaceous or 
submembranous , dark green and softly tomentose above, grey-green 
and densely tomentose on the lower surface; petioles 5-20 mm. long, 
canaliculate above, fulvous -tomentose; blades ovate or ovate-orbi- 
cular, 7-13 cm. long and 6-10 cm. vv'ide (upper leaves subtending 
individual cymes much smaller) , subcuneate or obtuse-acute at the 
base, obtuse-mucronate, round ed-mucronate, acute-mucronate or emar- 
ginate-mucronate at the apex; lateral veins 6-9 pairs; intercostal 
veins conspicuous, subparallel. Inflorescences axillary, racemes 
or panicles, composed of sessile or subsessile cymes of 1-5 flowers; 
raceme rachis 10-60 mm. or longer, tomentose with brown hairs; in- 
dividual cymes sessile or subsessile; pedicels and peduncles very 
short or absent; bracts and bracteroles small, lanceolate, 2-3 mm. 
long-tomentose. Sepals coriaceous or subcoriaceous, equal or slight- 
ly unequal, orbicular, ovate or ovate-orbicular; outer sepals 4-5 
mm. long, 3-4 mm. broad, densely tomentose or ferrugineous outside, 
glabrous inside, obtuse or rounded at the apex; inner sepals 5-6 mm. 
long, 5 mm. broad, sparsely tomentose or nearly glabrous outside, 
glabrous inside, rounded or emarginate at the apex. Corolla white, 
campanulate-infundibuliform, about 2 cm. long, entire or subentire 
at the margin, fulvous-pilose on interplicae. Stamens included; 
filaments villous; anthers oblong, 2-3 mm. long, dorsifixed. Ovary 
ovoid-conical, glabrous; styles bifid to the middle or lower, glab- 
rous; stigmas reniform or subbilobed. Capsules not known. 

Type: Paraguay: Caballero-cue (Zwischer Rio Apa und Rio Aqui- 
daban) , Trochnen Camp, mit Bursch bewaldete Anhohe, bis 4 m. hoch, 
kletternd, Weiss, K. Fiebrig 4764 , February, 1908-1909 (G-lectotype, 
BM, GH, L-isotypesJ). 

Dry highland of northern Paraguay near the Brazilian border 
(Map 14). It is known only from the type collection, which was 
collected in flower. 



230 P H Y T L G I A Vol. 17, no. 3 

Although the author of this species stated that it is related 
to j3. tomentosa , it seems to be more closely related to the IJra- 
zilian species, M. mattogrossensis , from which it differs only 
by a white corolla and sepals glabrous inside. Future study might 
prove n. subs ess ill s and j3. mattogrossensis to be cont^peci f ic. 

40. Bonamia mattogrossonsis Hoehne, An. Mem. Inst. Butantan 1 
Tfasc. 4): 45, tab. 4. 1922. 

Perennial, high climbing vines. Stems woody, terete, twining 
or scandent, 2-4 mm. thick or thicker, densely brown-tomentose or 
ferrugineous when young, becoming glabrous in age; old branches 
sparsely punctate or with scattered, white lonticels. T^eaves petio- 
late, soft, thick, subcoriaceous , densely soft-velutinous and dark 
green above, densely brown-tomentose or ferrugineous and light green 
underneath; petioles 1-2 cm. long, mostly 1.5-2 mm. thick, canali- 
culate above, densely tomentose or becoming sparsely puberulous in 
age; blades ovate, ovate-elliptic or ovate-acuminate, 5-12 cm. 
long, 3.5-8 cm. broad (slightly smaller on upper leaves), attenuate, 
subattenuate, obtuse or rarely subcordate at the base; acuminate, 
obtuse-mucronate or round ed-mucronate at the apex; lateral veins 
6-10 pairs, with distinct intercostal veins. Inflorescences axil- 
lary, sessile cymes of 3-7 flowers or on axillary short branches 
forming panicles composed of sessile cymes or axillary racemes; 
individual flowers sessile or subsessile; bracts and bracteoles 
small, 1-3 mm., lanceolate-linear, deciduous. Sepals coriaceous, 
equal or slightly unequal; outer two sepals ovate, 4-5 mm. long, 
3.5-4.5 mm. wide, densely tomentose or ferrugineous outside, dense- 
ly sericeous inside except glabrous center, obtuse-acute or broadly 
acute at the apex; in-out sepal (third sepal) oblique or orbicular- 
oblique, densely tomentose outside except glabrous inner margin, 
densely soft-sericeous inside on the outer margin, glabrous at the 
center and on the inner margin; inner t\-^o sepals orbicular or 
obovate-orbicular, sparsely or densely tomentose outside at the 
center, nearly glabrous at the margin, glabrous inside, rounded, 
truncate or slightly emarginate at the apex. Corolla purple, 
pale purple or violet, campanulate-infundibuliform, mostly 2 cm. 
long, subentire, entire or slightly lobulate at the margin, brown- 
pilose or villous with brownish long hairs on interplicae. Stamens 
included; filaments villous, at least lower parts; anthers oblong 
or narrowly elliptic-oblong, 2-3 mm. long, dorsifixed. Ovary oblong- 
conical or ovoid-conical, glabrous; styles bifid to the middle or 
lower, with distinct stylopodia, glabrous or with scattered hairs; 
stigmas reniforra or subbilobed. Capsules not known. 

Type: Brazil: Mato-Grosso: Commissao Rondon, Coxipo da Ponte, 
Cuiaba. flor. alvo-arroxeada, F.C. Hoehne 4655 , em Marco (1911) 
(R-lectotypel). 

Known only from the type location in the northern part of Mato- 
Grosso in western Brazil (Map 14). According to the author of the 
species, the plant gro\>?s in dry regions. Further collections are 
desired, since fruit, seeds and definite flowering and fruiting 



1968 Lijrint & '«Yard, Revision of Bonamia 231 

periods are not yet known. The only material available for the 
present study was collected in flower. 

The outstanding features of this species are (1) sessile 
flowers, (2) large leaves, slightly cuneate at the base, (3) outer 
sepals sericeous inside and (4) corollas purple. It is closely 
related to B. subsessilis or Paraguay, from which it differs only 
by its outer sepals being sericeous inside and a purple corolla. 
This species is separable, although not clearly, into two varieties. 

40a. Bonamia mattogrossensis Hoehne var. mattogrossensis . 

This variety is characterized by ovate-acuminate or ovate- 
elliptic leaves, acute, acuminate or rarely obtuse-acuminate and 
mucronate at the apex, cuneate or subcuneate at the base, and long 
raceme-rachis . 

Known only from the type collection. 

40b. Bonamia mattogrossensis Hoehne var. obtusifolia Hoehne, An. 
Mem. Inst. Butantan 1 (fasc. 4) :46, tab. 5. 1922. 

This variety differs from the typical variety by its smaller, 
ovate, obovate or broadly elliptic leaves, rounded, truncate or 
obtuse and mucronate at the apex, and short raceme-rachis or inflor- 
escence axillary and sessile. 

Type: Brazil: Mato-Grosso: Commissao Rondon, Coxipo da Ponte, 
Cuiaba, flor roxa, F.C. Hoehne 3039 , 3. 1911 (R-lectotype.'). 

Known only from the type collection from western Brazil. 



Little-Known Species 

The following species are poorly known and no specimens were 
available for study. The descriptions given here are based on 
the original descriptions. If material becomes available for com- 
parison, they may prove to be merely abnormal forms or local var- 
iants of the other species covered previously. They are included 
here to make this study as complete as possible. 

41. Bonamia abscissa (Choisy) Hall. f. Bull. Herb. Boiss. 5:812. 
1897. 
Breweria abscissa Choisy, in DC. Prodr. 9:438. 1845. 

Stems elongate, ferrugineous. Leaves petiolato, slightly ferru- 
gineous or glabrate; petioles 2.5 cm. long, ferrugineous; blades 
cordate-ovate, 5-7.5 cm. long, entire at the margin, slightly acu- 
minate at the apex. Flowers axillary, mostly solitary; peduncles 
not equalling petioles; pedicels ferrugineous. Sepals ovate-orbi- 
cular, 6-8 mm. long, subequal, ferrugineous outside, acutish at 
the apex. Corolla red, campanulate, 2.5-3.2 cm. long, truncate or 



232 PHYTOLOGIA Vol. 17, no. 3 

entire at the margin. Ovary villous; styles bifid almost to the 
base. Capsules glabrous. 

Typo: Madagascar, Bojnr . 

Reported from woods at Mooza in eastern Madagascar. 

In many characteristics this species resembles B. semidigyna , 
to which it must bo closely related. Ilallier treated B. abscissa 
under B. semidigyna in his earlier paper (1893) , Imt later ''IS??) 
he treated the two as distinct species. According to Hallier, 
this species differs from B. semidigyna by its red corolla with 
truncate or entire limb and uniflorous inflorescence. J 

42. Bonamia boivinii Hall, f. Bot. Jahrb. 18:91, 1894. 

Stems woody, elongate, twining, terete, glabrous; lower inter- 
nodes 10 cm. long. Leaves shortly petiolate, glabrous; blades 
ovate, 4 cm. long, 15 mm. broad, gradually smaller toward the apex 
of stem, falcate-recurved and folded, subacute at the base, acute 
and mucronate at the apex. Inflorescences dense, multiflorous, 
terminal, composed of dichasial cymes or subumbellate, shortly 
pedunculate; flowers small; peduncles short, 2 cm. long, longer J 
than petioles, finely subsericeous ; bracteoles small, aggregate, ' 
scale-like. Sepals coriaceous, orbicular, equal, glabrous, black, 
ciliate at the margin. Corolla (not yet unfolded) sericeous outside. 

Type: Northwest Madagascar: Ins. Nossi-be, Boivin , 1853. 

This species was offered by Hallier as new with an accompan- 1 
ying description far too brief for satisfactory comparison v;ith 
other species. From the description, it seems to be similar to 
B. d ens i flora , except in leaves. Hallier designated the type 
specimen as deposited at the herbarium of Boissier; this specimen 
cannot be located. 

43. Bonamia langsdorffii (Meissn.) Hall. f. Bull. Herb. Boiss. 

5:814. 1897. 

Breweria Langsdorffii Meissn. in Martius , Fl. Bras. 7:325. 
1869. 

Stems slender, perhaps twining, adpressed-pilose or glabrate. 
Leaves petiolate, subcoriaceous ; petioles 2-6 mm. long, slender, 
canaliculate; blades ovate or oblong-elliptic, 2.5-4.2 cm. long, 
1.7-2.5 cm. broad, entire or slightly wavy at the margin, slightly 
cordate at the base, obtuse-mucronate at the apex. Flowers solitary, 
axillary, pedunculate; peduncles unequal with the leaves, pubescent; 
pedicels as long as peduncles, the two together 1.8-2 cm. long; bracts 
two, opposite, minute, about 2 mm. long, acute. Sepals ovate, 8 mm, 
long, 4-5 mm, broad, coriaceous-herbaceous, equal, obtuse, glabrous. 
Corolla white (?), broadly infundibuliform, 2-2.5 cm. long, slightly 
less than 2.5 cm. in diameter at the limb, pilose outside on inter- 
plicae. Styles shorter than corolla, filiform, bifid, connate for 
lower 4 mm. 



1968 l^yint &. '.Yard, Revision of Bonamia 233 

Type: Brazil: Rio de Janeiro, Langs dorff . 

This species is very closely allied to, and perhaps conspecific 
with, B. burchellii , B. agrostopolis or _B. tomentosa , from which 
it differs by its solitary flowers. 

44. Bonamia capitata (Dammer) v. Ooststroom, Rec. Trav. Bot. Neerl. 

33:212. 1936. 

Prevostea capitata Dammer, Bot. Jahrb. 23 (Beibl. 57):36. 
1897. 

Shrubby plants; branches tomentose. Leaves closely spaced, 
sessile, coriaceous, pubescent above, grey-tomentose below, the 
margin revolute; blades lanceolate, rounded at the base, mucronate 
at the apex; nerves scarcely prominent below. Inflorescences ter- 
minal, of densely compacted, subglobose cjmies. Sepals lanceolate, 
setose-acuminate at the apex, the outer somewhat larger, densely 
pilose, ciliate on margins. Corolla blue, densely pilose on upper 
part (interplicae?). Styles bifid to the middle, pilose at the 
base; stigmas renifonn, capitate. Fruits not known. 

Type: Brazil: "Civitate Goyaz ad Fazenda da Boa Vista in 
Campo," Glaziou 21799 , Jan. 14, 1895; not seen. 

Known only from south-central Brazil. 

As noted by Dammer, this species appears well marked by its 
distinctive inflorescences. It would seem allied to B. tomentosa 
and B, subs ess ilis . 

45. Bonamia sedderoides Rendle, Jour. Bot. 46:178. 1908. 

Spreading undershrub. Stem 4-6 dm. long, 2 mm. thick, slender, 
covered with silky, whitish hairs. Leaves 1.3 cm. long, 3 mm. wide. 
Bracteoles 7-8 mm. long. Sepals 1.1-1.2 cm. long, 5 mm. broad. 
Corolla probabl}' 2.5 cm. long. Stamens 8 mm. long; anthers linear- 
oblong, 3 mm. long. Styles free nearly to the base. 

Type: Southeast Angola, in shrub-gro\>m pasturage on sandy 
alluvial soil at the foot of the Serra Ferreire de Amiral, western 
side, Gossweiler 2888 , February 9 (BM-holotype, K-isotype) ; not 
available. 



Doubtful and Excluded Species 

Doubtful Species 

Bonamia vignei Hoyle, Kew Bull. 1934:138. 1934. 

Although the author of this species stated that it is related 
to _B. cymosa (=_B. thunbergiana of the present treatment), it appears 
to be quite different from the latter in several features, especially 



231 P H T T L G I A Vol. 17, no. 3 

Ny its £iccrosccnt" sepals, which are not charactfristic of thr- g<'nus 
r^onamia . Since no specimen was availal)lG for the present study, 
its transfer to the genus Calycobolus is not attempted here. 

excluded Species 

Bonamia aJthoffiana Dammor, Pflnnz. Ostafr. C:329. 1895. 

=Convolviilus kJlimandschari Cngler, Ilochgeb. Trop. Afr. 348. 
1392. 
Bonamia angusti Folia (N'ash) Wilson, Jour. Arnold Arb. 41:306. ]960. 

=Styli5ma patens (Desr.) Myint, ssp. angusti folia (Nash) Myint, 
Drittnnia 18:112. 1966. 
Bonamia aguatica '''■/alt.) Gray, Man. ed. 5. 376. 1867. 

^Stylisma aguatica (Walt.) Raf. Fl. Tell. 4:83. 1838. 
Bonamia capons is (Baker) Burtt Darvy, Ann. Trans. Mus. 3:121. 1912. 

=Seddera capensis (Meyer) Mall. f. Rot. Jahrb. 18:86. 1893. 
Bonamia glonerata (Balf. f . ) Hall. f. Bot. Jahrb. 18:90. 1893. 

=S odd era glomerata (Balf. f . ) 0. Schwartz, Mitt. Inst. Allgemeine 
Bot. Hamburg 10:1971. 1939. 
Bonamia humiptrata (Walt.) Gray, Proc. Am. Acad. 5:337. 1862. 

=Stylisma humi strata (Walt.) Chapm. Fl. S. U. S. ed. 1, 346. 1860. 
Bonamia michauxii (FernT and Schub. ) Wilson, Jour. Arnold Arb. 41: 
306. 1960. 

=Stylisma aouatica (Walt.) Raf. Fl. Tell. 4:83. 1838. 
Bonamia patens (Desr.) Shinners, Castanea 27:75. 1962. 

=Stylisma patens (Desr.) Myint, Brittonia 18:110. 1966. 
Bonami a pickeringii (Torr. ox Curtis) Gray, Man. ed. 5. 376. 1867. 

=Stylisma pickeringii (Torr. ex Curtis) Gray, Man. ed. 2. 335. 
1856. 
Bonamia poranoides Hall, f. Bull. Herb. Boiss. 5:1007. 1897. 

=Metaporana densiflora (Hall, f . ) N.E. Brown, Kew Bull. 1914: 
169. 1914. 
Bonamia schizantha (Hall, f . ) Meeuse, Bothalia 6:665. 1957. 

=Seddera schizantha Hall. f. Bull. Herb. Boiss. 6:532. 1898. 
Bonamia spinosa Vierhapper, Oesterr. Bot. Zeitschr. 287. 1904. 

= ?Seddera sp. or ? Convolvulus socotranus Verdcourt, Kew Bull. 
1957:344. 
Bonamia suffruticoa (Schinz) Burtt-Davy, Ann. Transvaal Mus. 3:121. 
1912. 

=Seddera suffruticosa (Schinz) Hall. f. Bot. Jahrb. 18:88. 1893. 
Bonamia villosa (Nash) Wilson, Jour. Arnold Arb. 41:306. 1960. 

^Stylisma villosa (Nash) House, Bull. Torr. Bot. Club 34:149. 
1907. 
Bonamia volkensii Dammer, Pflanz. Ostafr. C:329. 1895. 

=Hewittia sublobata (L.f.) 0. Ktze. Rev. Gen. PI. 2:441. 1891. 
Br ewer i a africana (G. Don) Benth. and Hook. f. Gen. PI. 2:877. 1876. 

=Calycobolus africanus (G. Don) Myint, comb. nov. 
Breweria alsinoides Merrill, Interpr. Rumph. Herb. Amboin. 46. 1917. 

=Evolvulus alsinoides (L.) L. Sp. PI. ed. 2. 392. 1762. 
Breweria alternifolia Radlk. Abhandl. Nat. Ver. Bremen 8:413. 1884. 

=Calycobolus africanus (G. Don) Myint, supra. 
Breweria angustifolia Nash, Bull. Torr. Bot. Club 22:155. 1895. 



1968 Ifyint & Ward, Revision of Bonamj a 235 

=Stylisma patens (Desr.) Myint, var. angustifolia (Nash) Mvint, 

Brittonia 18:112. 1966. 

Rreweria aquatica (Walt.) Gray, Syn. Fl. N. Am. 2 (1) :217. 1878. 

=Stylisma aquatica (Walt.) Raf. Fl. Tell. 4:83. 1838. 
Breweria argentea Terrace, Ann. Inst. Bot. Roma 5:104. 1893. 

=Seddera latifolia Hochst. and Steud. Flora 27, Beil. 8, t. 5 
1844. 
Breweria baccharoides Baker, Kew Bull. 1894:68. 1894. 

=Seddera suffruticosa (Schinz) Hall. f. Bot. Jahrb. 18:88. 1893. 
Breweria campanulata Baker, Kew Bull. 1894:68. 1894. 

=Calycobolus camganulatus (Baker) Myint, comb. nov. 
Breweria capensis (Meyc^TBaker, in Dyer, Fl. Cap. 4 (2):80. 1904, 

-Seddera capensis (Meyer) Hall. f. Bot. Jahrb. 18:86. 1893. 
Breweria choisyana Steud. Nomencl. ed. 2, 1:224. 1840. 

=Seddera evolvuloides (Choisy) Wight, Icon. 4 (2):13, t. 1369. 
1848. 
Breweria Codonanthus Baker ex Oliver, Hook. f. Icon. pi. 23. 

=Calycobolus aft;icanus (G. Don) Myint, supra. 
Breweria conglomerata Baker, Kew Bull. 1894:68. 1894. 

=Seddera conglomerata (Baker) Hall. f. Bull. Herb. Boiss. 5-1008. 
1897. 
Breweria evolvuloides R. Br. Salt. Abyss. App. 65. 1814. 

=Seddera arabica (Forsk. ) Choisy, in DC, Prodr. 9:441. 1845. 
Breweria evolvuloides Choisy, Mem. Soc. Phys. Genev. 6:494. 1833. 
=Seddera evolvuloides (Choisy) Wight, Icon. 4 1^2) :13, t. 1369. 
1848. 
Breweria evolvuloides Vatke, Linnaea 43:523. 1882. 

=Seddera latifolia Hochst. et Steud. Flora, Beil. 8, t. 5. 1844. 
Breweria fastigiata Balf. f. Proc. Roy. Soc. Edin. 12:83. 1883. 

^Convolvulus socotranus Verdcourt, Kew Bull. 1957:344. 1957. 
Rreweria glaucata Peter, in Engler and Prantl. , Naturl. Pflanzenfam. 
4(3a):17. 18 9 7 . 
=Seddera glomerata (Balf. f . ) 0. Schwartz, Mitt. Inst. Allge- 
meine Bot. Hamburg 10:1971. 1939. 
Breweria glomerata Balf. f. Proc. Roy. Soc. Edin. 12:83. 1883. 

-Seddera glomerata (Balf. f . ) 0. Schwartz, 1. c. 1939. 
Breweria Hassleriana Chod. Bull. Herb. Boiss. Ser. II. 5:683. 
=Convolvulus hasslerianus (Chod.) O'Donell, Lilloa 23:430. 
1950. 
BrevN?eria heudelotii Baker, Kew Bull. 1894:68. 1894. 

=Calycobo3us heudelotii (Baker) Myint, comb. nov. 
Breweria hispida Franchet, Sert. Somal. p. 43. 1882. 

=Seddera somalensis (Vatke) Hall. f. Bot. Jahi-b. 18:90. 1893. 
Breweria humistrata (Walt.) Gray, Syn. Fl. N. Am. 2 (1):217. 1878. 

=St\0.isma humistrata (Walt.) Chapm. Fl. S. U. S. ed. 1, 346. 1860. 
Breweria intermedia Hochst. Flora 27, Beil. 8. 1844. 

=Seddera intermedia ![ochst. et Steud. Flora 27, Beil. 8. 1844. 
Breweria latifolia Hochst. Flora 27, Beil. 8. 1844. 

=Seddera latifolia Hochst, and Steud., Flora 27, Beil, 8, t, 5, 
1844. 
Breweria linifolia Spreng. Syst. 1:614. 1825. 

=Wahlenbcrgia linarioides (Lam.) A. DC. in DC. Prodr. 7r2) :440. 
1839. — 



236 PHYTOLOGIA Vol. 17, no. 3 

Hrev.'cria malvacea Klotzsch, in Pctors , Rcise Mossarab. 'tot. 245. t. 

37. 1861. 
=Astripomoca malvacea (Klotzsch) Moeuse, Bothalia 6:710. 1957. 
Brewpria mpxicana Ik-nsl. Biol. Central Am. Bot. 2:400. 1882. 

=Calycobolus velutinus (Mart, and Gal.) Houso, Bull. Torr. Bot. 
Club 34:14. 1907. 
Broworia michauxii Fern, and Schub. Rhodora 51:37. 1949. 

= Stylisma aqiiatica (Walt.) Raf. Fl. Toll. 4:83. 1838. 
Br p wen" a microcpphala Baker, Kew Bull. 1894:68. 1894. 

=Spddera welwitschii Hall. f. Bot. Jahrb. 18:88. 1893. 
Br ewer 1 a minima Gray, Proc. Am. Acad. 17:228. 1881-82. 

=Convolvulus simulans L.M. Perry, Rhodora 33:76. 1931. 
Breweria mirabilis Baker ex Oliver, liooker. f. Icon. PI. 23, t. 
2276. 1894. 
=Calycobolus £ani£ariulatus (Baker) Myint, supra. 
Breweria montevidensis Peter7 in Engler et Prantl. , Naturl. Pflan- 
zenfam. 4 (Abt. 3a) :16. 1897. 
=Convolvulus ottonis Meissn. inMartius, Fl. Bras. 7:311. 
1869. 
Breweria oxycarpa A. Rich. Tent. Fl. Abyss. 2:76. 1851. 

=Seddera arabica (Forsk.) Choisy in DC. Prodr. 9:441. 1845. 
Breweria parvi flora Arn. ex Steud. Nonencl. ed. 2, 1:224. 1840. 
=Seddera evolvuloides (Choisy) Wight, Icon. 4 (2) :13, t. 1369. 
1848. 
Breweria patens (Desr.) Fernald , Rhodora 42:298. 1940. 

Stylisma patens (Desr.) Myint, Brittonia 18:110. 1966. 
Breweria pedunculata Balf. f. Proc. Roy. Soc. Edinb. 7:83. 1884. 
=Seddera pedunculata (Balf. f . ) Hall. f. Bull. Herb. Boiss. 5: 
1010. 1897. 
Breweria pickeringii (Torr. ex Curtis) Gray, Syn. Fl. N. Am. 2 (1): 
217. 1878. 
=Stylisma pickeringii (Torr. ex Curtis) Gray, Man. ed. 2, 335. 
1856. 
Breweria rotundifolia Watson, Proc. Am. Acad. 23:281. 1888. 

=Evolvulus rotund if olius (Watson) Hall. f. Bot. Jahrb. 16:530. 
1893. 
Breweria scoparia Lindl. Fl. Med. 400. 1838. 

=Convolvulus scoparius L. f. Suppl. 135. 1781. 
Breweria sessiliflora Baker, Kew Bull. 1894:68. 1894. 

=Seddera suffruticosa (Schinz) Hall. f. Bot. Jahrb. 18:88. 1893. 
Breweria somalensis Vatke, Linnaea 43:523. 1882. 

=Seddera arabica (Forsk.) Choisy in DC. Prodr. 9:441. 1845. 
Breweria suffruticosa Schinz in Verb . Bot. Ver. Brand. 30:275. 1888. 
=Seddera suffruticosa (Schinz) Hall. f. Bot. Jahrb. 18:88. 1893. 
Breweria tenella (Desr.) Peter, in Engler et Prantl, Naturl. Pflan- 
zenfam. 4 (3a) :16. 1897. 
=Stylisma humlstrata (Walt.) Chapm. Fl. S. U. S. ed. 1, 346. 
1860. 
Breweria tiliaefolia Baker, Jour. Linn. Soc. Bot. 22:508. 1887. 
= Rapona madagascariensis Baill. Hist, des PI. 10:328. 1888. 
Breweria trichosanthes (Michx. ) Small, Fl. S. E. U. S. 595. 1903. 

=Stylisma patens YPesr. ) Myint, Brittonia 18:120. 1966. 
Breweria valerianoides Villar, Nov. App. 143. 1880. 



I 



1968 l^int & Ward, Revision of Bonania 237 

=Jacquemontia paniculata (Burm. f . ) Hall. f. Bot. Jahrb. 16: 
541. 1893. 

Breweria villosa Nash, Bull. Torr. Bot. Club 22:154. 1895. 

=Stylisma villosa (Nash) House, Bull. Torr. Bot. Club 34:149. 
1907. 
Breweria virgata Vatke, Linnaea 43:523. 1882. 

=Seddera virgata Hochst. et Steud. Flora, Beil. 8, t. 5. 1844. 



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1897. Bausteine zu einer Monographie der Convolvulaceen. 

V. Ubersicht uber die Gattung Bonamia , Bull. Herb. Boiss. 

5:804-820; 996-1003. 
Hochstetter, C.F. 1844. Seddera , Flora 27, Bes. Beil. 7, t. 5. 
Hooker, J.D. 1844. Trichantha, Icon. PI. tt. 666 and 667. 



238 P H Y T L G I A Vol. 17, no. 3 

House, II. D. 1907. Studips in thp North Amorican Convolvulaceaf III. 

Calycoboliis , Ik^namia , and Stylisma , Bull. Torr. B^^t. Club 34: 

143-149. 
Humboldt, A. v., fionpland , A., and Kunth , C.S. 1818. Dufourpa, 

3:88. In A. v. Humboldt, A. Itonpland and C.S. Kunth, Nova 

Genera ot Spccins Plantarum , P. School ^etc), I'aris. 
Hutchinson, .J. and Dalziel, J.M. 1931. Convolvulaccae, 2:208-219. 

In J. Hutchinson and J.M. Dalziel, riora of West Tropical 

Africa , The Crown Agents, IjOndon. 
.Jaumo Saint-Hilaire, J.H. 1805. Bonamia , 2:349. In J.H. Jaume 

Saint-Hilaire, Exposition dcs Fami He Naturellcs , Strasbourg 

and Co. , Paris. 
Karsten, H. and Triana, .1. 1856. Trichantha , Linnaea 28:437. 
Lanjouw, J. and Staflou, F.A. 1964. Index Herbariorum , 5th ed. pt 

1. Regnum Vegetabile vol. 31. Utrecht. 
Macbride, J.F. 1959. Flora of Peru, Publ. Field Mus. Bot. 13 C51) : 

455-536. 
Meeuse, A. D. J. 1957. The South African Convolvulaceae, Bothalia 

6:641-792. 
Meissner, C.F. 1869. Prevostea and Breweria , 7:323-327. In C.F.P. 

v. Martius , Flora Brasiliensis , Munchen. 
Myint, T. 1966. Revision of the genus Stylisma , Brittonia 18: 97-117. 
1968. Australasian species of Bonamia . Burma Jour. Life 

Science 1: 28-35. 
Nees von Esenbeck, C.G. and Martius, C.F.P. v. 1823. Dethardingia , 

Nov. Act. Nat. Cur. 11:80. 
O'Donell, C.A. 1959. Convolvulaceas argentinas , Lilloa 29:87-348. 
Ooststroom, S.J. v. 1932. Convolvulaceae, 4 (1):66-102. In A. 

Pulle, Flora Suriname , J.H. de Bussy, Ltd., Amsterdam. 
1954. Convolvulaceae, 4:388-512. In C.G. G.J. v. Steenis, 

Flora Males iana Ser. I, P. Noordhoff, Ltd. 
Peter, A. 1897. Convolvulaceae, 4 (3):l-40; 375-377. In A. Engler 

and K Prantl, Die Naturlichen Pflanzenfamilien , Leipzig. 
Poiret, J.L.M. 1810. Bonamia , 1:677. In J.B. Lamarck, Encyclopedic 

Method i que , botanique, Paris. 
Rafinesque, C.S. 1818. "A sketch of the Botany of South Carolina 

and Georgia" by Stephen Elliott, Am. Monthl. Mag. Grit. Rev. 

3:96-101. 
Roberty, G. 1952. Genera Convolvulacearum, Candollea 14:11-60. 
Roemer, J.J. and Schultes , J. A. 1819. Calycobolus , 5:4. In J.J. 

Roemer and J. A. Schultes, Systema Vegetabilium , Stuttgardiae, 

Gotta. 
Shinners, L.H. 1962. Synopsis of United States Bonamia including 

Breweria and Stylisma (Convolvulaceae), Castanea 27:65-77. 
Small, J.K. 1933. Manual of the Southeastern Flora . University 

of North Carolina Press, Chapel Hill. 1554 p. 
Sprengel, K. 1828. Re inward tia , 1:527. In K. Sprengel, Systena 

Vegetabilium , Goettingae. 
Verdcourt, B. 1957. Notes on East African Convolvulaceae (Part I), 

Kew Bull. 1957:334-347. 
1963. Convolvulaceae, p. 1-161. In C.E. Hubbard and E. 

Milne-Redhead, Flora of Tropical East Africa . The Crown 

Agents, London. 



1968 Myint & Ward, Revision of Bonamia 239 

Wilson, K.A. 1960. The Genera of Convolvulaceae in the SoutheastHrn 
United States, Jour. Arnold Arb. 41:298-317. 



Appendix 

New Names and Combinations 

Nev; species : 

B. brevipedicellata Myint & Ward 

Now varieties : 

_B. media var. emarginata Myint & Ward 
B. menziesii var. rockii Myint & Ward 

New forms : 

B. trichantha var. ovata forma glabrata Myint & Ward 

New combinations : 

B. elliptica (Smith & Schubert) Myint & Ward 

B. semidigyna var. semidigyna f. ambigua (Hall, f . ) 

Mv'int 0/ Ward 
_B. sulphur ea (Brandg.) Myint S- Ward 

New combinations in Calycobolus : 

£. africanus (G. Don) Myint 
C_. campanulatus (Baker) Myint 

C. heudelotii (Baker) Myint 



ADDITIONAL NOTES ON THE GEIRB VITEX . X 
Harold N, L'oldenke 



VITEX Tourn. 

Additional bibliography: J. F, Gmel. in L., Syst. Nat., ed. 13, 
pr. 1, 2: 890 (1789) and pr. 2, 2: 890. 1796; Steud., Nom. Bot. 
Phan., ed. 1, 228 & 888. 1821; Guinet <i Sauvage, Trav. Inst. Sci- 
ent. Ch^rif., ser. g6n., 2: 121. 195U; J. Bush-Brown, Shrubs b. 
Trees Home Landsc. 161, 19$, & 197. 1963; Martlnez-Crovetto, Bon- 
plandia 1: 177 & 198. 1963; E. Lawrence, South. Gard., ed. 2, 
139, 219, & 261. 1967; Doolittle k Tiedebohl, Southwest. Gard., 
ed. 2, 90, 170, & 171. 1967; Vyas, Joum. Bombay !,'at. Hist. Soc . 
61;: 219. 1967; Anon., Biol. Abstr. U9: 390 (I968) and U9 (8): S. 
185. 1968; Moldenke, Phytologia 16: 1;87~502, 507, 509, ^ 512 
(1968) and 17: 8—56 !k llli— 120. I968; L'oldenke, R4sun^ Suppl. 
16: 1—5, 7—13, 21, 25, 29, is. 30. 1968. 

Doolittle h. Tiedebohl (I967) point out that nenbers of this 
genus when cultivated in the southr/festem United States need 
care in transplanting, should be pruned in January, and tend to 
remain dormant until very late in the spring . 

VITEX UNIFLORA J. G. Baker 

Additional bibliography: Jacks, in Hook. f. & Jacks., Ind. 
Kew., pr. 1, 2: 1211; (1895) and pr. 2, 2: 1211;. 19l;6; Moldenke in 
Humbert, Fl. Madag. 171;: 71, 109—111, fie 273, fig. 16 (5 & g). 
1956; Moldenke, Phytologia 6: 203~-20li. 1958; Moldenke, P.^sumfi 
157 & 1;79. 1959; Jacks, in Hook, f . b. Jacks., Ind. Kew., pr. 3, 
2: 1211;. i960. 

Illustrations: Moldenke in Humbert, Fl. Madag. 171;: 109, fig. 
16 (5 & 6). 1956. 

VITEX URGEOIATA G. B. Clarke 

Additional bibliography: Forbes & Hemsl., Fl. Sin. 2: 259. 
I89O; Jacks, in Hook. f. L Jacks., Ind. Kew., pr. 1, 2: 1213 b. 
1211;. 1895; Dunn & Tutcher, Kew Bull. Misc. Inf. Addit. Ser. 10: 
201;. 1912; Lam. & Bakh., Bull. Jard. Bot. Euitenz., ser. 3, 3: 
55. 1921; H. N. Ridl,, Fl. Malay Penins. 633. 1923; P. Dop in Le- 
comte, Fl. Indochine 1;: 826. 1935; Fletcher, Kew Bull. Lj.sc. Inf. 
1938: 1;32 & l;3l;. 1938; Jacks, in Hook. f. & Jacks., Ind. Kew., 
pr. 2, 2: 1213 & 1211;. 19l;6; Anon., Kew Bull. Gen. Index 1929- 
1956, 293. 1959; Jacks, in Hook. f. L Jacks., Ind. Kew., pr. 3, 
2: 1213 & 1211;. I96O; Moldenke, Phytologia 8: 92. I96I; Hegnauer, 
Chemotax. Pfl. 3: 39. 1961;, 

Krukoff describes this species as a tree, 75 feet tall, the 
trunk 12 inches in circumference, fruiting in Novenber. Llaterial 
has been misidentified and distributed in herbaria as Teijsmanni - 
odendron coriaceum (C. B. Clarke) Kostenn. 

Additional citations: INDONESIA: GREAT iJR SUMDA ISLAImTjS: Sumatra: 

2l;0 



1968 Moldenke, Notes on Vitex 2l«l 

Krukoff h2hh (N, W~1750656), U339 (N, W— 1750719) . 

VITEX VANSTEENISI Moldenke 

Additional bibliography: Moldenke, Biol. Abstr. 27: 3121. 
1953} G. Taylor, Ind. Kew. Suppl. 12: l5l. 1959} Moldenke, Phyto- 
logia 8: 92. 1961. 

VITEX VAUTHIERI P. DC. 

Additional bibliography: Jacks, in Hook. f. & Jacks., Ind. 
Kew., pr. 1, 1: 296 (1893) and 2: 1211| (1395), pr. 2, 1: 296 
(19U6) and 2: 121ii (I9li6), and pr. 3, 1: 296 (I960) and 2: 12lU. 
I960} Moldenke, Phytologia 8: 93. 1961. 

An isotype of this species — Vauthier 193 — deposited in the 
herbarium of the Conservatoire et Jardin Botaniques at Geneva, 
was photographed there by Macbride as his type photograph number 
7882. 

Additional citations: BRAZIL: Rio de Janeiro: Sampaio 79U9 
( Ja— niaOO) ; Vauthier 193 [Macbride photos 7882] (W— photo of 
isotype) . 

VITEX VELUTINA (Koord. & Val.) Koord. 

Additional synonymy: Vitex velutina Koord. apud Stapf , Ind. 
Lond. 6: U79. 1931. 

Additional bibliography: A. W. Hill. Ind. Kew. Suppl. 6: 219. 
1926} Moldenke, Phytologia 8: 93 (1961) and 17: 30. 1968} Molden- 
ke, R6sum6 Suppl. 16: 30. I968, 

VITEX VENULOSA Moldenke 

Bibliography: Moldenke, Phytologia k: 6U — 65 (1952) and 6: 
210—211. 1958} Moldenke, R6sum6 lU3 & U79. 1959} G. Taylor, Ind. 
Kew. Suppl. 12: l5l. 1959. 

VITEX VERMOESENI DeWild. 

Additional bibliography: A. W. Hill, Ind, Kew. Suppl. 8: 2U9. 
1933} Moldenke, Phytologia 8: 93. I96I} Moldenke, R6s\ffii6 Suppl. 
12: 6 & 7. 1965} Moldenke, Phytologia 15: 251. 1967. 

Recent collectors describe this plant as a tree, 20 n. tall, 
called "mebassa" or "mevassa", grovri.ng in forests, and fruiting 
in March. The corollas on Monteiro & Muirta 209 are described as 
having been "clear green". 

Additional citations: ANGOLA: Cabinda: Monteiro ^ Murta 209 
(Ul)} Murta 39 (Ul) . 

VITEX VERTICILUTA A. Chev. 

Additional bibliography: Prain, Ind. Kew. Suppl. 5, pr. 1, 273. 
1921 J Moldenke, Phytologia 6: 213. 1958} Moldenke, R^sum^ li|0 & 
U79. 1959} Prain, Ind. Kew. Suppl. 5, pr. 2, 273. I960. 

Regardless of whether or not one considers Chevalier's descrip- 
tion of this taxon as adequate, his binomial is apparently invali- 
dated by the Vitex verticillata of DeCandolle & Lamarck, Fl. Franc. 
2: 363 (1778), and will have to be replaced if the taxon proves to 



2U2 PHYTOLOGIA Vol. 17, no. 3 

be distinct. 

VITEX VESTITA Wall. 

Additional synor^ymy: Vitex vestita "Wall, ex Kurz" apnid Anon., 
Kew Bull. Gen. Index 1929-19^6, 293. 19^9. Vitex vestita "-Wall. 
ex Schau." apud Backer £c Bakh., Fl. Java 2: 60$. 1965^. Vitex 
vestita Vahl ex I'oldenke, R^sujn^ Suppl. 13: 7, in syn. 1966. 

Additional h emended bibliography: Bocq., Adansonia 3= [itev. 
Verbenac] 2$3. 1063; Jacks, in Hook, f , h Jacks,, Ind, Kew., pr. 

1, 2: 1213 & 12m. 139!^; C, B, Clarke in J, Schmidt, Bot. Tida- 
skr. 26: 172, I90U; A. W, Hill, Ind, Kevr, Suppl. 7: 252. 1929} 
P'ei, Mem. Scl. Soc. China 1 (3) : [Verbenac, China] 112 & llU, pi, 
22, 1932; A, W. Hill, Ind. Kew. Suppl. 9: 297. 1938; Fletcher, 
Kew Bull. Misc. Inf. 1938: U32 & U36. 1938; Worsdell, Ind. Lond. 
Suppl, 2: 500, I9UI; Jacks, in Hook, f, b. Jacks,, Ind, Kew,, pr, 

2, 2: 1213 h 121ii, 1946; Anon., Kew Bull, Gen. Index 1929-1956, 
293. 1959; Jacks, in Hook, f, & Jacks., Ind, Kew., pr, 3, 2: 1213 
& I21U, I96O; Moldenke, Phytologia 8: 93. 1961; Moldenke, Dansk 
Bot. Arkiv 23: 92, 1963; Backer «c Bakh,, Fl, Java 2: 605. 1965; 
Moldenke, R6sum6 Suppl. 13: 7. 1966; Moldenke, Phytologia 15: 
111, 1967; Moldenke, R^sumfi Suppl. 16: 30, I968. 

Illustrations: P'ei, Mem, Sci. Soc, China 1 (3): [Verbenac, 
China] pi. 22. 1932. 

Recent collectors describe this plant as a shrub or small 
tree, 1.5 — 5 m. tall, with yellowish-green fruit, groTring on 
granite, in thickets, in primary forests, and "scattered by 
stream", at altitudes of 150 — 7000 feet, called "kajoe giak 
noenik", and fruiting in January, August, and September. Backer 
k Bakhuizen van den Brink (1965) describe the plant as follows: 
petioles 3.5 — 10.5 cm. long; leaflets 3, elliptic-ovate-oblong, 
long-acuminate at the apex, short-pubescent above, villous and 
copiously gland-dotted beneath, the median leaflet 11.5 — 20 cm. 
long, U.5 — 11 cm. wide, on a petiolule 1.5 — "i'S cm. long, the 
other leaflets smaller and on shorter petiolules; cymes 1 — U in 
each axil, tawrQr-pubescent; calyx and corolla with numero\i3 
rather large yellow glands on the outer surface; corolla yellow, 
its tube about 8 mm. long, glabrous except for a ring of hairs 
rather far below the insertion of the stamens and near the base 
inside. They comment that while the species has been recorded 
from Java "no Javan localities sire known. Probably the plant was 
collected in the Bogor Botanic Garden, where it was fonnerly 
cultivated." They give its distribution as the "Western part of 
Malesia". In my I963 work the distribution is given as "India, 
Burma, Indo-China, Thailand, and llalaya, north into southern 
China, east to the Lingga Archipelago, Svunatra, Java, Borneo, 
and the Lesser Sunda Islands". 

Chand states that it is a "rare forest tree" in Assam. The 
corolla is described as "whitish" on A_. Henry 12310 , as "yellow" 
on Toroes 28U , and "with a yellow spot in throat" on Chand 3U28 . 
The Boeea 70U9 and Toroes 28U & 1236 collections are accompanied 
by wood sample at the University of Michigam. Toroes lit97 has 



1968 Moldenke, Notes on Vitex 2U3 

leaves that approach those of f . glabrescena Moldenke. The Boeea 
8501, distributed as V^ vestita, is actually V^ gamosepala var. 
kunstleri King ^ Gamble. 

Additional citations: INDIA: Assam: Chand 3U28 (Mi), 3^73 (Mi). 
BURMA: Shan States: Khalil s.n. [Laikaw, 1893] (W--369$89) . Up- 
per Burma: Kingdon-Y/ard 22$01 (Bm). CHINA: YUnnan: A. Henry 
12310 (W— U59020, W— U5902IT7 ii F. Rock 7212 (W— 17^33Fy7'THAI- 
LAND: Hansen , Seidenfaden , & Smitinand 10808 (Ac, Cp); S^rensen , 
Lars en , & Hansen $209 (Cp) . MALAYA: Selangor: Kloss 3«n. [3.8. 
19lii] (W— 2318001). INDONESIA: GREATER SUNDA ISLANDS: Banka: 
Anta 608 (A). Sumatra: Boeea 70^9 (Mi, Mi), 9827 (Mi, Mi); Toro- 
es 28U (Mi, Mi), 1236 (Mi, Mi), lli97 (Mi)j Yates I6li8 (Mi, Mi), 
211iO (Mi). 

VITEX VESTITA f . GUBRESCENS Moldenke 

Additional bibliography: Moldenke, Phytologia 3: U89 (1951) and 
8: 93. 1961. 

Soepadno calls this plant a treelet, 1; m. tall, with yellowish- 
green fruit, groiiTing in sandy-loam soil on hillsides. The Toroes 
IU97 collection, cited under typical V. vestita Wall., has leaves 
which almost approach those of f . glabrescena in their pubescence. 

Additional citations: INDONESIA: GREATER SUNDA ISLANDS: Sumatra: 
Soepadmo I8I (S) . 

VITEX VESTITA f . MILLSH (Henderson) Moldenke 

Additional bibliography: A. W. Hill, Ind. Kew. Suppl. 8: 2U9. 
1933; Moldenke, Phytologia 3: U89 (1951) and 6: 216—217. 1958; 
Moldenke, R^sum^ I8I, 386, & 1;79. 1959. 

VITEX FESTITA var. SIAMICA Moldenke 

Bibliography: Moldenke, Phytologia I4.: 65 (1952) and 6: 217. 
1958} Moldenke, R6sum6 179. 1959. 

VITEX VESTITA f . WINKLERI Moldenke 

Additional bibliography: Moldenke, Phytologia 3: U89 (1951) 
and 8: 9U. 1961. 

VITEX VILLOSA Sim 

Additional bibliography: Prain, Ind. Kew, Suppl. k, pr. 1, 2li.8 
(1913) and pr. 2, 2U8. 1958; Moldenke, Phytologia 6: 217—218. 
1958; Moldenke, Rlsum6 l5l & U79. 1959. 

VITEX VOLKENSII GUrke 

Additional & emended bibliography: Durand & Jacks., Ind. Kew, 
Suppl. 1, pr. 1, U57. 1906; Prain, Ind. Kew. Suppl. U, pr. 1, 2U8. 
1913; A. W. Hill, Ind. Kew. Suppl. 8: 2li9. 1933; Durand & Jacks., 
Ind. Kew. Suppl. 1, pr. 2, U57. 19Ul; Prain, Ind. Kew. Suppl. U, 
pr. 2, 2U8. 1958; Durand &: Jacks., Ind. Kew. Suppl. 1, pr. 3, U57. 
1959; Moldenke, Phytologia 8: 9U. 1961; Hocking, Eaccerpt. Bot. 



2l4li PHYTOLOGIA Vol. 17, no. 3 

A.I4: 592. 1962. 

VITEX VONDROZEfJSIS Moldenke 

Additional bibliography: Moldenke, Biol. Abatr. 2?: 3121. 1953; 
Moldenke in Humbert, H. Madag. 17U: 7$, 121—123, & 273, fig. 18 
(U). 1956; Moldenke, Phytologia 6: 219—220. 1958; Moldenke, R6- 
surni 157 & U79. 1959; G. Taylor, Ind. Kew. Suppl. 12: I51. 1959. 

Illustrations: Moldenke in Humbert, Fl. Madag. 17U: 121, fig. 
18 (U). 1956. 

VITEX WATERLOTI Danguy 

Additional .<k emended bibliography: A. 7^. Hill, Ind. Y.en, Suppl. 
7: 252. 1929; Moldenke in Humbert, Fl. Madag. 17li: 76, 13>-136, 
& 273, fig. 21 (3—5). 1956; Moldenke, Phytologia 6: 220—222. 
1958; Anon., U. S. Dept, Agr. Bot. Sub j . Index 15: 1U361. 1958; 
Moldenke, Rlsum^ 157, 391, ?i hl9 . 1959. 

Illustrations: Moldenke in Humbert, Fl. Madag. 17U: 135, fig. 
21 (3—5). 1956. 

VITEX VTELLENSI DeTJild. 

Additional bibliography: A. V^ Hill, Ind. Kew. Suppl. 8: 2li9. 
1933; Moldenke, Phytologia 6: 222—223. 1958; Moldenke, R^sumS 
II43 & li79. 1959. 

VITEX 'iWlUnTSCHII Gttrke 

Additional bibliography: Durand & Jacks., Ind. Kew. Suppl. 1, 
pr, 1, U57. 1906; A. W. Hill, Ind. Kew. Suppl. 7: 252. 1929; F. 
R. Irvine, PI. Gold Coast U37. 1930; Durand & Jacks., Ind. Kew. 
Suppl. 1, pr. 2, U57 (19ia) and pr. 3, U57. 1959; Moldenke, Phy- 
tologia 8: 9U. I96I; Moldenke, R6sum$ Suppl. 12: 7. 1965; Mol- 
denke, Phytologia 15: 2U6— 2U7. 1967. 

Recent collectors describe this plant as a shrub or small 
tree, 0.5 — 35 ni, tall, often rhizomatous, with green or dark 
wine-colored fruit, growing in sandy soil, in wet or cense sec- 
ondary forests, Brachystegia woods, or regenerated forests after 
cultivation, at I630 — 1700 n. altitude, flowering also in Janu- 
ary, April, and November, fruiting in April and August, and 
called "munterengue" . The corollas are said to have been "white 
with the lip blue inside" on F. A_. Mendonca 2856 , "with a violet 
lip" on Torre 2096 , and "lilac with yellow throat" on E_. £, Men- 
des 2057. Irvine (1930) remarks that Thompson 37 represents a 
species "near V. welwitschii " . Herbarium material has been mis- 
identified and distributed as V. grisea J. G. Baker. 

Additional citations: COJJGOTeOPOLDVILLE : Louis U21 (B), 2l63 
(B). ANGOLA: Benguela: Gossweiler 12185 (Ul) . Cabinda: Lionteiro , 
Santos , & Murta 259 (Ul) . Huambo: £. £. Mendes 566 (Ul) . Huila: 
Antunes or Dekindt s.n. (Ul) ; E. J^ Mendes 20^ (Ul, Z) ; R. San - 
tos U58 (Ul). Luanda: Teixeira 10 (Ul) . PORTUGUESE EAST AFPJCA: 
Lourencjo Marques: F. A. Mendonca 2356 (Ul); Torre 2096 (Rf, Ul), 
6la7 (Ul), 7678 (Uiy. 



A NOTE ON BAUKENIA HAGENBECKII HARMS 
R. P. Wunderlln * 



Bauhlnla hagenbeckil Harms is an interesting and variable 
species which occurs in the Cache region of Paraguay and Brazil. 
The Cacho is an arid region conposed of alluvial soil of un- 
consolidated sands and clays which supposrts a vegetation com- 
posed of thickets of thorny scrub trees and openings of course 
grasses. Until this study was conducted, this species was 
known only from photographs of the type specimen. The type 
collection was made in "Gran Cacho, Brazil" by Hagenbeck in 
April, 1895. The only known existing type was destroyed by 
allied bombs and fire during World War II. A photograph and a 
fragnent (4 leaflets) of the type housed in the Field Museum of 
Natural History were examined by the author. This material is 
therefore designated as the lectotype of the species. 

Bauhinia hassleriana was described by Chodat in 190^ , one 
year after B. hagenbeckii , from the Cacho region of Paraguay. 
It was segregated into four forms (f . angustifolia , f . inter- 
media, f . latifolia , and f . acuminata ) and a variety (var. mar- 
ginata ) by Chodat and Hassler in the same paper. Several of 
these taxa are known to the author only from the anple type ma- 
terials and others from the description only. The forms angust- 
ifolia , intermedia , and latifolia were segregated on the basis 
of leaf width, the dimensiona of which apparently were arbitrar- 
ly selected and overlap on the type material examined by the 
author. Forma acuminata is differentiated by having the lobes 
of the leaflets acuminate, but this also is too variable for 
formal taxonomic designation. Variety marginata is different- 
iated by having the leaflets distinctly marginate. This char- 
acter is also not clear-cut and quite evident margins are found 
on other taxa of the species as well as on the type material of 
B. hagenbeckii . 

In vegetative and floral characters B. hassleriana is not 
separated from B. hagenbeckii and is best considered as being 
conspecific with it. 

The following is the synonymy of B. hagenbeckii and a de- 
scription of the species as I know it: 

BAUHINIA HAGE^BECKII Harms, Engl. Bot. Jahrb. 33- Beibl. 72:21. 

1903. (T: Hagenbeck s.n.I). 
Bauhinia hassleriana Chod. in Chod. & Hassl. Bull. Herb. Boiss. 

ser. 2. 4:690. 1904, ex char. 
Bauhinia hassleriana Chod. fonna acuminata Chod. & Hassl. Bull. 

Herb. Boiss. ser. 2. 4:690. 1904. (T: Hassler 7076!). 



2h$ 



2U6 P H y T L G I A Vol. 17, no. 3 

Bauhlnla hasslerlana Chod . fonna angustl folia Chod . & Hassl . 

Bull. Herb. Boiss. ser. 2. ^:690. 190U, ex cJ-iar. 
Bauhlnla hasslerlana Chod. forma Intermedia Chod. & Hassl. Bull. 

Herb. Bolss. ser. 2. 4:690. 1904. (T: Hassler 7898 .'). 
Bauhlnla hasslerlana Chod. forma latl folia Chod. & Hassl. Bull. 

Herb. Boiss. ser. 2. 4:690. 1904. (T: Hassler 7656 ). 
Bauhlnla hasslerlana Chod, var. marginata Chod. & Hassl. Bull. 

Herb. Bolss. ser. 2. 4:690. (T: Hassler 6958 ;). 

A much branched shrub or small tree. Branches pubescent, 
soon glabrescent. Leaves bifollolate; leaflets obliquely ovate 
to linear-oblong, apices obtuse to subacute, 2-6 cm long, 0.5- 
2.0 cm wide, upper leaflets generally narrower than lower, 
puberulent or subglabrous below, usually pubescent near petiole, 
glabrous above, 1-to 4-nerved (depending on width of leaflets), 
veins conspicuously reticulate below, less so above, margins 
often distinct; petioles slender, 1-2 cm long, puberulent to sub- 
glabrous. Flowers solitary to few in axillary clusters, appress- 
ed puberulent; buds 5-6 cm long at maturity; calyx cylindrical, 
tube 2.5-4.0 cm long, lobes 2.5-3-5 cm long, splitting and re- 
flexed at maturity; petals lanceolate to elliptic-lanceolate, 
clawed, 2.0-2.5 cm long, 6-8 rm wide, white; anthers 10, alter- 
nately long and short, filaments glabrous, 2.0-2.5 cm long; 
pistils slightly longer than stamens, glabrous or sparingly 
pilose; pedicels 10-12 mm long. Mature legume dimensions un- 
known, light tan, dehiscent. 

Type: Hagenbeck s_.n. (F) from "Gran Cacho, Brazil" is select- 
ed as the lectotype. 

SPECIMENS EXAMINED: BRAZIL: "Gran Cacho", Hagenbeck s.n. (F- 
type; photo-US, MO). PARAGUAY: Amambay: In rocks along Rio Apa 
near Bella Vista, Hassler 7898 (^D, MICH, US, UC, F) . Boqueron: 
In sand along bank of Rio Yacare, Hassler 7076 (F, GH, UC). 
Cordilleras: In dry rocky area near Valenzuela, Hassler 6958 
(photo-MO, US, F; MICH, F, MO, UC). 



* Department of Botany, Southern Illinois University, Carbondale. 



A NEW SPECIES OF PIRIQUETA AUBLET (TURNERACEAE) 
FROM MATO GROSSO, BRAZIL 

Carlos Alberto F. de Moura 
Instituto de Botaclca, Sao Paulo 

Plrlqueta corumbenels C, Moura sp. ncrr, - Herba caule 25 - 
35 cm alt., cyllndrico, alinpllci v. pauclramoso, pilie stellari- 
bus (radius centralis longlssimus, radlls basallbus brevloidbus ) 
flavo-a\irel8, dense vestito; pills slnpllclbos brevlsslmls te- 
nulbus, albldls, Intennlxtis. Stlpulae nullae. Folia pet lolls 
2 - 4, mm longls; lamina elllptlca v, fere late lanceolate, 3 - 
6 cm longa, 2 - 3 cm lata, 1.1/2 - 2-plo longiora quam latlora, 
basl obtusa v. subrotundata, aplce obtusa, In ambltu crenata ba> 
sl excepta, undlque pllosa, supeme pills plurlradlatls, radio 
Intermedlo malore, flavescentibus, Infeme denslsslne pllosa, pi- 
lls brevlorlbus, multlradlatls, radio Intermedlo non v, vlx lon- 
glore, pallldo-f3avescentlbus, utrlmque ad nervos pllosa, nervls 
supeme Impressls, Infeme prominent Ibue . Flores axlllares soll- 
taril, dollchoetyll. Pedunculi longlsslml, 35 - 50 mm longl, pe- 
dlcelll 5 - 8 mm longi; bneteolae nullae. Calyx 15 - 17 ran lon- 
gus, vlx In 1/4. alt. ooalltus, extus dense hirsutus, flavo-aureus ; 
tubus calyclnus obconicus, Intus glaber; lobl lanoeolatl bravl- 
aplculatl, tri- v, pentanervlbus. Petala calycem vlx ca. 1 mm 
longe superantla, vlolacea, glabra, obovata, 13 - 1^ nsa longa, 7 

- B ma lata, basl obtusa, aplce rotundata v. subtznancata, sub In- 
sertlone in calycis tubun marglnlbus decurrentla; corona vlx 1 
mm longa. In parte anterlore lac erato-fixobr lata. Fllamenta basl 
vlx 1 mm longe tubo calyclno adnata, glabra, 4. - 6 mm longa; an- 
tharae effloratae lancoolatae, 2 - 2,5 mm longae, dorso in I/3 
alt. afflxae. Ovariun ovatum, 2,5 mn longun, dense pllosun, 30- 
ovulatum. Still glabri, erectl, 3 - 5 mm longl, aplce breve 4f- 
partltl; stigmata multiflagellata digltata, 1 - 1,3 nm longa. 
Fructus globulosus, pilos\is, 6 nm longus, 6 on dlametro, dorso 
sub pube tuberculatus . Semlna Immatura. 

Holotype in the Jardim Botanlco do Rio de Janeiro, accession 
n« 8^7H» collected at Fazenda Aguassuzlnho, Municipality of Co- 
rumba, State of Mato Grosso, Brazil, October 17, 1953, by E, Pe- 
relra, W, Egler & Graziela Barroso 388. 

This specimen is similar to Plrlqueta aurea (Camb. ) Urban. 
However, a more detailed examination shows that the specimen in 
question has certain characteristics which distinguish it from 
that species. Thus, in P. aurea. the hairs of the stem and leaves 
are identical in being irregularly stellate with all of the rays 
well-developed; the peduncles do not exceed 27 mm in length; the 
petals are oblanceolate-ouneate and surpass the calyx lobes by 8 

- 15 mm; the style is divided irregularly to form the thin stig- 
matic branches. In the new species the stem hairs possess a cen- 
tral ray much Iwiger than the other rays; these other rays remain 
at the base as a low crown surrounding the central ray. The leaf 
hairs are shorter than the stem hairs. The hairs of the upper sur- 
face of the leaf resemble those of the stem in having a long cen- 

2U7 



2b8 



PHYTOLOGIA 



Vol. 17, no. 3 



tral ray with short basal rays, whereaa thoea of the lover surfa- 
ce have a shorter central ray and a larger number of basal rays. 
The peduncles are very long, 35 - 50 am. The petals are obovate 
and do not surpass the calyx lobes by Bore than 1 nt. The style 
Is divided at the tip Into /t. regular short branches; each of these 
then divides into A - 5 digitate stlgBatic branches. 

Because of the above differences I an describing it as a nev 
species, naming it for the municipality in which it was collected. 

I wish to extend my thaclcB to Dr. George Elten who reviewed 
the English text and to Dr. Gerhard Gottsberger for Inking in my 
drawings. 







Fig, 1, Piriqueta coruabwisia . Dlagraaatic drawing of a flower 

showing the relative length of corolla and calyx lobes. 

Fig. 2. Pistil with hairy ovary, 3 styles and rtlgmatlc branches. 

Fig. 3. Stem hair. 

Fig. ^. Hair on lover face of leaf. 

Fig. 5. I«af showing venation. 



1u,-o 



6L< RR 

l^.i PHYTOLOGIA 

Designed to expedite botanical publication 



Vol. 17 October, 1968 No. 4 

V 

CONTENTS ' ^^^^ 

NEW YO.RK 

REED, C. F., /^/r/e.v TAe/y/7/er/V/s ANICAL GARDC43 

LITTLE, E. L., Jr., Two new pinyon varieties from Arizona . . . .329 
DEGENER, O. & L, The eruption in Hiiaka Crater, Island of Hatiaii .343 
MOLDENKE, H. N., Notes on new and noteworthy plants. LI . . . .344 

MOLDENKE, A. L., Book rei lews 345 

MOLDENKE, H. N., Additional notes on the Eriocaulaceae . XIII . .348 



Published by Harold N. Moldenke and Alma L. Moldenke 

303 Parkside Road 
Plainfield, New Jersey 07060 
U.S.A. 

Price of this number, $1.50; per volune, $6.75, in advance, 

or $7 at close of volume . 



INDEX THELYPTERIDIS 
Clyde F, Hoed * 



Thelypteria Schmidel has bean proposed for conservation by 
Holttum (1968) over Thelypterls Adanson (» Pberis). In so doing he 
has presented a brief historical outline of the problems Involved 
in establishing Thelypteris Schnddel, Femald and Weatherby (1929) 
early presented the pros and cons regarding Schmidel 's publication 
of Thelypteris and gave reasons why the name was legitimately pub- 
lished as a generic name* Schott (1834) was the first to use the 
binomial Thelypteris palustrls. thus establishing the genotype. 

Ferns with "thelypteroid" characteristics have been descri- 
bed originally in many different genera, mainly In Nephrodium. As- 
pidium. Polypodlura. Dryopteris. Lastrea. Cyclosorus. Meniscium. 
C^nnno gramma. Abacopteris. Phegopteris and Stegnogramma . They have 
been transferred to Dryopteris . Thelypteris. Phegopteris. Lastrea. 
Cyclosorus. or have also most recently been put into niunerous new 
genera by Chlng (I963). 

Nieuwland (1910) presented Dryopteris as a synonym of Thely- 
pteris and thereby started a long line of confusion. Not only were 
palustrls. simulata and noveboracensis put into Thelypteris. but 
also fragrans. marglnalis. filix-^nas. goldlana. bootli . cristata 
and spinulosa were added, Chrlstensen (1913 and 1920) in his Mono- 
graph of the genus Dryopteris treated the genus as a large one, in- 
cluding in it those ferns with "thelypteroid" characteristics, 

Chlng (1936), in his Revision of the Genus Dryopteris in the 
Sikkim-Hlmalayan Region, accepted the ferns with "thelypteroid" 
characteristics as a separate genus, Thelypteris. and included in 
it species described la Lastrea. Glaphyropt eri s . Leptogramma and 
other genera. Thus he made Thelypteris a large genus, describing 
many new species and transferring about 80 species into the genus 
at that time. 

Chrlstensen (1938) In the Manual of Pterldology recognized 
the essential differences between the dryopteroid and thelypter^ 
oid ferns by dividing his subfamily (XII) Dryopteridoideae into 
two tribes: Dryopterideae and Thelypterideae, In the Thelypterl- 
deae he included Thelypteris Schmidel ( Lastrea Bory, Dryopteris 



* Reed Herbarium, Baltimore, Maryland; Research Botanist and Plant 
Kxplorer for United States Department of Agriculture; Collaborator 
in Department of Botany, amithsonian Institution, 

2h9 



250 P H Y T L G I A Vol. 17, no. h 

part, auctt.) with about 500 apeciea, Phegoptftrla Pee (- Oymno- 
carplum Newman), Glaphyropteria Presl, Stclropteris C.Chr., Para - 
polystichum Keyas., Pterldrys C.Chr. et Ching, Monachosorum Kunze, 
Monacho sorella Hayata, Cyclosorus Link (Meniecium Schreber, Lcpto- 
jyamma J.Sm. , Stegnoj^ranmia Blume, Sphaerostephanoa J.Sm. ) and 
Goniopti^rla Preal. 

H. Its (1939) emended Thelyptaria Schmidel and divided the 
genua into aeveral sections: Suthelyptcris H. Ito (Type: Th. palu- 
atris Schott), Parathelypteris H. Ito (Type: Th, glanduligera Ching), 
Metathelypteris H. It6 (Type; Th. gracilescens Ching) and Macro- 
thelypteris H. Ito (Type: Th, oligophlebia Ching). However, he main- 
tained the genera Glaphyropteria Presl (Type: Ca. . decuasata Preal), 
with aect. Euglaphyropteria H. Ito and aect. Cyclogramma H. Ito (Gl. 
similans H. Ito); Phegopteria Fee (Type: Ph. polypodioidea Fee), 
with sect, Euphegopteris Christ and sect. Lastrella H. Ito (Type: 
Fh, decuraive-pinnata F*e)j Gymnocarpivun Newm. (Type: G, dryopteria 
Newm. ). vd.th sect. Eugymnocarpium and sect. Currania Ching (Type: 
Gynmocarpium gracilipes Ching); Leptograimna J.Sm. (Type: L, totta 
J.5n. ); Cyclosorus Link (Type: C, gongylodea Link) and Heniscium 
Schreb, (Type: M» reticulatum Swartz), with sect, Sumeniscium H,Ito 
and sect. Goniopteridopsis H. Ito (Type: M, urophyllum H. Ito). 

Ching (19A0) established the family Thelypteridaceae, with 
twelve genera and about 800 species. He divided the family into 
three tribe 3: Thel^terideaq, Goniopterideae and Dictyoclineae , In 
the tribe Thelypterideae he placed Thelypteris Schmidel (Type: Th. 
palustris Schott). Lastreopsia Ching (Type: L, recedena (J.Sm.) 
Ching j. t^odematium Kunze (Type; H, crenatum (Forsk.) Kuhn), Glaphy - 
ropteris Presl (Type: Gl , decuasata (L.) Presl), Parapolystichuzn 
(Keyserling) Ching (Type; P. effusum (Sw.) Qiing) and Leptogramma 
J, So,, emend. Ching (L, totta (Willd,) J.Sm. ). In the tribe Gonio- 
pterideae he placed Cyclosorus Link (Type: C. gongylodes (Schkuhr) 
Link), Stegnograama Blume (Type; S, aspidioides Blume). Goniopteris 
Preal (Type: G, crenata (Sw, ) Presl), Abacopteris Fee, emend, Ching 
(Type: A, lineata (Blume) Ching) and Meniscium Schreb. (Type; M, 
reticulatum (L.) Swartz). In the tribe Dictyoclineae he placed 
Dictyocline Moore (Type; D, griffithii (Hook, et Thorns.) Moore. He 
pla ced Sdhaero at ephano a J.Sm. vType: S. polycarpa (Blume) Copel.) 
in the monotypic family Sphaerostephanaceae ChiBg. 

Copeland {I9h7) in Genera Filicxim established the family As- 
pidiaceae (S.F.Gray) Copel,, including in it several families estab- 
lished by Ching (1940), as Dictyoxiphiaceae, Thelypteridaceae, 
Sphaerostephanaceae, Woodsiaceae, Ifypoderriaceae, Didymochlaenaceae, 
and Elaphoglossaceae. On the basis that Thelypteris Schmidel was 
invalidly published, Copeland took up Laatrea Bory and transferred 
many species to this genus, using Polypodium oreopteris Elirh, as 
genotype. This arrangement also permitted Lastrea thelypteris (L,) 
Bory to become usable. The concept of Lastrea presented by Copeland 



1968 Reed, Index Thelypteridis 2^1 

included Leptogramma J.Sm, (Type: L, totta (Willd.) J.Sm,), Amauro- 
pelta Kunze (Type: A, breutelii Kunze). ^aphyropteris Presl (Type: 
(XL , d ecu 3 sat a (L.) Presl) , Phegopteris ?6c (Type: Pblypodium poly- 
podioldes L«). Oochlamys Fee (Type; 0, rivoirel Fee). Steiropteris 
C.Chr., Cyclogramma Tagawa (Type: Thelypteris simulans Ghing). 

Copeland in Genera Filicure also maintained the follovdng gen- 
era: Cyclosorus Link, including Abacopteris Fee (Type: £, gongy- 
lo.de s (Schkuhr) Link) and transferred many species into this genus; 
Currania Copel. (Type: C, gracilipes Copel.)j Ampelopteris Kunze 
(Type: A, elegans Kunze); Sphaerostephanos J.Sii. in Hook, et Bauer 
(T^e: S. asplenioides J . Sm. ) ; Stegnogramma Blume (Type: S, as- 
pidioides Blume); Ctoniopteris Presl (Lectotype: G, viyiparum ( Raddi ) 
Brack.); Meniscium Schreber (Type: M. reticulatum (L.) Swart z)j and 
Dictyocline Moore (Type: D. griffithii Moore). 

Pichi-Sermolli (1953) argued that since the legitimacy of 
Thelypteris Schmidel was uncertain, the best solution was to give 
up the name and to protect Lastrea versus Thelypteris. Fuchs (I963) 
also proposed using Lastrea, using L, thelypteris (L.) Presl and L. 
limbo sperma (Allioni) Holub et Pouzar apud Holub (I96I) as repre- 
sentatives (Flora of Hungary). Fuchs also maintained the generic 
names Dryopteris. Gymnocarpium and Riegopteris , 

Holttum (195^) distinguished four genera in the family Thely- 
pteridaceae: Thelypteris . Cyclosoin is, Abacopteris and Ampelopteris . 

Ching (1963) in his reclassification of the family Thely- 
pteridaceae divided the family into the three tribes he had estab- 
lished in 1940 and then subdivided these into subtribes, allowing 
for 18 genera, transferring the Asiatic mainland species previously 
(1936) placed in Thelypteris . Many of the sections Ching elevated 
to genera, especially those of H. ItS (1939), leaving only three 
species in Thelypteris . 

Tribe Thelypterideae Ching (1940): Subtribe Thelypteridi- 
nae Ching: Thelypteris Schmidel, Lastrea Bory, Parathelypt eri s 
(H.Ito) Ching, Metathelypteris (H.ItS) Ching and Hypodematium 
Kunze; Subtribe Phegopteridinae Ching: Macrothalypteris (H.Ito) 
Ching, Phegopteris F6e, Pseudophegopteris Ching, Cy do gramma 
Tagawa, and Leptogramma J.Sm. 

Tribe Goniopterideae Ching (1940): Subtribe Pseudocyclo- 
sorinae Ching: Qlaphyropteria Ching, Pseudocyclosorus Ching, 
and Mesoneuron Ching; Subtribe Cyclosorinae Ching: Cyclosoirus 
Link and Stegnogramma Blvune; Subtribe Goniopteridinae Ching: 
Ampelopteris Kunze; Subtribe Menisciinae Ching: Abacopteris Fee, 
emend, Ching. 

Tribe Dictyoclineae Ching (1940). Dictyocline Moore. 



252 PHYTOLOGIA Vol. 17, no. h 

Morton (I963) on the other hurid widened the concept of Thiejj- 
pterla Schmldel by including the generic segregates proposed by 
Ching, Copeland, Aloton, Holttum and Tardieu-blot up to that tin*. 
Therefore, Morton's ultimate concept of Thelypteris is niich like 
that of CJiing in 1936. However, Morton divided the genus into the 
following subgenera and sections. 

Thelypteris Schmidel is divided into three subgenera: 
Subgenus Thelypteris (Lectotype: IJL* palustris Schott)j Sub- 
genus Lastrea (Bory) Alston (Type: Polypodium oreopteris SJirh.), 
with sect. Lastrea and sect, Qlaphyropteris (Pre si) Mjorton (Type: 
Polypodium deeusaatum L. );and subgenus Cyclosorus (Link) Morton 
(Type: Aspidium gongylodes Schkuhr), with sect, Gyclosorus. 
sect, Steiropteris (C.Chr.) Morton (Lectotype: Polypodium del- 
toldeum Swartz) .sect. Leptograuna (J.Sm, ) Morton (Lectotype: 
L» totta J.Sra, « Th, pozoi (Lagasca) Morton), sect. Goniopteris 
(Pre si) Ntorton (Lectotype: Polypodium crenatum Swartz = Th. 
poitiana (Bory) Proctor^ and sect, Meniscium (Schreber) Morton 
(Tyioe: Polypodium reticulatum L, ). 

Momose made extensive studies of many genera of ferns and of 
species-complexes within genera. His studies dealt mainly with the 
gametophjrte stage. His studies are reflected in the Studies on the 
Gametophyte of Feme, I-XXIX, published in the Journal of Japanese 
Botany, volumes 13-18, 1937-1942, The prothalli of the Thelypter- 
oid ferns are reported in parts XVI and XVII (1941). Most of the 
species studied are those found in Japan. 

Iwatsuki (1964-1965) presented his classification of Thely- 
pteris Schmidel, based upon his studies of the thelypteroid ferns 
of Japan and adjacent regions. His studies have been quite exten- 
sive and quite thoixiugh. He recognizes the following genera in 
the Thelypteridaceae: Stegnogramma Blume, with three sections; 
Thelypteris Schmidel, with 14 subgenera; and Meniscivim Schreber, 
with 4 sections, 

1. Stegnogramma Blume (1828). (Type: St, aspidioides Bl.). 

a. Sect. 1, Leptogramma (J.Sm.) K.Iwats. (I963) (Type: 
Polypodium tottum Willd., non Thunb, = St. pozoi 
(Lagasca) K.Iwats. ). 

b. Sect, 2. Stegnogramma . (Type: St, aspidioides Bl,). 

c. Sect. 3. Dictyocline (Moore) K.Iwats, (1963 ). (Type: 
St. griff it hii (Moore) K.Iwats.), 

2, Thelypteris Schmidel (I763). (Type: Aero sti chum thely- 

pteris L . ). 
a. Subgen. 1, Phegopteris (Presl) Ching (1936). (Type: 
Polypodium phegopteris L. ), 

1, Sect, 1, Phegopteris . (2 species). 

2, Sect. 2, Ustrella (H.Ito) K.Iwats. (I964) . (Type: 
Polypodium de cursive-pinnatua van Hall) ( 50 species). 



1968 Reed, Index Thelypteridis 253 

b. Subgen. 2. Cyclogramma (Tagawa) K. Iwats. (1964). (l^rpe: 
Thelypteris simulans Ching = Th» auricula ta) . (7 species), 

c. Subgen. 3. Thel vpteris . (Lastrea Bory^ Types Poly- 
podium oreopteris Ehrh.). 

1. Sect. 1. Metathelvpteria H.Ito (1939). (Typ«s Aspi- 
dium gracilescens Blurae). (lA species), 

2. Sect. 2. Thelypteris . (Type: Acrostichum thelypteris 
L. = Th, palustris Schott). (300 species). 

d. Subgen. 4. Cyclosoriopsis K, Iwats. (I964). (Type: 
Polypodiuin dentatum Forsk.). (150 species). 

e. Subgen, 5. Cilaphyropterls (Presl) Alston (1958). (Type: 
Polypodlum decussatum L. ). (110 species). 

f. Subgen. 6. Glaphyropt eridop si s (Ching) K. Iwats, (I964). 
(Type: Polypodium erubescens haUL. ex Hook.), (20 

specieiy^ 

1. Sect. 1. Glaphyropt eridopsi 3 . (3 species). 

2. Scot. 2, Hesoneuron (Chingr"K. Iwats. (Type: Aspi- 
dium crassifoliun Blume). (10 species), 

3. Sect. 3. Neocyclosorus K, Iwats. (I964). (Type: 
Aspidium heterocarpon Blume). (Several species). 

g, Subgen, 7. Steiropteris (C.Chr.) K, Iwats. (1964), 
(Type: Polypodiuin deltoideum Swartz), (I3 species), 

h, Subgen. 8, Cyclosorus (Link) Morton (I963), (Type: 

Aspidium goggilodus Schkuhr. (100 or more species), 
i. Subgen, 9. Sphaero st ephano s (J,Sni, ) K. Iwats. (I964). 

(T^e: Sph , asplenioides J.Sn, = Th, polycarpa (Blume) 

K, Iwats. (6 species), 
j. Subgen. 10. Haplodictyum (Presl) K. Iwats. (1964). 

(Type: H. heterophyllum Presl, 1849). 
k, Subgen, U, Pneumatopteris (Nakai) K. Iwats. (1964). 

(Type: Aspidium callosum Blume), 

1. Sect, 1. Pneumatopteris , (20 species), 

2. Sect, 2. Macrccyclosorus K. Iwats. (I964). (Type: 
Aspidium megaphyllum Mett. (20 species), 

1. Subgen. 12. Abacopteris (Fee) K. Iwats, (I964). (Type: 

Aspidium line at um Blume, 1828), 
m. Subgen. 13. Dimorphopteria (Tagawa et K.Iwats.) K. 

Iwats, (1 species), 
n, Subgen, 14, Cyrtomiopsis K, Iwats, (I964), (Type: 

Aspidium boydiae Eaton)" (1 species), 
3. Meniscium Schreber (1791). (Type: Polypodium reticulatum 
L. (80 species) . 

a. Sect, 1, Asterochlaena (C.Chr,) K.Iwats. (I964). 
(Type: Polypodium reptans Cknel.), (50 species), 

b. Sect. 2, Cioniopteris (Presl) K,Iwats. (I964). (Type: 
Polypodium crenatum Swartz = Meniscium poiteanum 
(Bory) K.Iwats.) (20 species). 

c. Sect, 3. Ampelopteris (Kunze) K, Iwats. (I964). (Type: 
A, elegans Kunze = Meniscium proliferum (Retz,) Sw, ), 

d. Sect, 4, Meniscivun , (12 species). 



25U P H Y T L G I A Vol. 17, no. U 

ThelTptiridacsae may or may not b« a natural family of 
ferns separate from the rest of the dryopteroid ferns. Perhaps it 
is still best to treat the thelypteroid ferns as a subfamily of 
the Aspidiaceae. Christensen had proposed Subfamily Dryopteridoldeae 
Tribe Theljrpterideae (1938) and Copeland placed the family Thely- 
pteridaceae Ching in the Aspidiaceae Copel. (1947. 

Aspidiaceae subfam, Thelypteridoldeae Reed, subfam. nor. 
Basionym: Polypcdiaceae subfam. Uryopteridoideae tribe Thely- 
pterideae Christensen, Manual of Pteridology, 544. 1938; 
Thelypteridaceae tribe Thelypterideae Ching, Sunyatsenia, 5s 
238. 1940; Acta Phytotax. Sinica, 8(4): 295. 1963. Type 
genus: Thelypteris Schmidel. 

Whether the Theljrpteridaceae or the subfamily Thelypteridoldeae 
is composed of many genera as proposed by Ching (1963), or by a few 
genera as proposed by Iwatsuki (1964-1965), or by one large genus, as 
originally proposed by Ching (1936) and as emended and enlarged by 
Morton (1963), due to the wide range of variability of venation, of 
sporangial and indusial characteristics and of frond shape, and due 
to the gradual intergradation of all segregates as more species are 
put into Thelypteris. it seems to this author best to treat Thely- 
pteris as one large genus '.vith numerous subgenera and sections. 
Iwatsuki's treatment presents the genus in the most coherent form. 
However, I do agree with >'x)rton (1963) and Ching (1963) that the 
Meniscium-Goniopteris-otefinogranma- conplex belongs with the Cyclo- 
3orus- co!aplex and place them as additional subgenera or sections 
vinder Thelypteris . 

Thelypteris subgen, Stegnogramma (Bl'ime) Reed, stat, nov, 
Basionym: Stegnogramma Blume. Enum. PI. Jav. , 172. 1828, (Type: 
St . aspidioides Blume). 

Thelypteris subgen. Leptogramma (J.3m. ) Reed, stat. nov. 
Basionym: Leptogranma J. Sra., Joum, Bot., 4: 51. 1841. (Type: 
Polypodium totta Willd,, non Thunb, = Th» pozoi (Lagasca) Morton. 
Syn,: Thelypteris subgen. Cyclosorus sect, Leptogramma (J.Sm, ) 
Morton, Amer. Fern Joum., 53: 153. 1963. 

Thelypteris subgen, Dictyocline (Moore) Reed, stat, nov, 
Basionym: Dictyocline Moore, Card. Chron., 1855: 854. 1855. 
(Type: D. griffithii Moore). 

Thelypteris subgen. Meniscium (Schreber) Reed, stat. nov. 
Basionym: Meniscium Schreber, in Linn. Gen. PI,, ed VIII, 2: 
757. 1791, (Type: Polyoodium reticulatum L. ). Syn,: Thelypter is 
subgen. Cyclos orus sect. Meniscium (Schreber) Morton, Amer, Fern 
Joum., 53: 154. 1963. (80 species). 

sect, 1. Meni scium, (12 species), 

sect, 2, Asterochlaena (CChr.) Reed, stat, nov. Basionya: 
Dryopteris subgen, Goniopteris sect, Asterochlaena C.Chr., 
Biol. Arb. til. Eug. Warn., 84, 1911, (Type; Polypodima 
reptans anel., 1791). (50 species), 
sect, 3, Goniopteris (Presl) Reed, stat. nov, Basionym: 
Goniopteris Presl, Tent. Pterid,, 181. 1836, (Type: 



1968 Reed, Index Thelypteridis 2$$ 

Polypodium c renatum S wartz = ^. poiteana (Bory) Proctor), 
Syn.: Thelypterls subgen, Cyclosorus sect, Gonioptaris 
(Presl) >k)rton, Amer, Fern Joxim,, 53: 154. 1963, p.p. 
(20 species), 
sect, 4. Afflpelopterla (Kunze) Reed, stat. nov, Basionym: 
Ampelopterls Kunze, Bot, Zeit., 6: 114. 1848. (Type: 
A. elegans Kunze), (1 species). 



Most of the fossil species which belong in Thelypteris have 
been described in the genera Aspidium. Goniopteris, Phegopteris. 
Lastrea ( La straea ), Cyclosorus or Dryopteris . The most comprehen- 
sive papers dealing with fossil ferns of this complex are those by 
Alex. Braun (Ueber Fossile Qoniopteris- Arten. Zeitschr. Geol. Ges,, 
4: 553-556, 1852) and Ettinghausen (Die Famkrttuter der Jetztwelt, 
160-203, illus, I865). The species are mainly based on venation 
of fragments of pinnae. Many have proven to belong to other genera. 
However, there are some which have sporangia or/and indusia which 
prove their affinity to Thelypteris, for which reason they are 
transferred into The3:jrpteris here. 



THELYPTERIS Schmidel 



Thelypteris Schmidel, Icon. PI., ed. J.C.Keller, 45, t. 11, I3, 
Oct. 1763; Schott, Gen. Gil., ad t. 10. 1834 j emend. H. It^, 
in Nakai et Honda, Nova Flora Japonica, Polypodiaceae, Diyo- 
pteroideae, I: 123. 1939; Morton, Amer. Fern Joum., 53(4): 
153-154. 1963; Ching, Bull. Fan Mem. Inst. Biol. Bot., 6: 250, 
1936; Holttum, Taxon, 17(3): 330. I968, gen. conserv, propos.; 
non Thelypteris Adanson, Fam. des Plantes, 2: 20. July-Aug. 
1763 = Pteris). Type: Polypodium thelypteris L. * Thelypteris 
palustris Schott. . 

Meniscium Schreber in L. Gen. PI., ed. 8, II: 757. 1791. Type: 
Polypodium reticulatum L. = Thelypteris reticulata (L.) 
Proctor. 

Lastrea Bory, Diet. Class. Hist. Nat., 6: 588, 1824. Type: Poly- 
podium oreopterig Ehrh, = Thelypteris limbospenna (All.) 
Fuchs. 

Stegncgramroa Elume, Enum. PI. Jav., 172. 1828, Type: Stegnogram- 
ma aspidioides Blume * Thelypteris aspidioides (willd.) Tiyon. 

Cyclosorus Link, Hort, Berol., 2: 128. 1833. Tjrpe: Aspiditia go^- 
gilodus Schkuhr = Thelypteris totta (Thunb.) Schelpe. 



2^6 P H Y T L G I A Vol. 17, no. U 

Gonlopteris Presl, Tent. Pt^rid., 181, 1836. Type: Pclyrodlua 

YJvlcaruni Raddi - Thelypt-rirj vlvlpara (Raddl) htr'i. 
Spha^ro at cphano s J. Smith in Hook, et Baker, Oen. Fil., 21. 

1839. Type: SphaeroateLhanoa a3plenioid''3 J. Smith - Thely- 

pterie polycai-pa (Blume) K, Iwats. 
Mesochlaena R, Brown ex J. Smith, Journ, Bot,, 3^ 18. 1840, 

Type: Sphaero Stephanos asplenioides J. Smith ■ Thelypterie 

polycarpa (Blume) K, Iwat s , 
Leptogramma J. Smith, Journ. Bot., 4: 51. 1841. Type: Polypod- 

ium tottum '*/illd., non Thunb, » Thelypteris pozoi (Lagaeca) 

ttorton. 
Amauropelta Kunze, Famkr,, 1: 86. 1843. Type: Amauropelta 

breutelii Kunze = Thelypteris lirabata ( Swart z) Pi*octor. 
Abacopteris Pee, Congr, Sci. France, X, 1: 178. 1843. Type: 

Aspidium lineatum Blume - Thelypteris lineata (Blume) K. Iwats. 
Ampelopteris Kunze, Bot. Zeit., 6: 114. 1848. Type: Ampelo - 

pteris elegans Kunze = Thelypteris prolifera (Retz) Reed. 
CELaphyropteris Presl, Abh. Bohm. Ges. Wiss., V, 5: 344. 1848. 

Type; Polypodium decussatum L. - Thelypteris decussata (L.) 

Proctor. 
Haplodictyum Presl, E^im. Bot., 50. 1849. Type: Haplodictyum 

heterophyllum Presl « Thelypteris heterophylla (Presl) K. Iwats. 
Pronephrium Presl, Epim. Bot., 258. 1849. Type: Aspidium line- 
atum Blume - Thelypteris lineata (Blume) K. Iwats, 
Phegopteris (Presl) Fee, Gen, Fil., 242. 1852. Type: Phegopteris 

polypodioides Fee - Thelypteris phegopteris (L.) Slossen ex 

Rydb. 
Oochlamys Fee, Gen. Fil., 297. 1852, Type: Oochlamys revoirei 

Fee = Thelypteris opposite (Vahl) Ching. 
Hemestheum Newman, Phytologist, 4: app. XXII. 1851. Type: Poly- 
podium thelypteris L, = Thelypteris palustris Schott. 
Dictyocline Moore, Gard, Chron,, 1855: 854. 1855. Type: Dictyo- 

clina. griffithii Moore = Thelypteris griff ithii (Moore) Reed. 
Pneumatopteris Nakai. Bot. Mag, Tokyo, 47: 179. 1933. Type: 

Aspidium callosua Blume » 'Rielj^jteris callosa (Blume) K. Iwats. 
Steiropteris (C.Chr.) CChr, in Verdoom, Man, Pterid., 544. 

1938, Type: Polypodium deltoideua Swartz = Thelypteris del- 

toidea (Swartz) Proctor, 
Cyclograraraa Tagawa, Acta Phytotax. Geobot,, 7: 53. 1938. Type: 

Thelypteris simulans Ching - Thelypteris auriculata (J.Smith) 

K. Iwats, 
Menisorus Alston. Bol, Sec. Brot,, 30: 20, 1956. Type: Meniscium 

pauciflorua Hook, " Thelypteris pauciflora (Hook.) Reed. 
Dimnrphopteris Tagawa et K, Iwats, ex K, Iwats., Acta Phytotax. 

Geobot., 19: 8, I96I, Type: Dimorphopteris moniliformis Tagawa 

et K, Iwats, - Thelypteris moniliformis (Tagawa et K, Iwats.) 

K. Iwats, 
ParathelypterJB (H.Ito) Ching, Acta Phytotax, Sinica, 8: 300, 

1963. Type: Aspidium glanduligerum Kvinze * Thelypteris glandu- 

ligera (Kunze) Ching, 



1968 Reed, Index Thelypteridls 2^7 

Metathelypteris (Holto) Ching, Acta Riytotax. Sinica, 8: 305o 

1963. Type: Aspidium gracile scans Blume = Thelypteria 

graciie scans (Blume) Ching. 
Macrothelypteris (H.Ito) Ching, Acta Phytotax. Sinica, 8: 3O8. 

1963, Type: Nephrodlum oliHophlebium Baker = Thelypteris 

torresiana (Gaudich,) Alston, 
Pseudophef^opteris Ching, Acta Phytotax. Sinica, 8: 313, 1963. 

Type: Polypodium pyrrhorhachi s Kunze = Thelypteris paludosa 

(Blume) K,Iwat8. 
Glaphyropt eridop si s Ching, Acta Phytotax. Sinica, 8: 320. 1963o 

Type: Polypodium erubescens Wall, ex Hook. = Thelypteris 

erubescens (Wall, ex Hook.) Ching. 
Pseudocyclosorus Ching, Acta Kiytotax. Sinica, 8: 322, I963. 

Type: Aspidium xylodes Kunze = Thelypteris xylodes (Kunze) 

Ching. 
Mesoneuron Ching, Acta Phytotax. Sinica, 8: 325. 1963, Type: 

Aspidium crassifolium Bliune = Thelypteris crassifolia (Blume) 

Ching. 
Anisocaniplum Presl, Epim, Bot., 58. 18A9. Type: Anisocampium 

eumingianum Presl = Thelypteris aristata (Fee) Reed, 



Thelypteris abbiattii Reed, nom. nov, Baslonym: Goniopteris b\ir~ 
kartii Abbiatti, Darwiniana, 13(2-4): 556, f, 6, pi. 4. 19^47 
Argentina. 

Th. abbottiana (Maxon) Ching, Fan Mem. Inst, Biol, Bull,, 10: 250. 
1941. Basionym: Dryopt eris abbottiana Maxon, Joum. Wash. 
Acad. Sci., 14: 89. 1924. Hispaniola, 

Th. abortiva (Blume) Reed, comb. nov. Basionym: Aspidium abor- 
tivum Blume, Enura. PI, Jav,, 154. 1828, Malesia. 

Th. abrupta (Desv,) Proctor, Rhodora, 61: 305. (1959) I960. Basio- 
nym: Polypodium abruptum Desv. . Prodr., 239. 1827. West Indies- 
Brazil. 

Th. acanthocarpa (Copel.) Reed, comb, nov, Basionym: Dryopteris 
acanthocarpa Copel., Philip. Joum. Sci,, Bot, 6: I36, t. 17. 
1911. Borneo. 

Th. achalensis (Hieron. ) Abbiatti, Darv/iniana, 13(2-4): 566. 1964. 
Basionym: Aspidiu m ach-ilense Hieron., Engl. Bot. Jahrb., 22; 
371. I896 (1897); Synonym: Aspidium conterminum var, olii;osorum 
Griseb., Symb., 344, n. 2252, partim. 1879, non Aspidium oligo- 
sorum (Willd.) Kunth. 

Th. acrostichoides (Michx.) Nieuwl., Amer. Midi. Nat., 2: 277. 1912. 
Basionym: Nephrodium acrostichoides l^chx., Fl. Bor. Amer., 2: 
267. 1803. = Polystichum. 

Th, aaiminata (Panz. in Christm. et Panz.) Morton, Amer, Fern Joum., 
48: 139. (1958) 1959. Basionym: Polypodium acuminatum Panz, in 
Christm. et Panz., Nat. Hist., 14: 181, t. XCIX, f. 2, 1783, Syn- 
onjnn: Polypodium sophoroides Thunb, , Trans, Linn. Soc., 2: 341. 
1794; Aspidium oshimense Christ, Bull. Boiss,, II, 1: 1018. 1901; 



258 P H Y T L G I A Vol. 17, no. h 

Dryopt'^rls ogatana Koidz., Bot. l^fr. Tokyo, 39: 10. 1925; Cyclo - 
eorus yamawakii H. Ito, Journ. Jap. Bot., 27: 3^0. 1952. 

Thelyptwris acuminata forma cristata (Tagawa) Reed, coob. nov. basio- 
nym: Dryopteris acuminata forma cristata Tagawa, Acta Phytotax. 
Geobot., 1: 189. 1932. Japan (Honshu, Kyushu). 

Th. acuminata var, kuliangensis (Ching) K.Iwats., Acta Fhytotax. 
Geobot., 21: 40. 1964. Basionym: Cycloaorua a end oat us var. 
kullanp.ensis Ching, Bull. Fan Mem. Inst. Biol., 8: 192. 1932. 
China (Puki en), Japan (Shikoku, Kyushu), Taiwan. 

Th. acuminata VB.r. ogatana (Koidz.) K.Iwats., Acta Phytotax. Geo- 
bot., 21: 40. 1964. Basionj'TTi: Dryopteris ogatana Koidz., Bot. 
Mag. Tokyo, 39: 10. 1925. Japan (Honshu). 

Th. acuminata forma pilosa (H.Itb) Reed, cocb. nov. Basionym: 
Cyclosorus acuminatus forma piloaus H. Ito, Bot. Mag. Tokyo, 
51: 712. 1937. 

Th. adenophora (C.Chr.) Reed, comb. nov. Basionym: Dryopteris ade - 
nophora C.Chr., Ind. Fil., 251. 1905; Nephrodium hirsutua Presl, 
ipim. Bot., 48. 1849, non N^. hirsutum Don, 1825, nee Bory, 1823, 
Luzon, Celebes, 

Th, adenostegia (Copel,) Reed, comb, nov, Basionym: Dryopteris 

adenostegia Copel., Univ. Calif, Publ, Bot., IS: 220. 1942. New 
Guinea. 

Th. adscendens Ching, Bull, Fan Hem. Inst. Biol., 6: 332. 1936, 
China (Kwangtung, Kwangsi). 

Th. adnascens (Ching) Ching sensu Shimizu-H, , Trans, Nat. Hist. 
Soc. Formosa, 28: 238, f. 1. 1938. Basionym: Dryopteris adnas- 
cens Ching sensu Suzuki-3. et 3himizu-H. , Journ, Taihoku Soc. 
Agr. Forest., 2: 187. 1937. Taiwan, This name was published 
in this manner. 

Th, aequatorialis (Copel.) Reed, comb. nov. Basionym: Dryopteris 
aequatorialis Copel., Univ. Calif. Publ. Bot., 19: 298. 1941. 
Ecuador. 

Th. af finis (Presl ex Ett.) Morton, Contrib. U.S. Nat. Herb., 38(2): 
50. 1967. Basionym: Meniscium affine Presl ex Sttingsh., Denk- 
schr. Akad. Wiss. Math. Naturw. (Wien), 23: 94, t. 13, f. 3. 
I864. Sjmonym: Dryopteris dispar Maxon et Morton, Bull, Torr. 
Bot. Club, 65: 364. 1938, Brazil. 

Th. afra (Christ) Reed, comb. nov. Basionym: Dryopteris af ra Christ, 
Bull. Soc. Bot. France, 55 (Mem. 8B): 107. 1908. Synonym: Dryo - 
pteris dewevrei Christ ex Bonap., Not. Pterid., 14: 207. 1924. 
West Trop. Africa (Guinea - Congo). 

Th. afzelii (C.Chr.) Tard. in Humbert, Fl. Madagascar, Fam. 5, 1: 
282. 1958. Basionym: Drvopteris afzelii C.Chr., Ark. f. Bot., 
14(19): 2, t. 1, f. 3. 1916. Madagascar. 

*Th. aizuensis (Suzuki) Reed, comb. nov. Basionym: Cyclosorus 
aizuensis Suzuk_i, Sci. Rept. Fac. Arts & Sci., Eukushima Univ., 
No. 10: 18, t. 1, f. 1-4. 1961. Miocene; Japan (Aizu Basin, 
Fukushiraa Pref,), 

Th, alata (L.) Reed, comb. nov. Basionym: Polypodium alatum L., 
Sp, PI., 2: 1086. 1753. West Indies. 



1968 Reed, Index Thelypteridis 259 

Thelypteris alatella (Christ in K.Schum. et Laut.) Reed, comb, nov. 
Basionym: Nephrodium alatellum Christ in K. Schum. et Laut., 
Fl. Deut. SUdsee, 112. 1901. New Guinea, 

Th, albidipilosa (Bonap.) Reed, comb, nov, Basionym: Dryopteris 
albidipilosa Bonap., Not. Pterid., 15: 9. 1924. East trop, Af- 
rica (Oubangui, Congo). 

Th, albociliata (Copel.) Reed, comb, nov, Basionym: Dryopteris 
albo-ciliata Copel., Journ. Arnold Arb., 10: 177. 1929. Mew 
Guinea, 

Th, albosetosa (Copel,) Reed, comb, nov, Basionym: Dryopteris al- 
bosetosa Copel., Univ, Calif. Publ. Bot., 18: 221. 1942. New 
Guinea . 

Th. alfredii (Rosenst,) Ching, Bull. Fan Mem. Inst. Biol. Bot., 10: 
250. 1941. Basionym: Dryopteris alfredii Rosenst., Fedde Repert, 
22: 10. 1925. Costa Rica. 

•*Th, alpina (Presl in Sternb,) Reed, comb. nov. Basionym: Peco- 
pteris alpina Presl in Sternb,, Flora der Vorwelt, 2: 147, t. 
39, f. 5. IS38; Unger, Gen. et Sp. PI. Foss., 185. 1850. Syno- 
nym: Aspidi un at ember °;ii Ettingsh., Die Farnkr. der Jetztwelt, 
197. I865. Carboniferous: Alpine, Stangalpe, Styria, (Aff, 
Th. concinna et Th, noveboracensis), 

Th, alta (Brause) Reed, comb. nov. Basionym: Dryopteris alt a 
Brause, Sngl. Bot. Jahrb., 56: S6. 1920, New Guinea, 

Th, amboinensis (Willd.) Reed, comb. nov. Basionym: Aspidium am - 
boinense '.«'illd., Sp. PI., 5: 228. 1810. Amboina, 

Th, andina (Morton) Morton, Amer. Fern Journ., 51(1) s 38. 1961. 
Ba3ion3rra: Dryopteris andina Iforton, Journ. Wash. Acad. Sci,, 
28: 528. 1938, Bolivia. 

Th, andreana (Sodiro) Morton, Contrib. U.S. Nat. Herb., 38(2): 
50. 1967. Basionym: Meniscium andreanum Sodiro, Recens., 71. 
lS83j Crypt. Vase. Quit., 392. 1893. Ecuador, Santo Domingo, 

Th. aneitensis (Foum.) Reed, comb. nov. Basionym: Aspidium anei- 
tense Foum., Ann. Sci. Nat., V, 18: 297. 1874. Synonym: Dryo- 
pteris lenormandii CChr,, Ind. Fil., 274. 1905. New Caledonia, 
New Hebrides. 

Th, angulariloba Ching, Bull. Fan Mem. Inst. Biol., 6: 323. 1936, 
Synonym: Thelypteris simozawae Tagawa, Acta Phytotax. Geobot., 
6: 157. 193"^. China (Kwangtung, Hongkong), Japan (Kyushu, Hon- 
shu, Rytikyu). 

Th, angusta (Copel.) Reed, comb. nov. Basionym: Dryopteris anfflista 
Copel., Philip, Journ. Sci., Dot., 9: 3. 1914» New Guinea. 

Tho angustifolia (Willd.) Proctor, Bull. Inst. Jamaica, Sci, Ser. 
No, 5: 57. 1953. Basionym: Meniscium angustifolium V.llld,, Sp. 
PI., 5: 133. 1810. Tropical America, V/est Indies, 

Th, angustifrons (Miq. ) Ching, Bull. Fan Mem. Inst. Biol., Bot, 6: 
318. 1936, Basionym: Aspidium angustifrons Miq., Ann. Mus. 
Lugd, Bot., 3' 178. IB67. Synonyms: Athyrium cygtopteroides 
var, elatius Eaton, Proc, .Amer. Acad., 4 J 110. 1858; Dryopteris 
miyagii H, Ito, _^Bot. Mag, Tokyo, 49: 360. 1935; Dryopterig oki- 
nawaensis H. Ito, Bot. Mag, Tokyo, 49: 36O. 1935; Dryopteris 



260 P H Y T L G I A Vol. 17, no. U 

glanduligera var. hyaloategla H. Ito, Bot. Mag. Tokyo, Lfyi 363. 

1935; Lastrea mlquellana Tagawa, Acta Phytotax. Geobot., 15:1A. 

1953. Japan, Ryukyus, Okinawa lal., Taiwan. 
ThelTpterio angustipes (Gopel.) Reed, comb. nov. Basionya: Dryo - 

pteria angu stipes Ccpel., Philip. Joum. Sci., bot., 7: 60. 1912. 

Saurawak. 
Th. angustipinnata (C.Chr. et Tard. ex Tard.) Reed, comb, nov, Basi- 

onym: Cyoloaorua angustipinnatus C.Chr. et Tard ex Tard., Bull. 

Soc. Bot. France, 87: 369, t. 3. 19U. Tonkin. 
Th. anjenabenais (Tard.) Reed, comb. nov. Basionym: Abacopterig 

anjenabensis Tard., Adansonia, 5(4): U^h, t. 2, f. 5-7. 1965. 

Madagascar. 
Th. anoptera (Kxinze ex Kuhn) Reed, corab. nov. Basionym: Aspidiua 

anopteron Kunze ex Kuhn, Linnaea, 36: 113. I869. Brazil. 
Th. antillana Proctor, Rhodora, 63: 33-34. 1961. West Indies (St. 

Kitts, Dominica). 
Th. aoristisora (Harr.) Reed, comb, nov, Basionym: Polypodiiun ao- 

ristisorua Harr., Joum. Linn. Soc, 16: 30. 1877o Philippines. 
Th, appendiculata (Blume) Reed, comb. nov. Basionjnn: Qymnopiraaiia 

appendiculata Blume, Enum. addend., 1823; Fl. Javae Fil., 92, 

t. 39. 1828. Java, 
Th, aquatilis (Copel.) Reed, comb. nov. Basionym: Dryopteris aoua- 

tilis Copel., Philip. Journ. Sci. Bot., 6: 75. 1911. New Guinea. 
Th, aquatiloides (Copel.) Reed, comb, nov, Basionym: Dryopteris 

aquatiloides Copel., Philip. Joum. Sci. Bot., 7: 59. 1912. Sara- 
wak. 
Th, arborescens (Humb. et Bonpl. ex Willd. in L. ) Morton, Contrib. 

U. S. Nat. Herb., 38(2): 50. 1967. Basionym: Meniscium arbores- 
cens Humb, et Bonpl. ex '.'illd. in L. , Sp. PI., ed. 4, 5: 133. 

1810, Synonym: Thelypteris mollis (Mett.) Tryon, Rhodora, 69: 

7. 1967. Colombia, Venezuela. 
Th, arbusoila (Willd.) K.Iwats., Acta Phytotax, Geobot,, 21(5-6): 

170, 1965. Baionym: Aspidium arbusculum Willd, in L,, Sp, PI,, 

ed, 4, 5: 233. 1810, India-Male sia-Polynesia, Mascarene Isls, 
Th, arcana (Maxon & Morton) Morton, Contrib. U. S. Nat. Herb., 

38(2): 42. 1967. Basionym: Dryopteris arcana Maxon et l-torton. 

Bull, Torr. Bot. Club, 65: 352, t. 11. 1938. Ecuador. 
Th, archboldiae (Copel.) Reed, comb. nov. Basionym: Lastrea arch- 

boldiae Copel., Joum. Arnold Arb., 30: 436. 1949, Fiji Isls. 
Th, archboldii (C.Chr.) Reed, comb, nov, Basionym: Dryopteris ar^h- 

boldii CChr., Brittonia, 2: 297. 1937. New Guinea, Papua. 
Th, arcuata (Poir, in Lam.) Reed, comb. nov. Basionym: Polyp odium 

arcuatum Poir. in Lam., Encycl., 5: 528. 1804. West Indies. 
Th, arfakiana (Bak. in Becc. ) Reed, comb, nov, Basinym: Polypodium 

arfakianum Bak. in Becc, Malesia, 3: 45. 1886. New Guinea. 
The argentina (Hieron,) Abbiatti, Rev. Mus. La Plata, Ser, 2, Bot,, 

9: 19. 1958. Basionym: Aspidium argentinum Hieron. , Engl. Bot, 

Jahrb., 22: 367. 1896 (1897), Argentina - Bolivia. 
Th. arguta (Kaulf,) Moxley, Bull. So. Calif. Acad,, 19: 57. 1920, 

Basionym: Aspidium ar gut urn Kaulf,, Enum. Fil,, 242, 1624. = 

Dryopteris. 



1968 Reed, Index Thelypteridis 261 

Thelypteris arida (D.Don) Morton, Amer. Fern Joum. , 49: 113. 1959. 
Basionym: Aspidium aridum D.Don, Prodr. Fl, Nepal,, 4. 1825. 
Tropics of Asia: Singapore, Mariannas (Saipan), Taivran, Bismarck 
Archipelago, Philippine Isls, (Palawan). 

Th. aristata (Fee) Reed, comb. nov. Basionym: Goniopteris aristata 
Fee, Gen. Fil., 253. 1850-52. Synonym: Aspidium otarium Kxmze ex 
Ifett., Pheg, u. Asp., 34, n. 73. 1858; Anisocampium cumingianum 
Presl, Epim. Bot., 59. 1849, non Aspidium cumingiemum Kunze, 1840, 
Assam, S. India, Ceylon, 

Th, aspera (Presl) K.Iwats., Mem. Coll. Sci., Univ. Kyoto, Ser. B, 
31(3): 192. 1965, Basionym: Goniopteris aspera P resl. Tent, 
Pterid., 183. I836; Polyp odium asperum Presl, Rel, Haenk., 1: 24. 
1825, non L. , 1753. Synonyms: Dryopteris presliana Ching in 
CChr., Ind, Fil. Suppl. Ill: 95. 1934; Abacopteris philippi- 
arum Fee, Congr. Sci, France, 10 Sess,, 178. 1843 J Dryopteris 
gymnopteridif]x>ns Hayata, Icon. PI. Formos., 8: 148. 1919o Tropics 
of 5, Asia, Taiwan. 

Th. aspidioides (Willd.) Tryon, Rhodora, 69: 5. 1967. Basionym: 
Ceterach aspidioides Willd. in L., Sp, PI,, ed. 4, 5: 137. 1810, 
Trop, S. Amer. 

Th. aspidioides var. subhastata (G.Chr, ) Reed, comb, nov. Basionym: 
Dryopteris aspidioides var, subhastata C.Chr., Kgl, Dansk Vid, 
Selsk. Skr., 7: 287, 1907. Costa Rica, Columbia, Venezuela, Peru, 

Th. asplenioides (Swartz) Proctor, Bull, Inst. Jamaica, Sci, Ser., 
No. 5: 57. 1953. Basionym: Polyp odiiun asplenioides Swartz, in 
Schrad. Joum. Bot., 1800(2): 26. 1801. West Indies. 

Tho assamica (Bedd.) Reed, comb, nov, Basionym: Nephrodiua multi- 
lineatum var. assamicxu n Bedd., Joum. Bot,, 31: 228, 1893. Assam, 

Th. assurgens (Maxon) Tryon, Rhodora, 69: 5o 1967. Basionym: Dryo- 
pteris assurgens llaxon, Joum, Wash. Acad. Sci,, 34: 24. 1944o 
Pera. 

Th. asterothrlx (Fee) Proctor, Bull. Inst, Jamaica, Sci, Ser,, No, 
5: 57. 1953. Basionym: Goniopteris asterothrix Fee, Gen. Fil,, 
253. 1850-1352, Synonym: Dryopteris malacothrix Maxon, Proc. 
Biol. Soc. V/ash., 43: 87. 1930, Trop. Amer. 

Th, asterothrix var. bibrachiata (Jenm. ) Proctor, Bull. Inst. Ja- 
maica, Sci. Ser., No, 5: 58. 1953. Basionym: Nephrodium bibra- 
chiatum Jenm., Gard. Chron,, III, 15: 230. 1894. West Indies, 

Th. asyraetrica (Fee) Reed, comb. nov. Basionym: Goniopteris asyaetri- 
ca Fee, Gen. Fil., 253. 1850-1852, Philippine Isls. 

Th, atasripii (Rosenst.) Reed, comb. nov. Basionym: Dryopteris 
atasripii Rosenst,, Med. Rijks Herb,, No. 31: 6. 1917. New 
Guinea, 

Th. arthrothrix (Hook.) Reed, comb, nov, Basionjrm: Polypodium 

arthrothrix Hook., Sp. Fil,, 5: 14. I863. Madagascar, Mascarenes, 

Th, atrospinosa (C.Chr. ex Kjellb.et C.Chr.) Reed, comb. nov. 
Basionym: Dryopteris atrospinosa C.Chr. ex Kjellb. et C.Chr., 
Engl. Bot, Jahrb., 66: 43. 1933, Celebes. 

Th. atrovirens (C.Chr. in Christ) Reed, comb. nov. Basionym: Dryo- 
pteris atrovirens C.Chr, in Christ, Bull, Boiss., II, 7: 263, 



262 P H Y T L G I A Vol. 17, no. k 

1907; Vid. Selsk. Skrift., VII, U: 316, f. 39. 1907. Synonym: 
Laatrea coataricensj 3 Copel., Gen. Fil., I38. 19^7. Central Amer. 

Thelypteris attenuata (Kuntze) Morton, Contrib. U.3. Nat. Herb., 
38(2): 35. 1967. Ba3ionym: Dryopteris attenuata Kuntze. Rev. 
Gen. PI., 2: 812, 1891; Aapidium attenuatua Kunze ex Mett., Pheg. 
u. Asp., 96, n. 233. 1858, non Swartz, 1801; Laatrea attenuata 
J.Smith, Journ. Eot., 3: 412. I84I (nonen), non brack., 1854; 
Nephrodium attenuatum Fee. Gen. Fil., 305. 1850-1852 (noraen). 
Synonym: Dryopteri s atenobaaia C.Cbr., Ind. Fil., 294. 1905. 
Philippine Isls. (Seunar), Celebes, 

Th, augescens (Link) Munz et Johnston, .'imer. Fern Joum., 12; 75. 
1922, Basionym: Aapidiu m augescens Link. Fil. 3p., 103. 1841. 
Mexico-Costa Rica, Cuba, rtest Indies, 

Th . augeacens (var.) lindheimeri (A.Br, ex C.Chr.) R.P. St. John ex 
Small, Ferns Southeastern States, 241. 1938. Baaionym: Aapidium 
lindheimeri A.Br, ex C.Chr., Danske Vid, Selsk. Skr., VII, 10: 
182. 1913 (pro ayn.), =■ Thelypteris X lindheimeri cherry. 

Th, aureo-viridis (Rosenst,) Reed, comb. nov. Basionym: Dryopteris 
aureo-viridis Roserst., Fedde Repert., 13: 216. 1914. Taiwan. 

Tho auriculata ( J. Smith) K.Iwats., Acta Phytotax. Geobot., 19: 11. 
1961. Basionym: Phegopteris auriculata J.Smith, Hist. Fil., 
233» 1875; Polypodium auriculatum Wall, ex Hook., Sp. Fil., 
IV: 237. 1862, non L., 1753, nee Raddi, 1319, nee Presl, 1822, 
Synonyms: Polypodium appendiculatmn var. aquanaestipes Clarke, 
Trans. Unn. Soc, 11. Bot,, 1: 534, t. 79, f. 2, 1880; Dryo- 
pteris himalaye nsis C.Chr,, Ind. Fil. Suppl, III: 88, 1934; 
Thelypteris simula ns Ching, Bull. Fan Mem. Inst. Biol,, Bot, 
6: 280. 1936; Thelypteris subvillosa Cning, Bull, Fan Kern. Inst. 
Biol,, Bot. 6: 270. 1936." Taiwan, Nepal, China (Yunnan), 
Sikkim-Himalaya, Java, 

Th, auricolifera (v.A.v.R. ) Ching, Bull, Fan Mem. Inst. Bid., 

Bot. 10: 250. 1941. Basionym: Dryopteris auriculifera v.A.v.R., 
Bull. Buit., Ill, 5: 197. 1922. Lingga Isl., Malesia. 

Th, aurita (Hook.) Ching, Bull. Fan Mem. Inst. Biol., Bot. 6: 
266. 1936, Basionym: Gymnogramma aurita Hook., Icon, PI., t, 
974. 1854, Sikkim-Tonkin, 

Th, austera (Brause) Reed, comb, nov, Basionym: Dryopteris aus- 
tera Brause, Engl, Bot. Jahrb., 56: 108. 1920, New Guinea, 

Th, austro-philippina (Copel.) Reed, comb. nov. Basionym: Dryo- 
pteris austro-philippina Copel.. Philip. Journ, Sci., 40: 300. 
1929, Philippine Isls, (J4indanao). 

Th. badia (v.A,v,R. ) Ching, Bull. Fan Mem. Inst. Biol., Bot. 10: 
250. 1941, Basionym: Dryopteris badia v.A.v.R.. Bull. Jard, 
Bot. Buit., II, nr. XVI: 9. 1914. Synonym: Dryopteris linearis 
Copel., Philip, Joum. Sci,, 12C: 56. 1917. Sumatra, Borneo. 

Th, bakeri (Harr. ) Reed, comb. nov. Basionym: Nephrodium bakeri 
Harr., Joum. Linn. Soc., I6: 29. 1377. Philippine Isls. (Panay), 

Th, balansae (Ching) Reed, comb, nov, Basionym: Cycloaorus bal an- 
sae Ching, Bull. Fan Mem. Inst. Biol,, Bot. 8: 200. 1938. Tonkin. 



1968 Reed, Index Thelypteridis 263 

Thelypteris balbisii (Spreng.) Ching, Bullo Fan Mem. Inst. Biol. 
Hot., 10: 250. 1941. Basionym: Polypodiu m balbi sil Spreng,, 
Nova Acta Caes, Leop. Carol., 10: 228. 1B21. Synonyms: Aspidi xim 
sprengelii Kaulf., Flora, 1823(1): 365. 1823; Nephrodiuro sher- ' 
rlngii Jenm. , Joum. Bet., 1?: 261. 1879. West Indies (Cuba, 
Jamaica, Haiti, Puerto Rico)^ Guatemala, l^Caragua, C.R,, Venezuela, 

Th. banaensis Tard. et C.Chr., Not. Syst. (Parisj, ?: 66. 1936. 
Annam. 

Th, bangii (CChr.) Tryon, Rhodora, 69: 5. 1967. Basionym: Dryo- 
pteris bangii C.Chr., Danske Vid, Selsk. Shrift., VII, U: 333, 
f, 52 II. 1907; I.e., VII, 10: 190. 1913. S, Amer. (Bolivia, 
S. Brazil). 

Tho baramensis (C.Chr. ex C.Chr. et Holtt.) Ching, Bull, Fan Mem. 
Inst. Biol. Bot., 6: 287. 1936. Basionym: Dryopteris baramensis 
C.Chr. ex C.Chr. et Holtt., Gardens Bull. Straits Settlements, 
7: 2A6. 1934. 

Th, bartlettii (Copel,) Reed, comb, nov, Basionym: Dryopteris bart- 
lettii Copel., Univ. Calif. Publ. Bot., Ik: 374, t. 58. 1929. 
Sumatra, 

Th, batacorum (Rosenst,) Reed, comb, nov, Basionjon: Dryopteris 
batacorum Rosenst., Fedde Repert., 13: 217. 1914. Sumatra. 

Th. batjanensis (Rosenst.) Reed, comb. nov. Basionym: Dryopteri s 
bat.1a.nen sis Rosenst., Med. Rijks Herb., No. 31 J 5. 1917. Batjan 
Isl. 

Th. beccariana (Ces.) Reed, comb. nov. Basionjna: Nephrodium bec- 
carian um Ces., Atti Ac. Napoli, 7(8): 23. 1876. Borneo, 

Th. beddcmei (Bak. in Hook, et Bak.) Ching, Bull. Fan Memc Insto 
Biol, Bot., 6: 308. 1936. Basionym: Nephrodium beddomei Bak. 
in Hook, et Bak., Sjm. Fil,, 267. 1867. S, India, Ceylon, 
Malaysia, Taiv;an, Philippine Isls,, Java, 

Th, belensis (Copel.) Reed, comb, nov, Basionym: Dryopteris 
belensis Copel., Univ. Calif. Publ, Bot,, 18: 220, 1942. 
New Guinea. 

Th, benguetensis (Copel.) Reed, comb, nov, Basionjrm: Gyclosorus 
benguetenais Copel,, Philip, Joum, Sci,, 81: 28. 1952. Philip- 
pine Isls, (Luzon), 

Tho bergiana (Schlechtend, ) Ching, Bull, Fein Hem, Inst. Biol. Bot,, 
10: 251. 1941. Basionym: Polypodium bergianum vSchlechtend,, 
Adumbr, , 20, t. 9. 1825. Synonyms: Aspidium maranA"uense Hieron,, 
Engl, Pfl.-fajn. Ost. Afr., C: 85. 1895; Dryopteris palmil C.Chr,. 
Ark, f, Bot., 14: 1, t. 2, f. 6. 1916; Nephrodi um sewellii Bak., 
Joum. Linn. Soc, 15: 418, 1876; Nephrodium anateinophlebium 
Bak,, Joum. Linn. Soc, 16: 202. 1877. S. Africa, Madagascar. 

Th, berroi (C.Chr.) Reed, comb, nov, Basionym: Dryopteris berroi 
C.Chr,, Kgl. Danske Vid. Selsk. Skr., VII, 10(2): 185, f. 24. 
1912, Synonym: Cyclosorus berroi (C.Chr.) Abbiatti, Darvdniana, 
13(2-4): 567. 1964. Uruguay, Paraguay, Argentina. 

Th, biaurita (Bedd.) Reed, comb. nov. Basionym: Nephrodium bi- 
auritum Bedd., Handb. Suppl., 68. 1892. Assam. 



26U P H y T L G I A Vol. 17, no. h 

Thelypteris bi color (Bonap.) R«od, comb. nov. Basionym: Dryopteric 

bicolor Bonap., Notes Pterid., U: 20i*. IV^-ii*. Gabon, 
Th, bifonnata (Rosenst.) Tryon, Rhodora, 69: 5. 1967. Basionym: 

Dryopterlfl bifonnata Rosenst., Fedde Repert., 7: 300. 19C'9. Peru. 
Th. biolleyi (Christ in Pittier) Procter, Bull. Inst. Jamaica, Sci. 

Ser., No. 5: 58. 1953. Basionym: Aspidium biolleyi Christ in 

Pittier, Prim. Fl. Costaric, 3: 31. 1901. Costa Rica-Panaiaa, 

Jamaica. 
Th, bipinnata (Cope].) Reed, comb, nov. Basionym: Dryopteris bi- 

pinnata Copel., Philip. Joum. Sci. Bot., 9: 2. 191/*. New Guinea. 
Th. blanda (F6e) Reed, comb. nov. Basionym: Phegopteris blanda 

Fe'e, 86 Mem., 91. 1857. Mexico- Costa Rica. 
Th, blastophora (Alston) Reed, comb. nov. Basionyir: Cyclocorus 

blastophorus Alston, Bol. 3oc. Broter., Ser. II, 30: 12. 1956. 

S. Nigeria, 
Th. boholensis (Copel.) Reed, comb. nov. Basionym: Cyclosorus bo- 

holensig Copel., Philip. Joum. Sci., 81: 31, t. 23. 1952. Philip- 
pine Isls. (Bohol). 
Th, boliviensis (l-Iorton) Morton, V^ier. Fern Joum., 51: 38. 1961, 

Basionym: Diyopteris boliviensis Morton. Joum. V/ash. Acad. Sci., 

28: 527. 1938. Bolivia. 
Th, bonapartii (Rosenst.) Alston, Joum, Wash. Acad. Sci., hS{7)i 

233. 1958, Basionym: Dryopteris bonapartii Rosenst., Fedde Re- 
pert., 7: 303. 1909. Ecuador, Colombia. 
Th, boninensis (Kodama ex Koidz.) K.Ivrats., Acta Phytotax. Geobot,, 

21: /vl. 1964. Basionym: Dryopteris boninensis Kodama ex Koidz., 

Bot. Mag. Tokyo, 38: 109. 192A. Bonin Isls, 
Th, boottii (Tuckerm. ) Nieuwl., Amer. Midi. Nat., 1: 226. 1910. 

Basionym: Aspidium boottii Tuckenn., Hovey's Mag. Hort., 9: 145. 

1843. = X Dryopteris boottii. 
Th, boqueronensis (Hieron.) Tryon, Rhodora, 69^ 5. 1967o Basionym: 

Dryopteris boqueronensis Hieron., Hedvdgia, 46: 329. 1907, 

Colombia. 
Th, bordenii (Christ) Reed, comb, nov, Basionym: Dryopteris borde - 

nii Christ, Philip. Joum. Sci, Bet., 2: 204. 1917. Philippines. 
Th, bomeensis (Hook,) Reed, comb, nov, Basionym: Polypodivun bor- 

neense Hook., Sp. Fil., 5: 11. I863. Synonym: Dryopteris labu- 

anensis C.Chr., Ind. Fil., 273. 1905. Borneo. 
Th, boydiae (Eaton) K.Iwats., Mem. Coll. Sci. Univ. Kyoto, Ser. B, 

31(1): 36. 1964. Basionym: Aspidium boydiae Eaton, Bull. Torr, 

Bot. Club, 6: 361. 1879. Hawaii. 
Th, brachyodus (Kunze) Ching, Bull. Fan Mem. Inst. Biol. Bot., 6: 

286 (err. "brachyodon" ). 1936. Basionym: Polypodium brachyodus 

Kunze, Linnaea, 9: 48. 1834. West Indies, Costa Rica - Peru, 

Galapagos, Malesia, 
Th, brachypoda (Bak. in Im Thuna) Morton, Fieldiana, Bot., 23: 10- 

11. 1951. Basion3TB: Nephrodium brachypodum Bak, in Im Thurn, 

Timehri, 5: 213, 1886; Bak., Trans. Linn. Soc. II. Bot., 2: 290, 

1887, Venezuela, Mt. Roraima. 



1968 Reed, Index Thelypteridis 265 

Thelypteris brackenridgei (Mett. ) Reed, comb, nov, Basionym: 

Aspidiiim brackenridgei Mett.. Ann, Sci, Nat,, IV, 15: 75. 1861. 
Polynesia, Bismarck Archipelago - Tahiti, 

Th. bradei (Christ) Reed, comb. noVo Basionym: Dryopteris bradei 
Christ, Bull, Soc. Hot. tteneve, H, 1: 225. 1909. Costa Rica, 

Th, brasiliensis (C.Chr.) Morten, Amer. Fern Joum,, 51: 38. 1961. 
Basionym: Dryopteris decussata var. brasiliensis C.Chr., Danske 
Vid, Selsk, Skr,, VII, 10:TSl. 1913. Brazil, 

The brassii (C.Chr.) Reed, comb, nov, Basionym: Dryopteris bras si i 
C.Chr., Brittonia. 2: 295. 1937, Nev/ Guinea (Papua), 

Tho brausei (Hieron.) Alston, Journ. Wash. Acad. Sci., 48(7): 233. 
1953. Basionym: Dryopteris brausei Hieron,, Hedvdgia, 46: 337> 
t, 6, f. 11, 1907, Ecuador, Colombia. 

Th. brittonae (Slosson ex tiaxon in Britt.) Reed, comb, nov, Basio- 
nym: Dryopteris britt onae Slosson ex Maxon in Britt., Pterido 
Porto Rico, 475, 1926, Puerto Rico, Hispaniola, 

*Th, brongniartii (Ett.) Reed, comb. nov. Basionym: Aspidium 
bronffiiartii Ett., Die Famkr, der Jetztwelt, 200. I865. Syno- 
nyms: Pecopteris unita Brcngn., Hist. Veg, Foss., 1: 342, t, 116, 
f . 1-5. 1828; Cyatheites unitus Geinitz, Verstein. der Stein- 
kohl, in Sachsen, 25, t. 29, f. 4-5. 1855. Upper Carboniferous: 
Germany (Saarbrllck, Gaislauter, Wettenau, Oberhohndorf ), France 
(Alias, St. Etienne) Aff, Th. unita, 

Th, brooksii (Copel.) Reed, comb, nov. Basionym: Dryopteris brook- 
sii Copel., Philip, Journ. Sci. Bot,, 3: 345. 1909. Borneo. 

Th . brunnea (Wall, ex C.Chr,) Ching, Bull. Fan Mem, Inst, Biol,, 
6: 269. 1936. Basionym: Dryopteris brunnea Wall, ex C.Chr,, 
Ind. Fil., 255. 1905; Po lino odium brunneum Wall., nomen nud.; 
Phegopteris brunnea J.Smith, Hist. Fil,, 233 (brunea), 1875 
(nomen), = Th. paludosa. 

Th. brunnea var. glabrata (Clarke) Ching, Bull, Fan Mem, Inst. 

Biol., 6: 271. 1936. Basionym: Polypodium distans var, glabrata 
Clarke, Thans. Linn. Soc, II. Bot., 1: 544. 1880. 

Th o brunnea var. hirtirachis (C.Chr.) Ching, Bull. Fan Mem„ Inst, 
Biol,, 6: 27I0 1936. Basionym: Dryopteris hirtirachis C.Chr., 
Ind. Fil.jSuppl. II: 15. (1916) 1917. = Th. paludosa, 

Th , brunnea var, pallida Ching, Bull. Fan Mem. Inst, Biol., Bot. 
11: 62. 1941. 

Th, bnonnescens (C.Chr. ex Kjellb. et C.Chr,) Reed, comb. nov. 
Basionym: Dryopteris brunnescens C.Chr. ex Kjellb. et C.Chr., 
Engl, Bot. Jahrb,, 66: 44. 1933. Celebes, 

Th. bukoensis (Tagawa) Ching, Bull, Fan Mem. Inst. Biol., Bot, 6: 
272. 1936. Basionym: Dryopteris bukoensis T agawa. Acta Phyto- 
tax. Geobot., 1: 89. 1932. Japan (Honshu). 

Th. bunnemeyeri Reed, nom. nov, Basionym: Leptogramma celebica 
Ching, Sinensia, 7: 99, t. 5. 1936. Celebes. 

Th. burkartii Abbiatti, Darvd.niana, 13(2-4): 550-551, f. 4, pi, 2. 
1964. Argentina. 

Th. burmanica (Ching) Reed, comb. nov. Basionym: Cyclosorus 
burmanicug Ching, Bull. Fan Mem. Inst. Biol,, Bot. 8: 205, 
1938. Burma. 



266 PHYTOLOGIA Vol. 17, no. h 

Thelypteris cabrerae (Weatherby)Abbiatti, R«v, Mxis. La Plata, Ser, 

II, Bot. 9: 19. 1958. Basionym: Dryopteris cabrerae V/'eatherby, 

Bol. 3oc. Urgent, fiot., 3: 31, f. 1-5. 19i*9. Argentiria. 
Th, calcarata (Blune) Ching, Bull. Fan Kern. Inst. Biol. Bot., 6: 

288. 1936, Basionym: Aspidium calcaratum Blunie, Enum. PI. Jav., 

159, 1828, Oriental tropics: 3. China - India - Java - Malesia - 

Samoa. 
Th, callensii (Alston) Reed, comb, nov, Basionym: Cyclosorus cal - 

lensii Alston, Bol. Sec. Broter., Ser. II, 30: 13. 1956. Congo. 
Th, callosa (Blurae) K.Ivrats., Mem. Coll. Sci., Univ. Kyoto, Ser.B, 

31(1): 3h. 1964, Basionym: Aspidium callosum Blume. Snum, PI. 

Jav., 152. 1828, Java, Sumatra. 
Th, calva (Copel.) Reed, comb, nov, Basionjrm: Dryopteris calva 

Copel., Leaflets Philip. Bot., 3: 808. 1910. Mindanao. 
Th. CAlvata Ching, Bull, Fan Mem. Irst, Biol,, 3er II, 1: 313. 

1949. China (Kwangtung). 
Th, calvescens (Ching) Reed, comb, nov, Basionym: Cyclosorua cal- 

vescens Ching, Bull. Fan Mem. Inst, Biol., Bot. 8: 225. 1938. 

China (Kweichow), Tonkin. 
Th, cana (J.Smith) Ching, Bull. Fan Mem. Inst. Biol., Bot. 6: 291. 

1936. Basionym: Lastrea cana J. Smith, Cat. Cult. Ferns, 57. 

1857. Sikkim, 
Th, canadasii (Sod.) Alston, Joum. Wash. Acad. Sci., 48(7): 234. 

1958, Basionym: Nephrodium canadasii Sodiro, Rec,, 48. 1883j 

Crypt. Vase. Quit., 236. 1893. Ecuador. 
Th. canescens (Blurae) K.Iwats,, Mem. Coll. Sci., Univ, Kyoto, 

Ser. B, 31(1)5 34. 1964. Basionym: Polypodimr canescens Blume. 

Snum. PI. Jav., 133. 1328, Java, Celebes, Philippine Isls.,FiJi. 
Th. cinlaonensis (Copel.) Reed, comb, nov, Basionym: Dryopteris 
canlaonensis Copel., Philip. Journ. Sci., 40: 300. 1929. Philip- 
pine Isls. (Negros). 
Th. caribaea (Jenm.) Morton, Amer, Fern Journ., 53: 65. 1963. Basi- 
onym: Mephro d ium c aribeum J enm. , Joum, Bot„, Brit, & For., 24: 

270. 1886, ^j'imaica. 
Th, castanea (Tagav;a) Ching, Bull. Fan Memo Inst, Biol., 6: 315. 

1936, Basionym: Dryopteris castanea Tagawa, Acta Phytotax. 3eo- 

bot., 4: 132. 1935. M. Taiwan, 
Th, cataractorum (Wagner & Grether) Reed, comb, nov, Basionym: 

Cyclosorus cataractonua Wagner Sz Grether, Univ. Calif. Publ, 

Hot., 23: 50, t. 16. 1948, Ins. Admiralty. 
Th, caucaensis (Hieron.) Alston, Joum. Wasli. Acad, Sci,, 48(7): 

233. 1958, Basionym; Nephrodium caucaense Hieron. . Engl, Bot, 

Jahrb., 34: 444. 1904, Costa Rica - Colombia - Bolivia. 
Th, caudata (Ching) Reed, comb, nov, Basionym: Leptogramma. cau- 

data Ching, Sinensia, 7: 98, 6. 4. 1936, China (Fukien), Taiwan. 
Th, caudiculata (Sieber in Kunze) Reed, comb, nov, Basionym: 

Aspidium caudiculatun Sieber in Kunze, Linnaea, 24: 28. I85O. 

Comores, Mauritius. 
Th. caudipinna Ching, Bull. Fan Mem. Inst. Biol., 6: 288. 1936. 

China (Hainan). 



1968 Reed, Index Thelypteridis 26? 

Thelypteris cavatensis (Copel.) Reed, comb, nov, Basionyra: Las- 
trea cavltensla Copel., Philip. Journ. Sci., 81: 26. 1952. 
Luzon . 

Th. celebica (Baker) Reed, comb. nov. Basionym: Aero sti chum cele- 
blcura Baker, KewB'oll., 19C1: U5. 1901. Celebes. 

Th. cesatiana (C.Chr. ) Reed, corob. nov, Basionjir: Dryopteris cesa- 
tiana C.Chr., Ind. Fil., 257. 1905. Synonym: Meniscium beccari- 
anum Ces,, Rend. Acad. Napoli, 16: 27, 3'-'. 1877, non Nephrodiuir 
beccarianum Ces., 1876. Nev; Guinea, Fiji, 

Th. chaerophylloides (Poir. in Lam.) Proctor, Bull. Inst. Jamaica, 
Sci. Ser., No. 5: 58. 1953. Basionym: Polypodium chaerophyl- 
loides Poir, in Lam., Encycl., 5: 5A2. 180/;. west Indies. 

Th, chamaeotaria (Christ) Reed, comb. nov. Basionym: Dryopteris 
chamaeotaria Christ, Philip. Joum. Sci,, Bot. 2: 203. 1907. 
Malesia - Polynesia. 

Th, chaseana Schelpe, Journ. S. Afr. Bot., 31(4): 263, f. 1 e-f. 
Oct. 1965. Africa (Southvest, Rhodesia, Malawi, Zambia, Tan- 
zania, Angola). 

Th. cheilanthoides (Kunze) Proctor, Bull. Inst. Jamaica, Sci. Ser,, 
No. 5: 58, 1953. Basionym: Aspidium cheilanthoides Kunze, Lin- 
naea, 22: 578. 1849. Brazil, Trop. A-ner. 

Th. cheilanthoides var. resinofoetida (Hook.) Proctor, Bull. Inst. 
Jamaica, Sci, Ser,, No. 5: 58. 1953. Basionj/m: Menhrodium re- 
sinofoetidum Hook., Sp. Fil., 4: 105. 1862. Jamaica. 

Th, chinensis Ching, Bull. Fan Mem, Inst. Biol., 6: 311. 1936. 
China (Anwhei), 

Th, chlanipdophora (Rosenst.) Ching, Bull, Fan Mem, Inst. Biol., 
6: 287. 1936, Basionym: Dryopteris chlamydophora Rosenst., 
Med. Rijks Herb. Leiden, 31'. 5. 1917. Malaya, Borneo, prob, 
Sumatra. 

Th, christensenii (Christ) Reed, comb, nov, Basionym: Dryopteris 
christensenii Christ, Bull. Boiss., II, 7: 263. 1907; C.Chr., 
Vid. Selsk. Skr., VII, 4: 322, f. 46. 1907c Costa Rica -Panama. 

Th. christii (C.Chr.) Reed, con-.b, nov. Basionjmi: Dryopteris 

Christ il C.Chr., Ind. Fil., 257. 1905. Synonym: Meniscium opa- 
cura Bak., Journ. Bot., 1877: 166. 1877. Ecuador. 

Th. chrysoloba (Kaulf, ex Link) Ching, Bull, Fan Mem, Inst. Biol., 
Bot. 10: 251. 1941. Basionym: Aspidium chrysolobum Kaulf. ex 
Link, Hort. Berol., 2: 117. 1833. Brazil, Colombia. 

Th. chrysodioides (Fee) Morton, Contrib. U.S. Nat. Herb., 38(2): 
51. 1967. Basionjnn: Meniscium chrysodioides Fee, Gen. Fil., 
225. 1852. S. Amer. 

Th, chrysodioides var. goyazensis (Maxon & Morton) Morton, Contrib. 
U. S. Nat. Herb., 38(2): 51. I967. Basionym: Dryopteris chryso- 
dioides var. goyazensis Maxon & Itorton, Bull. Torr. Bot. Club, 
65: 374. 1938. Brazil. 

Th. chunii Ching, Bull. Fan Mem. Inst. Biol., Bot. 6: 284. 1936. 
China (Kvrangtung, Kweichov). 

Th. ciliata (V/all. ex Benth.) Ching, Bull, Fan Mem, Inst. Biol,, 
Bot. 6: 289. 1936. Basionyir: Aspidium ciliatum Wall, ex Benth., 



268 P H Y T L G I A Vol. 17, no. h 

Fl. Hongkong, ^55. 1661. Synonym: QryopterlB paeudocalcarata 
C.Chr., Ind. Fil. Suppl. Ill: 95. 1V3^. Hongkong, Nej al li to 
Malaya. 
*Thelypteris claibomiana (Berry) Reed, comb. nov. Baaionym: ^nio- 
pterls claibomiana Berry, Bull. Torr. Bot. Club, UU: 331, t. 22. 
1917. Eocene: Louisiana, Kissieaippi. 

Th. clarkei (Bedd.) Reed, comb, nov, Basionym: Pleocnemia clarkei 
Bedd., Ferns Brit. India, Suppl., 15, t. 368. 1?76. oynon./u.: lle - 
phrodium artinexiim Clarke, Trans. Linn. 3oc., II. bet., 1: 53*^. 
1880. Sikkim. 

Th . clintoniana (D.C.Eaton) House, New York State Mus. Bull., 233- 
23A : 69 . 1922 . Basionym: Aspidium cristatuip. var. cllntcnianum 
D.C.Eaton in A, Gray, Manual, 5th ed. : 665. 1867. - Dryopteris. 

Th, clemensiae (Copel.) Reed, comb. nov. Basionym: Dryopteris cle- 
mensiae Copel., Philip. Joum. Sci., 46: 213. 1931. Luzon. 

Th, clypeolutana (Desv.) Proctor, Rhodora, 6I: 3O6. (1959) I960. 
Basionym: Nephrodium clypeolutanum Desv. . Mem, Soc, Linn, Paris, 
6: 258. 1827. Endemic to Lesser Antilles. 

Th.coalescens (Baker) Reed, comb, nov, BasionjTti: Polypodiati coa— 
lescens Baker, Joum. Bot,, 1877: I64. 1877. Ecuador. 

Th, coarctata (Kunze) Tryon, Rhodora, 69: 5, 1967, Basionym: Aspi- 
dium coarctatum Kunze. Bot. Zeit,, 18^5: 287. 1845. Caracas, Co- 
lombia, 

Th, Columbiana (C.Chr.) Morton, Leaflets of V.'estem Bot., 8(8): 194. 
1957. Basionym: Dryopteris columbiana C,Chr,, 7id, Selsk, Skr,, 
VII, 4: 279, f. 8. 1907. Colombia, Panama, Galapagos Isl, 

Th, comosa (Morton) Morton, Aner. Fern Joum., 51 J 38. 1961, Basi- 
onym: Dryopteris como sa Morton, Joum. Wash. Acad. Sci., 28: 
528, 1938. Peru, 

Th, compacta (Copel.) Reed, comb, nov, Basionym: Diropteris com - 
pacta Copel,, Philip, Joum. Sci., Bot, 6: 137, f. 18, 1911, 
Borneo, 

Th. concinna (Willd.) Ching, Bull. Fan Mem. Inst. Biol,, Bot. 10: 
251, 1941. (Proctor, Bull, Inst. Jamaica, Sci. Ser., No. 5: 58. 
1953)0 Basionym: Polypodium concinnum V/illd., Sp. PI., 5: 201, 
1810. Trop, Amer., Jamaica, Cuba, Mexico - Costa Rica - Ecuador, 

Th. concinna var. elongata (Eoum, ) Reed, comb, nov, Basionym: 
Phegopteris elongata Foum., PI. Mex. , 1: 87. 1872, Mexico. 

Th, conferta (Brause) Reed, comb. nov. Basionym: Dryopteris con- 
ferta Brause, Engl, Bot. Jahrb., 49: 22, f . IF, 1912, New Guinea. 

Th, confluens (Thunb.) Morton, Contrib. U.S. Nat, Herb,, 38(2): 
71. 1967. Basionym: Pteris confluens Thunb., Prodr. PI. Cap., 
1800, S, Africa, Cape of Good Hope, 

Th. confusa (Copel.) Reed, comb, nov, Basionym: Dryopteris confusa 
Copel,, Philip, Joum, Sci., 6C: I46, 1911. Luzon, 

Th. connexa (Kuhn ex Baker) Reed, comb, nov, Basionym: Nephr odium 
connexum Kuhn ex Baker, in I'fartius Fl. Bras., 1(2): 489. 1870. 
S3monym: Dryopteris martini C.Chr., Indc Fil,, 276, 1905o Cayenne. 

Th, consanguinea (F^e) Proctor, Rhodora, 61: 306, (1959) I960. Basi- 
onym: Aspidium consanguineum F^e, lie Mem. Foug., 76, t. 20, f.3o 
1866, Guadeloupe, V.'est Indies, 



1968 Reed, Index Thelypteridis 269 

Thelypteris consanguinea var. aequalis (C.Chr. ) Reed, comb, nov, 

Basionym: Dryopteris consangiinea var, aequalis CoChr,, ^olths. 

Misc. Coll., 52: 380. 1909. Jamaica, Grenada. 
Th. consimilis (Fee ex Baker) Proctor. Rhodora, 68:^ 468, 1966. Basi- 
onym: (bnnnograinna gracilis " G. consimilis " Fee ex Baker, in 

Hook, & Baker, Syn. Fil., 377. 1868 (invalid), Jamaica, Guade- 
loupe . 
Th. consobrina (Maxon & Morton) Tryon, Rhodora, 69: 5. 1967. Basio- 

nyn: Dnropteris consobrina Maxon & Morton, Bullo Torr. Bot, dub, 

65: 356. 1938. ' Peru. 
Th. conteraina (Willd, in L. ) Re«d, comb, nov, Basionym: Aspidium 

conteminum Willd. in L,, Sp. PI., ed. 4, 5: 249. 1810. TropoAmer. 
Th. conterminoides (C.Chr.) Reed, comb, nov. Basionym: Dryopteris 

contenainoides C.Chr., Ind. Fil., 258. 1905. Synonym: Nephro- 

dium simulans Baker. Joum. Bot., 1890: 106. 1890. New Guinea. 
Th, contigua (Rosenst.) Reed, comb. nov. Basionym: Dryopteris con- 

tigua Rosenst., Med. Ri^ks. Herb. Leiden, No. 31: 8. 1917. Bor^ 

neo, Malaya, 
Th . contingens Ching, in herb.; pro syn., in Ching, Acta Hij't.otax. 

Sinica, 8: 310. 1963. ( Macrothelypteris contingens Ching, I963). 
Th. cooleyl Proctor, Rhodora, 68: 468-469. I966. St. Vincent. 
Th. copelandii Reed, nom. nov, Basionym: Cyclosorus dimojTphus Copel., 

Philip. Joum, Sci., 83: 99, t. 5. 1954o Luzon, 
Th. cordata (Fee) Proctor, Bull. Inst. Jamaica, Sci. Ser,, No. 5: 

59. 1953. Basionym: Phegopteris cordata Fee, Gen. Fil., 244. 

I85O-I852. Synonym: Polypodium cubanum Baker in Hook, i Bak., 

J^. Fil., 304. I867. West Indies (Cuba, Puerto Rico, Jamaica), 

Mexico. 
Th. coriacea (Brause) Ching, Bull. Fan Mem^ Inst. Biol., Bot. 10: 

251. 1941. Basionym: Dryopteris coriacea Brause, Engl. Bot. 

Jahrb., 56: 83. 1920. New Guinea, 
Th, cornuta (Maxon) Ching, Bull. Fan Mem. Inst. Biol., Bot. 10: 

251. 1941. Basionym: Diropteris cornuta Maxon, Joum, Wash. 

Acad. Sci,, 19: 245, fig. 1924. Tahiti. 
Th, costata (Brack.) Reed, comb. nov. Basionym: Goniopteris cos- 

tata B rack.. Expl. Exp., 16: 28. 1854. Society Isls. (FijiTT" 
Th, costulisora (Copel.) Reed, comb, nov, Basionym: Lastrea costu- 

lisora Copel,, Gen. Filo, 138. 1947. Synonym: Dryopteris basi- 

sora Copel., Philip. Joum. Sci., 6: 73. 1911, non D, basisora 

Christ, 1909. New Guinea, 
Th. crassa (Copel.) Reed, comb. nov. Basionym: Dryopteris crassa 

Copel,, Univ. Calif. Publ. Bot., 18: 220. 1942. New Guinea. 
Th. crassifolia (Blume) Ching, Bull. Fan Mem. Inst. Biol., Bot, 6: 

285. 1936, Basionym: Aspidium crassifolium Blume, Enum. PI. Jav, 

158. 1828. W, Malaysia - Malaya - Burma, Philippine Isls. 
Th. crassifolia var, motleyana (Hook.) Ching, Bull. Fan Mem. Inst, 

Biol., Bot. 6; 286, 1936. Basionym: Nephrodium motleyanum 

Hook., Syn. Fil., 266. 1867. Borneo, Malacca. 
Th. crinipes (Hook.) Reed, comb. nov. Basionym: Nephrodium crinl- 

pes Hook.. Sp. Fil., 4: 71. 1862. N. India, Malacca. 



270 P H Y T L G I A Vol. 17, no. h 

ThelypterlB cristata (L.) Nieuwl., Amer. Midi. Nat., 1: 226. 1910. 

Baaionym: PolyT^odlum cristatua L. , 3p. PI., 1090. 1753. ■ Dryo- 

pteris. 
The crjytata var„ clintoniana (D.C.Eaton) Weatherby, Rhodora, 21: 

177. 1919. Basionym: Aspldiurr crlgtatua var. cjintonlanun D.C. 

Eaton, in A„ Gray's Man., ed. 5: 665. 1867. - Dryopterla. 
Th. cruciata (Willd.) Tard„, Not. 3yst., 15(1): 90-91. 1954. Basi- 
onym: Aspidium cimciatuji V.'ilDd. in L. , 3p, PI., ed. 4, 5- 27P. 

1810, .jynon3TT!E: Fhegopteria roontbrlsonieria Fee, G*»n. Fil., 247. 

1850-.ie52; Polypodi'im sesallifolium Hook,, Sp. FU., 4: 251. 

1862; Polypodium bojerl Hook., Sp. Fil., 4: 20. 1962, Reunion, 

Seychelles, Maurice. 
Tb, crypta (Underw. et Maxon) Reed, comb, nov, Basionym: Poly- 
podium cryptum 'Jnderrf. et Maxon, Bull, Torr. Bot. Club, 29: 579, 

fig, 1902, Cuba. 
Th. cundngiana (Kunze) Reed, comb, nov, Basionym: Aspidium cuolngi- 

anum Kunze, 1: 17, t. 19, f . 2, 1840. Panama, 
Th, cijueata (C.Chr.) Reed, comb, nov, Basionym: DxTropteria cur, eat a 

C.Chr., Vid. Selsk. 3kr., VII, 10; 253, f. U2. 1913. Brazil. 
Th, carta (Christ) Reed, comb. nov. Basionym: Dryopteris carta 

Christ, Bull. Boias., II, 7: 263. 1907. Costa Kica, 
Th. cuspidata (Blume) K.Iwats., Mem. Coll. Sci. , Univ. Kyoto, Ser, 

3, 31(3): 192. 1965. Basionym: Meniscium cuspidatum Bluae. 

Jhum. n, Jav, , 114, 1828, Indochina (Tonkin, Annam), Himalayas, 

Hals-ccas, Philippine Isls. 
Th, cyatheoides (Kaulf,) Fosberg, Occ, Papers Bishop Mus., 23(2): 

30. 1962. Basionym: Aspidium cyatheoides Kaulf., Enum. Fil., 

234. 1824, Hawaiian Isls., New Guinea, Sumatra, 
Th, cyclolepis (C.Chr. et Tard,) Tagawa, Joum. Jap. Bot., 26: 20. 

1951. Basionym: Athyrivim cyclolepis C.Chr, et Tard., Bull, Mus, 

Paris, II, 6: 109, f. 3-4. 1934. Annam, 
Th, cylindrothrix (Rosenst.) K.Iwats,, Fl, Eastern Himalaya, 482. 

1966, Basionym: Dryopteris cylindrothrix Rosenst,. Fedde Re- 
pert,, 12: 246, 1913. Sikkim, 
Th, cyrtocaulos (v,A.v.R. ) Reed, comb, nov. Basionym: Dryopteris 

cyrtocaulos v,A.v,R,, Bull, Buit,, III, 5: 201. 1922, Suaatra. 
Th. cystopteroides (0,0. Eaton) Ching, Bull. Fan Mem. Inst, Biol., 

Bot. 6: 316, 1936, Basionjnn: Athyrium cystopteroides D.C.Eaton, 

Proc, Amer. Acad,, 4: 110, 1358. Synonym: Dnropteris abbrevi- 

atipinna Makino et Ogata, Joum, Jap. Bot,, 6: 10. 1929. Japan, 

Korea, Taiwan o 
Th, debilis (Kett.) Reed, comb, nov, Basionym: Phegopteris debi- 

lia Mett., Ann. Ludg. Bat., %: 223, t. 6, f, 1, I864. Araboina, 
Th. decadens (Baker) Reed, comb, nov, Basionym: Wephrodiua deca*^ 

dens Baker, Journ, Bot,, 1886: 183. 1886, Fiji, 
Th, decipiens (Clarke) Ching, Bull, Fan Mem, Inst. Biol,, Bot. 6: 

335. 1936, Basionym: Nephrodium gracilescens var. decipiens 

Clarke, Trans. Linn. Soc, II. Bot. 1(3): 514, t, 65, f. 2, 

1880. N. India, 



1968 Reed, Index Thelypteridis 271 

ThelypteriB decora (Doadn) Reed, comb, nov, Basionym: Dryopterls 
decora Doraln, Bibl. Bot., 85: 48. 1913. Australia (Queensland), 

Th, decurrenti-alata (Hook.) Reed, comb, nov, Basionym: Qynno- 
Kramma decurrenti-alata Hook,, Sp. Fil., 5: 142, t, 294. 1864. 
Japan, 

Th, decursive-pinnata (van Hall) Ching, Bull, Fan Mem, Inst, Biol,, 
Bot. 6: 275. 19360 Basionym: Polypoditm decursive-pinnatum van 
Hall, Nieuwe Verhandl. Nederl, Inst., 5: 204, tab, I8360 China, 
Japan, Tonkin, Kashndr, 

Th, decursive-pinnata forma simplex (Holt'6) Reed, comb, nov, Basi- 
onym: Phegopteids decursive-pinnata fonna simplex H,It^^ Nova 
Flora Jap,, 1: 154. 1939, Japan. 

Th, decursive-pinnata forma truncata (H,Ito) Reed, comb, nov, Basi- 
onym: Riegopterls decursive-pinnata forma truncata H,It'fe, Nova 
Flora Jap,, 1: 154. 1939, Japan. 

Th. decussata (L.) Proctor, Bull. Inst. Jamaica, Sci. Ser,, Noo 5: 
59. 1953. Basionym: Polypodium decussatum L., Sp, PI,, 2: 1093. 
1753* West Indies, Jamaica j Amer. trop. 

Th, decussata forma velutina (Sodiro) Reed, comb, nov, Basionym: 
Polypodium velutinum Sodiro, Rec, 59. 1683; Crypt. Vase, Quit., 
292. 1893. Costa Rica - Fr. Guiana, Ecuador, Peru. 

Th, deflexa (Presl) Tryon, Hhodora, 69: 5. 1967o Basionym: Nephro- 
dium deflaxum Presl, Rel, Haenk,, 1: 36, t, 5, f. 2, 1825, ^o- 
nyms: Dryopteris lindijgii CChr., Ind, Fil,, 275. 1905; Thely- 
pteris lindigii (CChr,) Alston, Joum, Wash, Acad, Sci,, 48: 
233. 1958, Colombia - Peru, 

Th, degener (Christ) Reed, comb, nov, Basionym: Dryopteris canes- 
cens var, de^ener Christ, Philip, Joum. Sci,, Bot, 2C: 199, 
1907, Philippine I sis, (Luzon), 

Th, degeneri (Copel,) Reed, comb, nov, Basionym: Qyclosorus de- 
generi Copel., Joum, Arnold Arb,, 30: 438, 1949o Fiji, 

Th, dejecta (Jenm, ) Reed, comb, nov, Basionym: Nephrodium dejecta 
Jenm., Card, Chron., Ill, 18: 640, 1895. Demerara. Brit, Quiana, 

Th. delicatula (Fee) Proctor, Rhodora, 6I: 306. (1959) I960. Basi- 
onym: Phegopteris delicatula Fee, 11® Mem. Foug,, 51, t. 20, 
f . 1, 1866. West Indies, Guadeloupe. 

Th. deltiptera (Copel.) Reed, comb, nov, Basionym: Dryopteris 
deltiptera Copel., Univ. Calif. Publ, Bot., 18: 220. 1942, New 
Guinea, 

Th, deltoidea (Swartz) Proctor, Bull. Inst. Jamaica, Sci. Ser., 
No. 5: 59. 1953; (K.Iwats,, Mem, Coll, Sci,, Univ, Kyoto, Ser« 
B, 31(1): 3I0 1964), Basionym: Polypodium deltoideum Swartz, 
Prodr,, 133. 1788, West Indies, Jamaica, 

Th, demerarana (Baker) Reed, comb, noVo Basionym: Polypodium de- 
meraranum Baker, Timehri, 5: 214. 1886, Brit, Guiana. 

Th, densa (Maxon) Tryon, Rhodora, 69: 5. 1967, Basionym: Dryo- 
pteris densa Maxon. Joum, Wash, Acad, Sci,, 34: 25. 1944, Peru, 

Th. densisora (C.Chr,) Reed, comb, nov, (Murillo, Cat, Illus, PI. 
de Cundinamarca, 2: 105o 1966 (noraen)), Basionym: Dryopteris 
densisora C,Chr,, Ind. Fil,, 26I. 1905. Synonym: Aspidium cos- 
tale Mett, ex Kuhn, Linnaea, 36: 111. I869. Costa Rica -Colombia 
- Venezuela o 



272 P H Y T L I A Vol. 17, no. U 

Th«lypteris dentata (Farsk.) E,St„ John, Aaer. Pern J»um., 26: 
Uh. 1936. (Allen, Fl. N. Zealand, 1: 52. 1961). BaBionyn: P»ly - 
podlum dentatum F«r3k„, Fl. Aegypt.-Arab,, 185. 1775. STnonyma: 
Polypodlum inolle Jacq., Collect,, 3: 188. 1769; Icon. PI, Rar,, 
t, 6A0. 1793; Dryopterla afcllls (Jacq J Hieron., Hedwigia, i*6: 
3/»8. 1907; Dryopterlo oblanclfplia Tagawa, Acta Ftiytotax. Geo- 
boto, 5: 190„ 1936; Aaridiua natalense Fee, Mem. 8: 102„ 1857; 
Nephrodiua hispidulum Peter, Peddo Repert,, Beth,, W): deecr, 
10, t. 4, f. 1-2. 1929. Pantropical: Yemen, Arabia, Tropo ATriea 
(Ivory Coast, Oubangui, Zambesla, Mocambique), India, China, 
Thailand, Taiwan, Tonga, Palau, Galapagos Isls,, Colombia, 
Paraguay, Uruguay, Argentina, Antilles. 

Th. dentata var, buchaxianii Schelpe, Joum, S, Afr, Bot., 31(4): 
265, f. Id. 1965. Mocambique, Natal, Rhodesia 

Th, dentata var. violascens (Lank) Reed, comb, nov, Baaionya: 

Aspidium violascens Link, Hort. Bot, Berol,, 2: 115. 1833. Syno- 
nym: Cyclosoirus dentatus var, violascens (Link) Abbiatti, Dar- 
winiana, 13(2-4): 540, 545. I9S4I Argentina, Brazil, 

Th, devolvens (Baker) Reed, comb, nov, Basionym: Nephrodium de - 
volvens Baker, Joum. Bot., 1885: 217. 1885. Brazil. 

Th. diaphana (Brause) Ching, Bull, Fan Mem, Inst. Biol., Bot. 10: 
251. 1941. Basionyn: Dryopteids diaptyana Brause, Engl. Bot. 
Jahrb., 56:^80. 1920. New Guinea, 

Tho dicarpa (Fee) Reed, comb. nov. Basionym: Nephrodiua dicarpua 
Fee, Gen. Fil., 305. 1850-1852, Reunion. 

Tho dichrotricha (Copel.) Reed, comb. nov. Basionym: Dryopteris 
dichrotricha Copel., Philip, Joum, Sci., 6C: 74. 1911. New 
Guinea. 

Th, dichrotrichoides (v.A.v.R,) Reed, comb. nov. Basionym: Dryo- 
pteris dichrotrichoides v.A,v.R, , Malayan Ferns Suppl. Corr., 
48. 1917. Synonyms; Dryopteris dichrotricha Copel., Philip, 
Joum. Sci,, 7C: 54. 1912; Dryopteris weberi Copel,, Philip, 
Joum. Sci., 38: 135, 1929. Philippine Isls. (Mindanao). 

Th. dicksonioides (Mett. ex Kuhn) Ching, Bull. Fan Mem. Inst, Biol,, 
Bot, 10: 251. 1941. Basionym: Phegopteris dicksonioides Mett, 
ex Kuhn, Linnaea, 36: 118, 1869o Hawaiian Isls., Tahiti. 

Th, dicranogramma (v,A,v,R,) Reed, comb, nov, Basionym: Dryopteris 
dicranogramma v.A.v.R,, Bull, Buit,, III, 5: 202. 1922. Sumatra, 
Malaya, 

Tho didymochlaenoides (Clarke) Ching, Bull, Fan Mea, Inst, Biol,, 
Bot. 6: 325. 1936. Basionym: Nephrodium gracilescens var. didy- 
mochlaenoides Clarke, Trans, Linn. Soc, Lond,, II, Bot, 1(8)1 
514, t. 68, f . 2. 1880, N. India. 

Th, didymosora (Parish in Bedd.) Reed, comb, nov, Basionym: Ne- 
phrodixim didymosorum Pari^ in Bedd., Ferns Brit. India, t, 200, 
1866, Tenas serin. Perak, Singapore, 

Th, dilatata (Hoffm.) House, New York State Mus. Bull,, 233-234: 
69. 1922, Basionym: Polypodium dilatatum Hoffm,, Deutsch, Fl,, 
2: 7. 1795. - Dryopteris. 



1968 Reed, Index Thelypteridis 273 

Thelypterls dllatata var, americana (Fisch, ex Kunze) House, New 

York State Mus. Bull. 233-234: 69. 1922, Basionym: Aspidlum 

spintilosua var, amerlcanum Fisch. ex Kunze, Amer, Joum* Sci., 

II, 6: 84. 1848. ■= Drjopteris. 
Th, dindnuta (Copel.) Reed, comb, nov, Basionym: Dryopterls dimi- 

nuta Copel., Philip. Journ. Sci., 40: 298. 1929 «, Mindanao, 
Th, dimorpha (Brause) Reed, comb. noVo Basionym: Dnropteris dimoi— 

pha Brause, Engl. Bot. Jahrb., 56: 100. 1920, New Guinea, 
Th. diplazioides (Moritz ex Mett.) Ching, Bull. Fan Mem. Inst. Biol,, 

Bot. 10: 251. 1941. Basionym: Aspidium diplazioides Morits ex 

Mett., Pheg, u. Asp., 83, no. 200. 1858 (Abh. Senskenb, Ges. 

Frankfurt, 2: 367. 1856). Ck)loobia, Venezuela, 
Th, diplazioides (Desv. ) Proctor, Bull. Inst. Jamaica, Sci, Ser. , 

No. 5: 59. 1953. Basionym: Gymnogra imia diplazioides Desv,, Mem, 

Soc. Linn, Paris, 6: 214 (diplazoidesTT 1827. - Th. linkiana . 
Th. dissimulans (Maxon et C.Chr. ex C.Qir, ) Reed, comb, nov, Basi- 
onym: Dryopterls dissimulans Maxon et C,Chr, ex C.Chr,, Vid, 

Selsk. Skr., 711, 10: 215. 1913. Cuba, 
Th, diatans (Hock.) Reed, comb. nov. Basionym: Nephrodium distans 

Hook., Sp. Fil., 4: 76 (adnota). 1862, Madagascar, Comores. 
Th. iistincta (Copel.) Reed, comb. nov. Basionym: Dryopterls di» ' 

tincta Copel., Unir. Calif. Publ. Bot., 18: 220, 1942, New 

Guinea. 
Th. divergens (Rosenst.) Reed, comb. nov. Basionym: Dryopterts 

divergens Rosenst., Fedde Repert., 13: 218. 1914. Sumatra, 
Th, diver siloba (Pre si) Reed, comb, nov, Basionym: Nephrodium di- 

versilobum Presl, ^im. Bot., 47. 1849. Luzon, Mindanao. 
Th, diversisora (Copel.) Reed, comb, nov, Basionym: Dryopterls 

diversisora Copel., Occ, Papers Bishop Mus,, 14: 54> t, 6, 1938. 

Rapa Isl, 
Tho diversivenosa (v,A,v.R,) Ching, Bull, Fan Mem. Inst, Biol,, 

Bot. 10: 251, 1941, Basionym: Diryopteris diversivenosa v,A,v.R,, 

Bull. Buit., II, No. 28: 23. 1918, Sumatra, 
Th, doodioides (Copel.) Reed, comb, nov, Basionym: Dryopterls doodi- 

oides Copel,, Philip, Joum. Sci., 60: 107, t. 11, 1936. Solomon 

I sis. 
Ih- dryopterls (L.) Slosson ex I^b., Fl. Rocky Mts., 1044. 1917. 

Basionym: Polypodlum dryopterls L., Sp. PI., 1093. 1753. " 

Dryopterls linnaeana, 
Th. dryopteroidea (Brause) Reed, comb. nov. Basionym: Alsophila 

dryopteroidea Brause, Engl, Bot. Jahrb., 56: 70. 1920. Synonyms: 

Cyathea atrispora Domin, Acta Bohem,, 9: 95. 1930; Dryopterls 

atrispora (Domin) C.Chr., Brittonia, 2: 296. 1937. New Guinea. 
Th, duclouxii (Christ) Ching, Bull. Fan Mem. Inst. Biol., Bot. 6: 

303. 1936. Basionym: Dryopterls duclouxii Christ, Bull. Acad. 

Geogr. Bot. Mans, 139. 1907. China. 
Tho dumetorum (Maxon) Tryon, Rhodora, 69: 5. 1967. Basionym: Dryo- 

pteris dumetorum Maxon, Journ. Wash. Acad. Sci., 34: 26. 1944o 

Peru. 



27U P H Y T L G I A Vol. 17, no. U 

Thelypteris duplosetooa (Copel.) Reed, conb. nov, Basionym: Cyclo- 
sorus duplosetoBus Copel,, Philip. Joum. Sci., 81: 31. 1952. 
Philippine I sis. (Palawan) . 

Th. dura (Copel.) Reed, comb, nov, Basionym: Dryopt'sris dura Copel., 
Leaflets Philip. Bot., 3: 805. 1910. Philippine Isls. (Mindanao). 

Th, ecallosa (Holtt.) Reed, comb. nov. Basionym: Cyclosorus ecal- 
losus Holtt., Gard, Bull. Singapore, 11: 269. 19A7. Malay Penin. 
(Pahang). 

Th. echinata (Mett.) Reed, comb. nov. Basionym: Aspidium echinatum 
Mett., Ann. Ludg. 3at., 1: 230. 1864, Malaya, New C^iinea. 

Tho echinospora (v.A.v.R, ) Reed, comb. nov. Basionym: Eh-yopteris 
echinospora v.A.v.R., Bull. Buit., Ill, 2: li*9. 1920, Sumatra. 

Th, edanyoi (Copel.) Reed, comb. nov. Basionym: Cyclosorus edan- 
yol Copel., Philip, Joum, Sci., 81: 37. 1952, Philippine Isls. 
(Panay), 

Th, eggersii (Hierono) Reed, comb. nov. Basionym: Nephrodiua eg- 
/tersii Hieron,, Engl. Bot. Jahrb, , 34: 441, 1904. Colombia. 

Th, elata (Mett. ex Kuhn) Schelpe, Joum. S. Afr. Bot., 31(4): 
265. 1965. Basionym: Aspidium elatum Mett, ex Kuhn, Fil. Afr,, 
131, 1868, Synonym: Nephrodium venule sum Hoolc,, Sp. Fil,, 4: 71. 
1862, non Desv, , 1827o MoCHobique, Oiinea cum ins\ilis, Fei^ 
nando Po, 

Th, elegantula (Sodiro) Alston, Joum. Wash. Acad. Sci., 48(7): 

233. 1958, Basionyra: Nephrodium elegantulum Sodiro, Crypt, Vase, 
Quit,, 243. 1893. Colombia, Ecuador. 

Th, elliptica (Rosenst,) Reed, comb, nov, Basionym: Dryopteris 
elliptica Rosenst., Med, Rijks Herb., No, 31: 6. 1917, Philip- 
pine Isls, 

The elmerorum (Copel.) Reed, comb, nov, Basionym: Dryopteris el- 
merorum Copel., Philip. Joum. Sci., 40: 295, t. 2. 1929o 
Philippine Isls. (Mindanao), 

Th, elwesii (Bak. in Hook, et Bak, ) Ching, Bull, Fan Mem, Inst. 
Biol,, Bot. 6: 308, 1936, Basionym: Nephrodium elwesii Bak, 
in Hook, et Bak., Syn. Fil,, 497, 1874. Sikkim, 

Th, engelii (Hieron,) Morton, Fieldiana, Bot,, 28: 11, 1951, Basi- 
onym: E)ryopteiria engelii Hieron., Hedwigia, 46: 339, t. 6, f. 
12. 1907. ^monym: Dryopteris pit tier! C.Chr., Sniths. Misc. 
Coll,, 52: 393. 1909. Venezuela, Colombia. 

Tho engleriana (Brause) Reed, comb. nov. Basionym: Dryopteris 
engleriana Brause, F>igl, Bot. Jahrb., 49: 19, 1912, New Guinea, 

Th, ensifer (Tagawa) K.Iwats,, Acta Riytotax, Geobot,, 21: 40. 
1964, Basionym: Dryopteris ensifer Tagawa, Acta Phytotax, Geo- 
bot., 6: 89. 1937, Synonym: Dryopteris sophoroides forma en si- 
pinna Hayata, Icon. PI. Formos., 4: 180, 1914. Taiwan, 

Tho ensiformis (C.Chr.) Tryon, Rhodora, 69: 6. 1967. Basionym: 
Diyopteris ensiformis C.Chr., Danske Vid. Selsk. Skr., VII, 
Naturv. Afd., 10(2): 269, f. 46, 1913. Costa Rica. 

Th, ensipinna (Brause) Ching, Bull. Fan Mem. Inst, Biol,, Bot, 
10: 251. 1941. Basionym: Dryopteris ensipinna Brause, Engl. 
Bot, Jahrb,, 56: 84. 1920 o New Guinea, 



1968 Reed, Index Thelyptcridis 275 

Thelypteris epaleata (C.Chr, ) Reed, comb. noVa Basionym: Dryo - 
pterJB epaleata C.ChTo, Ind, Fil. Siippl. Ill: 85. 1934o Syno= 
nym: Dryopteris francll Copel., Univ. Calif. Publ. Bot., 14: 
357. 1929, non C.Chr., 1925. New Caledonia. 

Th, equitans (Christ) Reed, comb, nov, Basionym: Nephrodium eqiii- 
tans Christ, Bull, Boiss,, II, 6: I63. 1906. Costa Rica. 

Th. erecta (Copel.) Reed, comb, nov, Basionym: Cyclosorus erecta 
Copel., Philip. Joum. Sci,, 81: 30, t. 22, 1952. Philippine 
Isls, (Leyte), 

Th„ eruboscens (Wall, ex Hook.) Ching, Bull, Fan Mem, Inst, Biol,, 
Bot, 6: 293. 1936. Basionym: Polvpodium erubescens Wall, ex 
Hook,, Sp, Fil., 4: 236, 1862. Synonym: Dryopteris reflexa 
Ching, Bull, Fan Mem, Inst. Biol,, Bot. 2: 193. 1931. N, India, 
SW China, Malaya, Laos, Tonkin, Taiwan, 

*Tho escheri (Heer) Reed, comb. nov» Basionym: Aspidiun escheri 
Beer, Tertiarfl, der Helvetiae (Schweiz), 1: 36, t. 10, f. 2. 
1855J I.e., 3: 153, t. lU, f. 9. 1859; Ett., Die Famkr, der 
Jetztwelt, 200, 1865. Synonym: Dryopteris escherd (Heer) LaMotte, 
Geol. Soc. Amer, Mem. 51: 151. 1952. Miocene: Switzerland (Up- 
per Rhone), Aff, Th. invisa et Th, raultilineata. 

The esquirolii (Christ) Ching, Bull, Fan Mem. Inst, Biolo, Bot, 6: 
301, 19360 Basionym: Dr yopteris esquirolii Christ, Bull, Acad, 
Geogr, Bot. Man, lAA. 1907, S. China, N, India, Taiwan, S, Korea, 
Japan (Honshu, Shikoku, Kyushu), I^rukyus, 

Th, esquirolii var, glabrata (Christ) K.Iwats,, Mem. Coll. Sci., 
Univ. Kyoto, Ser, B, 31(3): 182. 1965. Basionym: Dryopteris 
eberhardtii var. glabrata Christ, Not. Syst., 1: 37, 1909. 
Synonym: Thelypteris subochthodes Ching, Bull, Fan Mem. Inst, 
Biol,, Bot. 6: 305. 1936, S. Honshu, Shikoku, Kyushu, Korea 
(Isl, Quelpaert), China, 

*Tho ettinghausenii Reed, noa, nov, Basionym: Goniopteris poly- 
podioides Ett., Denkschr, K. Akad. Wiss,, Math.-Naturw, CI., 
8: 26, t. 2, f. 1-4. t. 3, f. 5. 1854. Synonyms: Lastraea 
polypodioides (Ett,) Heer, Fl. Tert Helvetiae (Schweiz), 3: I5I, 
t. 144, f. 1-3. 1859 J Phegopteris polypodioides (Ett.) Ett., 
Die Famkr. der Jetztwelt, I96, 18o5o Eocene: Switzerland. 

Th, euchlora (Sodiro) Reed, comb. nov. Basionym: Polypodium 
euchlorum Sodiro, Rec, 58. 1883 j Crypt. Vase. Quit., 290. 
1893. Ecuador, 

Th, euchlora var. inaequans (C.Chr.) Reed, comb. nor. Basionym: 
Dryopteris euchlora var. inaequans C.Chr., Kgl. Dansk. Vido 
Selsk. Skr., 7: 150. 1913. Panama, Nicaraguao 

Th, eugracilis (Copel.) Reed, comb, nov, Basionym: Lastrea eu- 
^p'acilis Copel., Gen, Fil,, 138. 1947. Synonym: Dryopteris 
gracilis Copel., Philip. Joum. Sci., 40: 294. 1929. Mindanao, 

Th, euphlebia (Ching) Reed, conb. nov. Basionym: Cyclosorus eu- 
phlebius Ching, Bull, Fan Mem. Inst. Biol., Bot. 8: 226. 1938, 
China (Kwangci, Kweichow), Tonkin, 

*Th, europaeae Reed, nom, nov, Basionym: Pecopteris deli ca tula 
Brongn., Hist, Veg. Foss., 1: 349, t. II6. 1828; Unger, Gen. 
et Sp. PI. FosSo, 181. 1850. Synonym: Aspidium delicatulum 
(Brongn,) Ett,, Die Famkr, der Jetztwelt, 198, I865. Upper 
Carboniferous: France (Fresnes), Germany (Saarbrttck); Switzer- 
land. Aff. The eontei*minum et Th, calearata. 



276 PHYTOLOOIA Vol. 17, no. h 

"nielyptoris eurostotrlcha (Baker) Raed, comb, nov, BasionTHt 
NephrodluMi euroatotrichiup B*ker, Joum, Bot., 1880: 3^9. 1880, 
Madagascar. 

Th, euryphylla (Roaenst.) Reed, comb, nor, Basionym: Dryopterla 
euryphylla RosenBt., Med. Rijks Herb. No. 31: 7. 1917. Sumatra. 

The evoluta (Bodd.) Heed, comb, nov, Basionym: Nephrodiua eTolutmc 
Bedd., Handb. Suppl,, 76. 1892, N. India. 

Th, exlgua (Kunzc ex Mett.) Reed, comb. nov. Basionyit: Aepidliui 
exlguun Kxmze ex Mett., Pheg. u. Asp., 76, no. 180. 1858; Haplo- 
dictyun exlguum Fee. Gen. Fil,, 309. 1850-1C52 (nom, confue.); 
Lastrea exl/^ua J .3a. , Joum. Bot., 3i 412. 1841 (nomen). Synonym: 
Phegopterls nervosa Fee, Gen. Fil., 244. 1850-1852, Philippines. 

The extensa (Blume) Morton, Amer. Fern Joum., 49: 113. 1959, Basi- 
onynt: AspiiLiua extensxan Blume. Enum. PI. Jav. , 156. 1828. Synonym: 
Nephrodiun wakefieldii Baker, Ann, Bot., 5: 326, 1891. Trop. Asia, 
Melanesia, Micronesia, N. Australia, E. Africa (Kenya), S. India, 
Ceylon, Burma-Male si a, Tonga, Java, Philippine Islso 

Th, falcata (Liebn. ) Tryon, Rhodora, 69: 6. 1967, Basionya: Menia- 
cium falcatum Liebm, , Dansk, Vid, Selsk. Skr., V, 1: 183. 1849. 
Synonym: Dryopterls .jurgensenii Maxon et Morton, Bull, Torr. Bot. 
Club, 65: 360. 1938, Mexico, 

Th, falcatipinnula (Copel,) Reed, comb, nov, Basionym: Dryopteris 
falcatipinnula Copel,, Philip. Joum. Sci., 6: 74. 1911. New 
Guinea, 

Th, faCcatula (Christ) Reed, comb, nov, Basionym: Dryopteris para - 
sitica var, falcatula Christ, Philip, Joum. Sci., 2C: 147. 
1907, Philippine Isls. (Mindanao, Palawan). 

Th, falciloba (Hook.) Ching, Bull, Fam Mem. Inst. Biol,, Bot. 6: 
298, 1936. Basionym: Lastrea falciloba Hook., Joum. Bot., 9: 
337. 1856, Himalayas, Burma, Tonkin, China. 

Th, farinosa (Brause) Reed, comb, nov, Basionym: Dryopteris fari- 
ng sa Brause, Engl. Bot, Jahrb,, 56: 111, 1920, New Guinea, 

Th, fasciculata (Foum,) Ching, Bull. Fan Mem. Inst. Biol., Bot, 
10: 251, 1941. Basionym: Aspidiua fascicvilatun Foam,, Ann. Sci, 
Nat., V, 18: 295. 1873. New Caledonia. 

Th, fatuhivensis (E,Brown in E, et F, Brown) Ching, Bull, Fan, Mem. 
Inst, Biol,, Bot, 10: 251. 1941. Basionym: Dryopteris fatu- 
hivensis E, Brown in E, et F, Brown, Bemice P, Bichop Mus, Bull., 
89: 27, f, 8, 1931. Marquesas, 

Th, feci Moxley, Bull, So, Calif, Acad, Sci,, 20: 35. 1921. - Th, 
puberula, 

Th, fendleri (D,C,Eaton) Reed, comb, nov, Basionym: Aspidium fend- 
leri D.C.Eaton, Mem, Amer, Acad,, n.s., 8: 210, IsSo, Vene- 
zuela, Colombia, 

Th, ferox (Blua.e) Reed, comb. nov. Basionym: Aspidium ferox B lume. 
Enum, PI, Jav., 153. 1828. S, Thailand, Malesia - Philippine 
Isls,, Perak, ?Kumoan, 

Th. filix-mas (L.) Nieuwl., Amer. Midi. Nat,, 1: 226, 1910, Basi- 
onym: Polypodium filix-mas L,, Sp. PI., 1090. 1753. - Dryopteris. 



1968 Reed, Index Thelypteridis 277 

Thelypteris finisterrae (Brause) Reed, comb. nor. Basionym: Dryo- 
pterls finisterrae Brause, Engl. Bot, Jahrb,, /t9: 20. 1912. New 
Guinea, 

The firraa (Baker ex Jenm. ) Proctor, Bull. Inst. Jaoaica, Sci. Ser. , 
No. 5: 60. 1953« Basionym: Nephrodiua fimmm Bak. ex Jennu, 
Joum. Bot., 1879: 260, 1879. Jamaica. 

Tho firnula (Baker) K.Iwats., Acta Phytotax. Geobot., 21(5-6): 170. 
1965. Basionym: Polypodium fimulum Baker, Kew Bull., 1893: 211„ 
1893. Borneo o 

*Th. fischeri (Heer) Reed, comb. nov. Basionym: Lastrea ( Gonio- 
pteris ) fischeri Heer, Fl. Tert. Helv., 1: 34, t. 9, f. 3. 1855. 
Synonyms: Cyclosorus fischeri (Heer) Kolakorskii, Akad, Nauk 
Greezinskoi SSR, Sukhum Bot. Sad, Monografii 1(1964): 23, t. 1, 
f. 1, 1964; Lastrea knight iana Newberry, U.S. Nat. Mas. Proc., 
5: 503. 1882 (1883); Aspidium goldianua Lesq., 7th Ann. Repto, 
U.S. Geol, Surv, Terr., 393. 1873 (1874); Dryopteris Integra 
Knowlton, U.S. Geol. Surv., Prof. Paper 155: 17, t. 1, f. 5, 
I93O; Dryopteris lesquereuxii Knowlton, Bull. U.S. Geol, Sui^,, 
696: 284. 1919 (1920); Dryopteris richardaoniana Knowlton, U.S. 
Geol. Surv., Prof. Paper 155: 20, t. 2, f. 3-5. 1930. Eocene: 
Oregon, Alberta, British Columbia, Colorado; Pliocene: Georgia 
in SSR; Cretaceous: Colorado, 

Th, flaccida (Blume) Ching, B\ai, Fan Mem. Inst. Biol,, Bot. 6: 336. 
1936. Basionym: Aspidium flaccidum Blume, Enum, PI. Jav. , I6I, 
1826, Malaya - India - China, Japan, 

Th, flavovirens (Rosenst,) Reed, comb, nov, Basionym; Dryopteris 
flavovirens Rosenst,, Fedde Repert,, 10: 334. 1912© New Guinea, 

Th, flexilis (Christ) Ching, Bull. Fan Mem. Inst, Biol., Bot, 6: 
283. 1936, Basionym: Aspidium flexile Christ, Bull, Acad, Geogr, 
Bot. Mans, 1902: 252, 1902, China (Szechwan, Kweichow). Ano- 
nyms: Aspidium melanorhizum Christ, Bull, Soc, Bot, Ital,, 1901: 
295. 1901, non Desv., 1827; Dryopteris subthelypteris CChr,, 
Ind. Fil,, 296. 1905. 

Th. forsteri Morton, Contrib. U.S. Nat, Herb,, 38(2): 60. I967, 
Basionym: Polypodium invisum G. Forst., Fl. Ins, Austr, Prodr,, 
1786. Tahiti, Polynesia, 

Th. foxLi (Christ) Reed, comb, nov, Basionym: Dryopteris foxli 
Christ, Philip. Joum. Sci., 2C: 208. 1907; ( Nephrodium foxii 
Copel., ms8. ). Philippine Isls. (Luzon - Mindanao). 

Th, fragilis (Baker) Alston, Bol. Soc. Broter., 2 ser,, 30: 25, 
1956, Basionym: Polyt>odium fragile Baker, Joum. Linn. Soc, 
16: 203. 1877, non L, Synonym: Dryopteris fragilis (Bak.) C.Chr., 
Ind. Fil., 266, 1905; Phegopteris fragilis Kuhn, v. Deck. Reis, 
3(3): Bot. 66, 1379. Madagascar, (Probably should be based on 
Kuhn). 

Th. fragrans (L.) Nieuwl., Amer. Midi. Nat,, 1: 226. 1910. Basio- 
nymz Polypodium fragrans L. , Sp, PI., 1089. 1753o = Dryopteris. 

^. fragrans var, hookeriana Femald, Rhodora, 25: 3, 1923. » 
Dryopteris, 



278 P H Y T L G I A Vol. 17, no, 1; 

Thelyptftris francoana (Foum.) R^<5d, conb. nor. Baeionyn: As- 

pidlum f rancoanua Poum. . Bull. Soc. Bot. France, 19: 255.. 1872. 
NicMrHgua, Costa Rica, Ecuador. 

Th, frieali (Brauae) Schelpo, Bol. Soc, Broteriana, Ser. 2A, 41: 
216, 1967 (1968), Basionym: Dryopt^ria frieali brauae in Fries, 
Wiss. Ergebn. Schwed. Hhodesia-Gongo Exped, 1911-12, Bot. 1: 1. 
1914. Rhodesia. 

Th. fukienenais (Ching) Reed, corab. nov. Basionym: Cyclosorua fuki - 
anenaia Ching, Bull. Fan Mem. Inat. Biol., Bot. 8: 209. 1938. 
China (Fukien). 

Th. fulgena (Brauae) Ching, Bull. Fan Meo. Inst. Biol., Bot. 10: 

251. 1941. Basionym: Dryopteris ful/^ens Brause, Sigl. Bot. Jahrb., 
56: 39. 1920. New Guinea. 

Th, funckii (Mett. in Triana et Planch.) Alston, Joum. Wapn. Acad, 
Sci., 43(7): 233. 1958. Basionym: Aspidium funckii Mett, in 
Triana et Planch., Ann. Sci. Nat., V, 2: 2^6. 1864, Colombia, 
Venezuela, Peru, Ecuador, Costa Rica, 

Th. funestra (Kunze) Alston, Kew Bull. 1932: 309. 1932. Basionym: 
Aspidium fune strum Kunze, Linnaea, 9: 96. 1834. » Ctenitia pro- 
tenaa var, funestrao 

Th, furva (Maxon) Tryon, Rhodora, 69: 6. 1967. Basionym: Dryo- 
pteris furva Maxon, Joum. Wash. Acad. Sci., 34: 24. 1944, Peru. 

Th. galanderi (Hieron. ) Abbiatti, Darwiniana, 13(2-4): 566. 1964. 
Basionym: Aspidium galanderi Hieron., Engl,, Bot. Jahrb,, 22: 
369. 1896 (1897). Argentina. 

Th. gardneriana (Baker) Reed, comb, nov, Basionym: Nephrodiuw 
gardnerianua Baker, Fl. Braa., 1(2): 474. 1870; ( Aspidium gar*- 
nerianum Kunze, msc). Synonym: Dryopteris densiloba C.Chr., 
Ind. Fil., 261. 1905. Brazil. 

Th. germaniana (Fee) Proctor, Rhod^ora, 61: 3O6. (1959) I960. Basi- 
onym: Phegopteria germaniana Fee, 11 Mem. Foug., 55, t. 13, f. 
2. 1866. We at Indies. Cuba, Guadeloupe, 

Tho ghieabreghtii (Hook.) Morton, Contrib, U.S. Nat. Herb., 38(2): 
45. 1967. Baaionym: Polypodiiun crenatum var. ghieabreghtii 
Hook., Sp. Fil., 5: 3. 1864. 5ynonytt: Goniopteri8 mollis F^e. 
Gen. Fil,, 252. 1850-1852. Mexico - Panama, 

Th, gigantea (Mett.) Tryon, Rhodora, 69: 6. 1967; Morton, Con- 
trib. U.S. Nat. Herb., 38(2): 51. 1967. Baaionym: Menis- 
eium giganteum Mett., Fil. Lochl., 1: 19. 1856. ^rnonym: Dryo- 
pteris aimplicifrons C.Chr., Ind. Fil., 486. 1906. Peru. 

Tho glabra (Brack.) Ching, Bull. Fan Mem. Inat. Biol., Bot. 10: 
251. 1941. Baaionym: Laatrea glabra Brack. . Exped. Exp,, 16: 
200, 1854, Hawaii, Tahiti, 

Th, glabrata (Mett, ex Kuhn) Tard., Not. Syat., 14: 344. 1952; 
Mem, IFAN, 28: 120. 1953. Basionym: Aspidium s 3-*bratuM Mett, 
ex Kuhn, Fil. Afr,, 133. 1868, - Athyrium. 

Th o glabrata var. hirauta Tard., Not. Syst., 14: 344. 1952. 

Th« glandulifera (Brack, ) Reed, coaib. nov. Basionym: Gonjo- 
pteria glandulifera Brack., Expl. Exp., 16: 29. 1854. Samoan 
lals. (Tutuila). 



1968 Reed, Index Thelypteridis 279 

Thelypteris glanduligera (Kunze) Ching, Bull, Fan Mem. Inst. Biol,, 
Bot, 6: 320. 1936, Basionym: Aspidium glanduligerum Kunze, Anal, 
Pteridc, kh. 1837, Synonym: Dryopteris thelypteris var, koreana 
Nakai, Bot. Mag. Tokyo, 45: 97. 1931. Himalayas - Japan, 

Th. glanduligera var, atripes Tard, in Lecomte's Flore Gen, de 
I'Indochine, 7(2), fasc. 8: 365. 1941. Annam. 

Tho glanduligera var. elatior (D.C.Eaton) Kurata, in Namegata et 
Kurata, Enun, Jap. Pterid., 317, 343. 1961. Basionym: Athyrium 
eystopteroides var, elatius D,C, Eaton, ProCo Amer. Acad,, 4: 

110. 1858. Japan, 

T h. glanduligera var. hyalostegia (Holto) H.Ito, Bot. Mag, Tokyo, 

52: 589. 1938, Basionym: Dryopteris glanduligera var, hyalo~ 

stegla H.Ito, Bot. Mag. Tokyo, 49: 363. 1935. » Th. angustifrons. 
Th, glanduligera yaro koreana (Nakai) H.Ito, No-ra Flora Jap., 1: 

130. 1939. Basionym: Dryopteris thelypteris var, koreana Nakai, 

Bot, Mag, Tokyo, 45: 97. 1931. Korea, 
Th. glanduligera Ching, var. typicaj H.Ito, Nova Flora Jap., 1: I3O, 

1939. Taiwan; Himalayas - Japan, 
Th. glandulosa (Desv,) Proctor, Rhodora, 61: 306, (1959) I960, Basi- 
onym: Polyp odium glandulosum Desv,, Berl, Mag., 5s 317. 1811, 

West Indies, 
Th. glandulosolanosa (C.Chr, ) Tryon, Rhodora, 69: 6, 1967. Basionym: 

Dryopteris glanduloso-lanosa C.Chr,, Dansk Bot. Aik,, 9(3): 61, 

1937. Peru. 
Th. glaucescens (Brause) Ching, Bull. Fan Mem. Inst. Biol., Bot. 

10: 251. 1941. Basionym: Dryopteris glaucescens Brause, Engl, 

Bot. Jahrb., 56: 85. 1920, New Guinea. 
Th, glaziovii (Christ) Reed, comb, nov, Basionym: Aspidium gla~ 

ziovii Christ, Bull. Boiss., II, 2: 633. 1902, Brazil. 
Th, globulifera (Brack,) Reed, comb, nov. Basionffn: Lastrea filo- 

bulifera Brack., Expl. Exp., 16: 194. 1854.. Hawaii, 
Th. glutinosa (C.Chr.) Morton, Amer. Fern Joum„, 53: 66, I963. 

Basionsrm: Dryopteris glutinosa C.Chr., Svensk Vet. Akad, Handl,, 

111, 16(2): 18. 1937. Haiti. 

Th, goedenii (Rosenst.) Reed, comb, nov, Basionym: Dryopteris 
goedenii Rosenst,, Fedde Repert., 4: 292, 1907. S. Brazil, 

*Th. goeppertii (Ett,) Reed, comb. nov. Basionym: Aspidium goep- 
perti Ett., Die Farnkr. der Jetztwelt, 198, 1865. Synonym: 
Aspidites deeussatus Goepp,, Syst, Fil. Foss,, 369, t. 26, 
f . 1-2. I836; Pecopteris d ecu 3 sat a Goepp., Ubersicht d. foss. 
Flora Schlesiens, 215. 1844. Upper Carboniferous: Silesia. 
Aff. Th. oligocarpa et Th, multilineata. 

Th. goggilodus (Schkuhr) Small, Ferns S.E. States, 248 cum tab., 
475, 1938. Basionym: Aspidium goggilodus Schkuhr, Krypt, Gew,, 
1: 193, t. 33c. 1809. - Th, totta. 

Th, goldiana (Hook, ex Goldie) Nieuwl., Amer. Midi. Nat., 1: 226o 
1910. Basionym: Aspidium goldianum Hook, ex Goldie, Edinb, 
Philos. Joum., 6: 333. 1822, » Dryopteris, 

Th, gongylodes (Schkuhr) Small, Ferns S.E. States, 248, 475. 1938; 
Morton, Contrib. U.S. Nat. Herb., 38(2): 73. 1967; K.Iwats., 
Mem. Coll. Sci., Univ. Kyoto, Ser. B, 31(1): 31. 1964« Basic- 



280 P }i Y T L G I JL Vol. 17, no. h 

nyva: Aspldiua j^ongylodes (pro err, ^o^gilodus) Schkuhr, Krypt, 
Gew., 1: 193. 1809. - Th. totta. 

Tholypteria gracilescens (Blame) Ching, Bull. Fan Kea. Inst. Biol., 
Bot. 6: 327. 1936, Basionym: Aapidluja gracilescena Bluiae, SnujB. 
PI. Jav. , 155. 1828. Synonym: Dryopt^ris aublaxa Hayata. Icon. 
Fl. Formosa, /*: 183, f . 121. 191A; Dryopterls arisanensis Rosenst,, 
Hedwigia, 56: 3^0, 1915. S. India, Perak, Maleaia, Taiwan, Philip- 
pine I si 8. to Javm and New Guinea, China (Yunnan), Japan (Kyushu). 

Tho gracilis (Hew.) Proctor, Bull. Inst. Jamaica, 3ci. Ser., No, 5: 
60. 1953. Basionym: Gfannnogramma gracilis Hew., Mag. Nat. Hist., 
II, 2: /*57. 1838. West Indies. 

Th, grammitoides (Christ) Ching, Bull, Fan Mem. Inst. Biol., Bot, 
6: 317. 1936. Basionym: Aspidium grammitoides Christ, Bull. 
Herb, Boiss., 6: 193. 1398. Philippine I sis. (Luzon). 

Th, grantii (Copel,) Ching, Bull. Fan Mem. Inst. Biol., Bot. 10: 
251. 19WL, Basionym: Dryopterls grantii Copel,, Bemice P, Bis- 
hop Mu8, Bull., 93: 8, t. 7C. 1932. Society Isls., Tahiti. 

The granulosa (Presl) Reed, comb, nor, Basionym: Polypodium granu- 
le sua Presl, Rel. Haenk,, 1: 24, t, U, f. 2. 1825. Luzon, Timor. 

Tho gregaria (Copel.) Reed, comb, nor, Basionym: Cyclosorus gre- 
narius Copel,, Joum. Arnold Arb, , 24: 440. 1943, New Guinea, 

Th, gretheri (Wagner) B.C.Stone, Micronesica, 2: 3. 1966, Basionym: 
Laatrea gretheri Wagner, Pacific Sci,, 2(3): 214, f. 1. 1948. 
Mariana I si, (Rota). 

Th, griffithii (Moore) Reed, comb. noT. Basionym: Dictyocline 

griffithii Moore, Card. Chron., 1855: 854. 1855; Ind. Fil., LH. 
1857, Synonyms: Aspidium griffithii (Moore) Diels, Nat. Pfl*- 
fam., 1(4): 186, 1899; Stegnograrama griffithii (Moore) K.Iwats., 
Mem. Coll. Sci,, Unir, Kyoto, Ser, B, Biol, 31(1): 20, 1964. 
Himalayas to Japan, Taiwan, Tonkin, N. India. 

Tho griffithii rar, wilfordii (Hook,) Reed, comb. noTo Basionym: 
Hemionitis wilfordii Hook,. Fil, Exot., t. 93, 1859. Synonyms: 
Dictyocline griffithii var, wilfordii (Hook.) Moore, Ind, Fil., 
317, 1861; Dictyocline griffithii var. pinnatifida (Hook.) Bedd., 
Ferns Brit. Ind., t. 155. 1866; Dictyocline wilfordii J.Sm., 
Hist. Fil., U9. 1875. Japan, 

Th, grisea (Baker in Hook, et Bak.) Ching, Bixll, Fan Mem. Inst. Biol., 
Bot. 6: 331. 1936, Basionym: Nephr odium griseum Baker in Hook, 
et Bak,, Syn. Fil,, 271. 1867, S. India, 

Th, gmmovfii (Belle) Reed, comb, nov, Basionym: Aspidium grunowii 
Belle, Bonplandia, 3: 123, 1855. Ins, Caboverdicae, 

Th, goadalupensis (Wikstr, ) Proctor, Bull. Inst, Jamaica, Sci. Ser., 
No. 5: 60. 1953. Basionym: Polypodium guadalupense Wikstr., 
Vet, Akad. Handl., 1825: 435. 1826, Synonym: Polypodium sco- 
lopendrioides L., Sp. PI., ed. 2, 1585. 1763, non L. , 1753. 
Guadeloupe. 

Th, gueintziana (Mett,) Schelpe, Joum. S. Afr. Bot., 31: 262. 
1965, Basionym: Aspidium gueintzianum Mett,, Pheg. u. Asp., 
83, n. 201. 1858. Trop. and S. Africa, Natal, Reunion. 



1968 Reed, Index Thelypteridis 281 

Thelypteris guentheri (Rosenst.) Ching, Bull. Fan Mem. Inst. Biol,, 
Bot. 10: 251. 1941. Basionym: Dryopteris guentheri Rosenat., 
Fedde Repert., 25: 59. 1928, Bolivia. 

Th. guineensis (Christ in A, Cheval.) Alston, Bull, Brit, Mas, 

(Nat, Hist,), Bot. 1: 48. 1952. Basionym: Dryopteris guineensis 
Christ in A. Cheval., Joum, de Bot., 22: 22. 1909. Fr. Guinea. 

Th, gustavii (Bedd.) Reed, comb. nov. Basionym: Nephrodium gustavi 
Bedd., Joum, Bot., 1893: 227. 1893. India. 

Th. gymnccarpa (Copel.) Morton, Amer. Fern Joum., 56(4): 179. 1966, 
Basionym: Dryopteris gymnocarpon Copel. in Elmer, Leaflets Phi- 
lip, Bot., 3: 807. 1910, Philippine I sis. (Mindanao), 

Th, gymnocarpa subsp, amabilis (Tagawa) tforton, Amer, Fem Joum,, 
56(4): 179. 1966. Basionym: Leptogramma amabilis Tagawa, Acta 
Phj'totax. Geobot,, 7: 76. 1938, Ryuksu Isls,, Okinawa, 

The gymnopoda (Baker) Reed, comb, nov, Basionym: Nephrodixim gycK 
nopoda Baker, Trans. Linn, Soc,, II, Bot,, 4: 252. 1894. Syno- 
nym: Dryopteris athyriocarpa Copel,, Philip. Joum. Sci, Boto, 
3: 344o 1909. Borneo. 

Th, haenkeana (Presl) Reed, comb, nov, Basionym: Nephrodium haen- 
keanum Presl, Epim, Bot., 46. 1849, Synonym: Nephrodium ser- 
ratum Presl, 1825, non Aspidium serratum Swartz, I8O6, nee Menis- 
cium serratum Cav, , I8O3. I'felesia - Polynesia, 

Th, halconensis (Copel.) Reed, comb, nov, Basionym: Cyclosorus 
halconensis Copel,, Philip, Joum. Sci,, 81: 29. 1952. Philip- 
pine I sis, (Mindoxxi). 

Th, hallieid (Christ) Reed, comb, nov. Basionym: Aspidium hal~ 
lieri Christ, Ann. Jard. Buit,, II, 5: IO6. 1905. Borneo, 

Iho hamifera (v.A.v.R, ) Reed, comb, nov, Basionym: Dryopteris 

hamifera v.A.v.R,, Bull. Jard. Bot. Buit., II, No. XVI: 12, 1914. 
Sumatra. 

Th, handroi (Brade) Reed, comb. nov. Basionym: Dryopteris handroi 
Brade, Arquiv. Jard, Bot. Rio de Janeiro, 18: 24. 1962-65, Brazil. 

Th, harcourtii (Domin) Ching, Bull, Fan Mem, Inst. Biol,, Bot, 

10: 251. 1941. Basionym: Dryopteris harcourtii Domin, Kew Bull., 
1929: 219. 1929; Mem. R. Czech. Soc, Sci., n.s., 2: 198, t. 34, 
f. 1. 1929, Dominica, 

Th, harrisonii (Baker) Reed, comb, nov, Basionym: Nephrodium 

harrisonii Baker, Ann. Bot,, 5: 326 ( harrisoni) , 1891, Synonym: 
Nephrodium stenophyllum Baker, Joum. Bot,, 1884: 363. 1884, 
non Sod. 1883 . Costa Rica. 

Th. harx'eyi (Hett. ex Kuhn) Proctor ex Iwats,, Amer. Fem Journ., 
53: 133. 1963. Basionym: Aspidium harveyi Mett. ex Kuim, Lin- 
naea, 36: 115. 1869. Synonym: Dryopteris euaensis Copel,, Univ, 
Calif. Publ. Bot., 12: 391. 1931. Polynesia, Tonga Isl, 

Th, hastata (Fee) Proctor, Bull. Inst, Jamaica, Sci, Ser,, Noo 5: 
60. 1953. Basionym: Goniopteris hastata Fee, 11 Mem., 65, t. 
18, f. 1 (p.p.). 1866, West Indies, Jamaica, 

Th, hastato-pinnata (Brause) Reed, comb, nov, Basionym: Dryopteris 
hastato-pinnata Brause, Engl. Bot. Jahrb., 56: 112, 1920. New 
Guinea. 



2B? P H Y T L G I A Vol. 17, no. h 

Tholypteris hattorii (M.Ito) Tagawa, Acta Phytotax. Goobot., 5* 
195. 1936. Basionym: Dryopttris hattorii H.Ito, Bot. Mag. 
Tokyo, 49: 359. 1935. Japan (K:Ai8yu, Sikoku, Honayu), China 
(Szechuan). 

Th, hattorii var. nemoralis (Ching) Kurata in Namegata et Kurata, 
Enum. Jap. Pterid., 317, 343. 1961. Bagionym: Thelypteria ne- 
mo rails Ching, Bull. Fan Mem. Inst, Biol., Bot. 6: 338. 193^ 
Japsm, Cent. China, 

Th, hawaiiensis Reed, nom. nov, Basionym: Stegj^ograama sandvi - 
censis Brack., Expl. Exp., 16: 26, t. U, t. 2. 1B54. Hawaii. 

Th, heineri (C.Chr.) Reed, comb, nor, Basionym: Dryopteris hei - 
meri C.Chr., Fedde Repert., 6: 3BO-381. 1909. Brazil. 

Th. henrlqueslJ (Baker in Henri q. ) Tard., Not. Syst,, 14: 344. 
1952, Basionym: Polypodium henriquesii Baker in Henriq,, 
Bol. Soc, Brot., 4: 154, t. 1. (1886) 1887, Guinea, S. Thome. 

Th. herbacea Holtt., Gard. Bull, Singapore, 11: 268. 1947. Malay 
Peninsula, 

Th, herzogii (Rosenst,) Tryon, Rhodora, 69: 6, 1967= Basionym: 
Dryopteris herzoj^ ii Rosenst., Meded. Rijks Herb, Leiden, Nr. 
19: 15. 1913. BoUvia, 

Th, heterocarpa (Blume) Morton, Amer. Fern Joum., 49: 113. 1959. 
Basionym: Aspidium heterocarpon Blume, Enura. PI. Jav, , 155. 
].828. Micronesia (Kusaie, Palau, Ponape), Malesia, Hongkong, 
Malaysia to New Guinea, Bismarck Archipelago, 

Th, heterocarpa var, glaucostipes (Bedd.) Reed, comb. noVo Basi- 
onym: Nephrodium glauco stipes Bedd., Handb. Suppl., 80, 1892. 
Perak, Malaya, Borneo, Sumatra, Mentawi Isls, 

Th, heteroclita (Desv.) Ching, Bull. Fan Mem. Inst. Biol., Bot, 
10: 252, 1941. Basionym: Polypodium heteroclitijg Desv,, Berl. 
Mag,, 5: 318. 1811. West Indies, Jamaica. (Proctor, Bull. 
Inst. Jamaica, Sci. Ser,, No. 5: 60. 1953). 

Th, heterophlebia (Baker) Reed, comb, nov, Basionym: Polypodium 
heterophlebium Baker, Joum. Bot., 1884: 363. 1884. Costa Rica. 

Th, heterophylla (Pre si) K.Iwats,, Mem, Coll. Sci., Univ. Kyoto, 
Ser. B, 31(1): 32. 1964. Basionym: Haplodictyum heterophyllxut 
Pre si, Epim, Bot., 51. 1849. Philippine Isls. 

Th, heteroptera (Desv.) Tard. in Humbert, Fl, Madagas,, Fam. 5: 
276, f . 38 (6-10). 1958, Basionym: Nephrodium heteropterum 
Desv,, Prodr,, 256. 1827. Reunion, Madagascar. 

Th, hewittii (Copel.) Reed, comb, nov. Basionym: Dryopteris 

hewittil Copel., Philip. Joum. Sci., Bot, 3: 3U.. 1909. Borneo. 

Th, hexagonoptera (Michx.) Weatherby, Rhodora, 21: 179. 1919. 
Basionym: Polypodium hexagonoptenun Hicfvx. , Fl. Bor. Amer,, 
2: 271. I8O3. Texas to N. Florida, N to Minnesota and Quebec, 

Th. hexagonoptera foma furcata (Reed) Reed, comb, nov, Basio- 
nym; Dryopteris hexa g onoptera fonna furcata Reed, Amer, Fern 
Joum,, 37: 83, 1947. Pennsylvania. 

Th, hexagonoptera forma simonsii (Reed) Reed, comb. nov. Basio- 
nym: ^22Et£Ei2. hexagonoptera forma simonsii Reed, Amer, Fern 
Joum,, 35: 105. 1945. I'laryland. 



1968 Reed, Index Thelypteridis 283 

Thelypteris hickenii (Abbiatti) Reed, comb, nov, Basionym: qyclo- 

sorus hickenli Abbiatti, Darwiniana, 13(2-4): 537-538, f. 1, pi. 

1. 1964. Argentina, 
Tho hieronjinusii (CChr.) Reed, comb, nov, Basionym: Dryopteris 

hieronymusii CChr., Vid. Selsk. Skr., VII, 4: 307. 1907. Colon- 

bia. 
Th, hillii (Baker) Reed, conb. nov, Basionym: Polypodium hlllil 

Baker in Hook, et Bak., Syn. Fil,, 505. 1874. Sjmonym: Gonio- 

pteris ghiesbrechtii Bail.. Handb. Ferns Queensland, 40. 1874. 

Australia (Queensland ) , 
Th, himalaica (Ching) Reed, conb. nov. Basionym: Leptogramma hi- 

malaica Ching, Sinensia, 7: 100, t. 6. 1936, N.W.India, 
The hirsutipes (Clarke) Ching, Bull, Fan Mem. Inst. Biol., Bot. 6: 

314. 1936. Basionym: Nephrodium gracilescens var, hirsutipes 

Clarke, Trans. Linn. Soc., II. Bot., 1(8): 514, t. 67, f. 1. 

1880, N, India, China (Yunnan), Japan, Khasia. 
;ni, hirtirachis (CChr.) Ching, Bull. Fan Mem. Inst, Biol,, Dot. 

6: 271, 1936, Basionym: Dryopteris hirtirachis CChr., Ind. 

Fil. Suppl. II: 15. 1917. = Th. paludosa. 
Th, hirtisora (C.Chr,) Reed, comb, nov, Basionym: Dryopteris 

hirtisora CChr,, Contrib. U.S. Nat. Herb., 26: 277 « 1926, 

China (Yunnan), N. Thailand, 
Th, hirto-pilosa (Rosenst.) Reed, conb. nov. Basionym: Dryopteris 

hirto-pilosa Rosenst,, Med. Rijks Herb., No. 31: 7. 1917. 

Philippine Isls. 
Th, hispida (Brause) Reed, comb. nov. Basionym: Dryopteris his- 

pida Brause, Engl. Bot. Jahrb,, 56: 102. 1920, Synonym: Dryo- 
pteris hispidulifcrmis CChr., Ind. Fil, Suppl. Ill: 88. 1934. 

New Guinea, 
Th, hispidifolia (v.A.v.R, ) Reed, comb, nov, Basionym: Dryo- 
pteris hispidifoli a v.A.v.R. , Bull. Jard. Bot. Buit., II, No, 

XX: 15. 1915. Borneo. 
Th. hispidula (Decne.) Reed, comb, nov, Basionym: Aspidium his- 

pidulum Decne., Mouv. Ann. Mus., 3 J 346. 1834. Eo Malesia, 

Philippine Isls,, New Guinea, Samoa, 
Th. hokouflnsis (Ching) Reed, comb. noVo Basionym: Cyclosorus ho- 

kouensis Ching, Bull. Fan Mem, Inst, Biol,, Set. II, 1: 289. 

1949. China (Yunnan), 
Th, horridipes (v,A.v,R, ) Reed, comb, nov. Basionym: Dryopteris 

horridipeg v.A.v.R., Bull. Buit., II, No. 28: 23. 1918. Sumatra. 
Th. hosei (Baker) K.Iwats., Acta Phytotax. Geobot., 21(5-6): 170. 

1965. Basionym: Meniscium hosei Baker, Journ. Linn. Soc, 22: 

230. 1886, Borneo. 
Th. hostmannii (Klotzsch) Morton, Contrib. U.S. Nat. Herb,, 38: 

59. 1967. Basionym: Polypodium hostmannii Klotzsch, Linnaea, 

20: 397. 1847. Surinam. 
Th, houi (Ching) Reed, comb, nov, Basionym: Cyclosorus houl Ching, 

Bull. Fan Hem. Inst. Biol., Ser. II, 1: 290. 1949. China (Kwei- 

chow) . 



28U P H y T L Q I A Vol. 17, n&. li 

Thelypterle hudaonian* (Brack.) Reed, comb, nor, Baflionym: Ne::,hro- 
dium hudaonlanua Brack., Expl. Exp., 16: 188, t, 25. 165U. Sand- 
vrLch I si 9, 

Tho hunst«iniana (Brause) Reed, comb. nor. Basionym: DryopterlB 
hunatolnlana Brause, Engl. Bot. Jahrb., 56: 79. 192Ci. New CJuinee. 

Th, hydrophila (Fee) Proctor, Rhodora, 6l: 306. (1959) I960. Basi- 
onym; Phe/^opterls hydrophila Fee, 11 Mem. Poug., 56, t. 13, f« 
3. 1866^ Guadeloupe. 

Th. illicita (Christ) Reed, comb. nor. Basionym: Dryoptarie il- 
licit Chriat, Bull. Soc. Bot. Geneve, II, 1: 225. 1909. Costa 
Rica. 

Th, imbricata (Liebm, ) Reed, comb, nor, Basionym: Polypodiua ii>- 
bricatum Liebm., Vid. Selsk. Skr., V, 1: 210. 1849. Synonym: 
Aspidium Tarians Mett. ex Kuhn, linnaea, 36: llA. 1869. Mexico, 

Tho immersa (Blume) Ching, Bull, Fan Mem. Inst. Biol., Bot. 6: 
306. 1936. Basionym: Aspidium immersum Blume, Enuin^ PI, Jav., 
156, 1328, Malaysia, Java, Hainan, Philippine Isle (Mindanao). 

Th, inrponens (Cea,) Reed, comb, nor, Basionym: Polypodium icpo- 
neng Cea,, Rend. Ac, Napoli, 16: 27, 29. 1877, New Guin«a. 

Th, inqiressa (Desv.) Reed, comb, nov, Basionym: Mephrodium !»- 
greasum Desv, . Prodr., 259. 1827, Timar, 

Th, incerta (Domin) Reed, comb, nov, Basionym: Dryopteris incerta 
Doain, Bibl. Bot., 85: U9. 1913. Australia (Queensland). 

Th. incluaa (Copel.) Reed, comb, nor, Basionym: Dryopteris in- 
elusa Copel,, Univ. Calif, Publ, Bot., U: 373, t. 57. 1929. 
Sumatra. 

Th, ineonspicua (Copel.) Ching, Bull. Fan Mem, Inst, Biol., Bot, 
10: 252, I94I0 Basionym: Dryopteris ineonspicua Copel., Philip. 
Journ. Sci., 12C: 55. 1917, Borneo. 

Th. indica (v.A.v.R,) Reed, «orab, nov, Basionym: Dryopteris in- 
dica v.A.v.R., M«l, Ferns, 224. 1909. India, Malacca, 

Th, indochinensis (Christ) Ching, Bull, Fan Mem, Inst. Biol,, Bot. 
6: 327. 1936. Basionym: Dryopteris indochinensis Christ, Joum, 
de Hot., 21: 231, 263. 1908. Anname 

Th. insignis (Mett. in Triana et Planch.) Ching, Bull. Fan Mem. 
Inst, Biol., Bot. 10: 252, 1941. Basionym: Aspidium insigne 
Mett, in Triana et Planch,, Ann, Sci, Nat,, V, 2: 247. I864, 
Colombia, 

Th. insularis (K.Iwats,) K.Iwats., Mem. Coll. Sci,, Univ. Kyoto, 
Sor. B, 31(3): 195. 1965, Basionym: Abacopteris insularis 
K.Ivrats, , Acta Phytotax. Geobot., 18: 6, 1959. Ryukyus, Okinawa 
Isl, 

Th, intermedia (Muhlo ex Willd.) House, New Yoric State Mus. Bull, 
233-234: 69, 1922, Basionym: Polypodium vel Aspidium inter- 
medium Muhl. ex Willd. in L., Sp, PI,, ed. 4, 5: 262, 1810. •= 
Dryopteris, 

Th, interrupta (Willd.) B.C.Stone, Micronesica, 2: 3. 1966, Basi- 
onym: Pteris interrupta Willd.. Phytogr., 1: 13, t. 10, f. 1. 
1794. Synonym: Aspidium pteroides Swartz, Schrad. Joum, Bot«, 
1800(2): 33. 1801, Micronesia (Yap, Palau, Saipan), Trop. Asia, 



1968 Reed, Index Thel^ypteridis 285 

N, Australia, Polynesia, Trop, Africa (Oubangui)o 

Thelypteris inrisa (Swartz) Proctor, Rhodora, 61: 306. (1959) I960; 
l.c, 63: 34. 196lj Morton, Contrib. U.S. Nat. Herb., 38(2): 6l. 
1967. Basionym: Aspidium in visum Swartz, Schrad, Journ. Bot., 
1800(2): 34. 1801. West Indies, Jaraaica„ 

Th, inrisa rar, aequatorialis (C.Chr.) Morton, Contrib, U.S. Nat, 
Herb,, 38(2): 61. 1967o Basionyra: Dryopteris oli/tophylla rar. 
aequatorialis CoChr., Dansk Vis. Selsk. Skr., VII, 10(2): 189. 
1913. Ecuador, Bolivia, PerUo 

Th, invisa var, kunzeana (Hook.) Morton, Contrib, U.S. Nat, Herb., 
38(2): 62, 1967, Basionym: Nephix)diuii kunzeanum Hook., Sp, 
Fil, , 4: 102. I862, Synonyms: Aspidium abruptua Kunze, Linnaea, 
9: 93. 1834, non Blume, 1828; Dryopteris oligophylla Maxon, 
Contrib. U.S. Nat. Herb., 10: 489. 1908, Trop. Amer., Peru. 

Th, invisa var, pallescens (C.Chr.) Morton, Contrib. U.S, Nat. 
Herb., 38(2): 62, 1967. Basionym: Dryopteris oligophylla var, 
pallescens C.Chr,, Dansk Vid, Selsk, Skr., VII, 10(2): 188, 
1913. Ecuador, 

Th, irayensis (Copel,) Reed, comb, nov« Basionym: Cyelosorus i ray- 
en sis Copel,. ftiilip, Journ. Sci., 81: 28, 1952. Philippine 
Isls. (Batan), 

Th, iridescens (v,A,v.R. ) Reed, comb, nov, Basionym: Dryopteris 
iride scans v.A.v.R,, Bull, Jard. Bot. Buit., II, No, 11: 11, 

1913. Sumatra. 

Tho jacobsii (Holtt.) Reed, comb, nov, Basionym: Cyclosorua .1a- 

cobsii H oltt,. Bluraoa, U(2): 530. 1962, Sarawak, 
Th. jamesonii (Hook,) Tryon, Rhodora, 69: 6, 1967. Basionym: 

Nephrodium .lamesonii Hook., Sp, Fil,, 4: 66, 1862, Ecuador - 

Peru, 
Th, japoni«a (Baker) Ching, Bull, Fan Mem. Inst. Biol,, Bot, 6: 

312. 1936. Basionym: Nephrodium .japonicum Baker, Ann, Bot,, 

5: 318, I89I, Japan, S, Korea, China, 
Th, Japoniea var, glabrata Ching, Bull, Fan Mem, Inst, Biol,, Bot, 

6: 312. 1936. S. Korea. 
Th. japonica var. musashiensis Hiyama, Journ. Jap. Bot., 26: 155o 

1951. Japan (Honshu) „ 
Th, japoniea var. typica; H.Ito, Nova Flora Jap., 1: 135. 1939. 

Japan, 
Th. Japonica var. typica forma viridescens H.Ito, Nova Flora Jap., 

1: 135. 1939. ( Dryopteris .japonica var, viridescens Makino, 

Gensyoku Yagai syokubutu Zuhu, 3i 225, 1933 (nom, subnud,), 

Japan. 
Th, Japonica var. typica forma vulgaris H.Ito, Nova Flora Jap,, 

1: 135. 1939. Japan. 
Th, Jerdonii (Ching) Reed, comb, nov. Basionym: Cyclosorus jer- 

donij- Ching, Bull. Fan Mem, Inst, Biol,, Bot, 8: 228. 1938. 

Himalaya, Sikkim, 
Th« Jiraenezii (Maxon et C.Chr.) Reed, comb, nov. Basionjrm: Dryo- 
pteris jimenezii Maxon et C.Chr., Amer. Fern Journ,, 4: 79. 

1914. Costa Rica. 



286 P H y T L G I A Vol. 17, no. !■ 

Thelypteris Juergenaii (iloaenat,) Reed, coa.b. noT, BaalonyiQ: 

Nqphrodlani JuerKcnsii hosenst,, Vid. oelsk. 3kr., Vll, 10: 256, 

f. /♦3d. 1913. brazil (Amazonas). 
Th. keyaserlana (ftosenat.) Heed, comb, nov, Baaionym: Di-yopteris 

keysaeriana Roaenst., Fedde ftepert., 10: 333. 191*-, ^3^ Guinea. 
Th. khasiensis Ghing, Bull. Fan Mem. Inst. Biol., Bot. 6: 284. 

1936. Aasaai. 
Th. klossli (Ridl.) Ghing, Bull. Fan Mem. Inst. Biol., Bot. 10: 

252, 1941. Basionym: Laatrea klosaii ftldl., Trans. Linn, Soc, 

II. Bot., 9: 257. 1916. New Guinea. 
Th. kotoensis (Hayata) K.Iwats., Ac^a Phytotax. Joobot., 21: 42, 

1964. Basionya: Dryopteris kotoen3i8 Hayata, Icon. PI. Foraios., 

5: 279. 1915. Synonym: Gyclosorus truncatue rar, kotoensis 

(Hayata) H.Ito, Bot, Mag. Tokyo, 51: 729. 1937. Taiwan (Kotsho 

I si., Botal Tobago), 
Th. kunthjLi (Desv. ) Morton, Gontrib. U.S. Nat. Herb., 38(2): 36, 

53. 1967. Basionym: Nephjodium kunthii Desv,, Mera. Soc, Linn, 

Paris, 6: 258, 1827, Synonyms: Dryopteris noi-malls G.Chr., 

Ark. f. Bot., 9(11): 31. 1910; Thelypteris normalis Moxley. 

1920. Mexico to Boliyia and Brazil, Veneeuela; acuthem U.S.; 

West Indies. 
Th . kunzeana (Hook.) Ghing, Bull. Fan Mem. Inst. Biol,, Bot, 10: 

252. 1941. Basionym: 'Jephrodium kunzeanuna Hook., Sp. Fil., 

4: 102. 1862. - Th. invisa ▼er. 
Th, kunzei (Fee) Reed, comb. nov. Basionya: Aspidiun kunzei Fee, 

10 Mem,, 37, t. 41, f. A-H. I865. oynonym: Dryopteris cochlagQrs 

G.Chr., Ind. Fil., 280. 1905. Mexico, 
Th. kusaiana (Hosok.) Reed, comb. nov. Basionym: Dryopteris kusa- 

ana Hosok., Trans, Nat. Hist. Soc. Formos., 26: 77. 1936. Caro- 
line I sis (Kusaie), 
Th, laete-strigosa (Clarke) K.Iwats, , Fl. Eastern Himalaya, 484. 

1966. Basionym: f?ephrodium glandulosua var. laete-strigosua 

Clarke, Trans. Linn. Soc, London, Ser. 2, Bot., 1(3): 532, 

pi. UCXIV, f. 2. 1300, India, Ceylon. 
Th. laevigata (Mett. ex Kuhn) Tryon, Rhodora, 69: 6. 1967. Basi- 
onym: rhegonteris laevigata Kett. ex Kuhn, Linnaea, 36: 112, 

I869. Peru, 
Th, laevis (Mett.) Reed, comb, nov, Basionym: Aspidium laeve Mett., 

Pheg, u. Asp., 104, n. 249. l'^58. Philippine Isls. (Luzon). 
Th, lakhiinpurensi s (Rosenst.) K.Iwats., Mem. Coll. Sci., Univ. 

Kyoto, Ser. E, 31(3): 19i*. 1965. Basionym: Dryopteris lakhiro - 

purensis P^senst,, Mem. Rijks Herb., No. 31: 7. 1917. Assam, 
Th, lanceola (Christ) Reed, comb, nov, Basionym: Dryopteris di- 

versiloba var, lanceola Christ, Philip. Journ, Sci., Bot. 2: 

200. 1907. Philippine Isls. 
Th, lanipes (C.Chr.) Alston, Joum. Wash. Acad, Sci., 48(7): 233o 

1958, Basionym: Dryopteris lanipes C.Chr., Smiths. Misc. Coll., 

52: 39i». 1909. Colombia, Guatemala, 
Th, lainitensis (Bedd.) Reed, comb, nov, Basionym: Nephr odium laru- 

tense Bedd,, Handb, Suppl,, 73. 1892. Perak, Borneo, Malaya. 



1968 Reed, Index Thelypteridis 28? 

IVielypteris latebrosa (Kunze ex Mett.) Reed, comb. noVo Basionym: 
Aspidium latebrosum Kunze ex Mett,, Pheg. u. Asp,, lOA, n, 248, 
1858, Java, Sumatra. 

Th, latifolia (Presl) Reed, comb, nor, Basionjnn: Nephrodium lati- 
follum Presl, Epim. Hot,, 45. 1849. Philippine Isls. (Luzon) o 

Th. latiloba Chang, Bull. Fan Mem. Inst, Biol., Bot. 6: 303. 1936. 
China (Kweichow). 

Th, latipinna (Hook, ex Bak, in Hook, et Bak.) K.Iwats., Acta. 
Phytotax. Geobot., 21(5-6): 166, 1965. Basionym: Nephrodium 
latipinna Hook, ex Bak. in Hook, et Bak., Syn. Fil,, 292. 1867, 
Jara, Hongkong, S. China, S to Malaysia, Assam, Ceylon, 

Th, lauterbachii (Brause) Reed, comb, nor, Basionyia: Dryopteris 
lauterbaehii Brause, Engl, Bot. Jahrb., 49: 18, 1912, New Guinea. 

Th, laxa (Franch. et Sav, ) Ching, Bull. Fan Mea, Inst, Biol,, Bot, 
6: 333. 1936. Basionym: Aspidium laxum Franch. et Sar,, Enum, 
PI, Jap,, 2: 237. 1876; l,c., p. 63I. 1879. Central and S, China, 
Japan, Korea, Taiwan, 

Th . laxa rar, dilatata (Koidz,) H.Ito ex Honda, Nom. PI, Jap., 520, 
1939« Basionym: Dryopteris laxa var. dilatata Koidz,, Acta 
Phytotax, Geobot. , 1: 28, 1932, (Japan), = Th, hattorii. 

Th, laxa var. typica; H,It6, Nova Flora Jap,, 1: I40, 1939. Japan. 

Th. laxa var, typica forma glabrescens H,It6, Nora Flora Jap,, 1: 
140, 1939. Japan, 

Th, laxa var, typica forma typica j H.Ito, Nova Flora Jap«, 1: 140, 
1939. Japan, 

Th, lepidopoda (C.Chr. ex Tard, et CGhr,) Reed, comb, nov. Basi- 
onym: Cyclosorus lepidopodua CChr, ex Tard, et C.Chr., Not, 
5yst,, (Paris), 7: 73. 1938 (" lepidopoda" ). Laos, Tonkin, Annam. 

Th, lepidula (Hieron,) Alston, Jounn. Wash. Acad. Sci,, 48(7): 233, 
1958. Basionym: Dryopteris lepidula Hieron,, Hedwigia, 46: 328, 
t. 4, f. 7. 1907. Colombia. 

Th. leprieurii (Hook.) Tryon, Rhodora, 69: 6, 1967. Basionym: 
Nephrodi'-m leprieurii Hook,, Sp, Fil,, 4: IO6, 1862, Trop. 
Amsr, , Guiana, Peru, Costa Rica, 

Th, leptocladia (Fee) Proctor, Bull, Inst, Jamaica, Sci, Ser,, 

No, 5: 61. 1953. Basionym: Goniopteris leptocladia Fee, 11 Hera,, 
60, t. 16, f, 1, 1866, Trop, Amer,, Jamaica, 

Th. leptogramraoides (Rosenst,) Reed, comb, nov, Basionyia: Dryo- 
pteris leptogrammoide s Rosenst., Fedde Repert,, 9: 63. 1910, 
Costa Ricfto 

"-Th, lothaea (Unger) Reed, conb, nov, Basionym: Aspidium lethaeum 
linger. Gen, et Sp. PI. Foss., 190. 1850, Miocene: Styria (Kain- 
berg). Aff. Th. mollis, Th. oreopteris (=Tho limbospenna) et 
Th, patens, 

Th. leucadenia (Copel,) Reed, coisb, nov, Basionym: Cyclosorus 
leucadenius Copel., Philip, Joum. Sci,, 81: 27. 1952, Luzon, 

Th, leucolepis (Presl) Ching, Bull. Fan Mem, Inst. Biol,, Bot. 6: 
345. 1936, Bftsionym: Lastrea leucolepis Presl, Epim, Bot., 39, 
1849. Philippine Isls., Polynesia, Tonga Isl., Fiji. 



288 PHYTOLOGIA Vol. 17, no. U 

Thelypterio leuconevTon (Pee) Schelpe, Jourri. 3. Afr. Bot., 31(i.): 
266. 1965. Baalonym: Ngphrodlura leuc o norron F^'e, 5'' Mem. Foug., 
306, t. 18, f . 3. 1052, Synonym: Nephrodiujai maurltianuxa Fee, 
5® Mem, Foug. , JOP,, 1^52. Natal, Mascareneo, f-ladagascar, 

Th, leucophlebia (Chflst) Refid, con±i. nov. Baaionyn: Aapidlum 

leucophlrbia Christ. Bull, Boisa., II, 4: 961. 1?0U, 3o3ta Rica. 

Th, leucothrlx (C.Chr.) Tryon, Rhodora, 69: 6. 1967. Baaionym: 
Dryopteris leucothi'ix C.Chr., Smiths. Misc. Coll., 52: 377, 
1909, Boliria. 

Th. leTingei (Clarke) Ching, Bull. Fan Mem. Inst. Biol., Bot. 6: 
273o 1936, Basionym: Qymnogranima aurita var^ lerin^ei Clarke, 
Trans. Linn. Soc, li. Bot., 1: 568. 1880. Synonya: Dryopteris 
purdoaii C.Chr., Bot. Gaz., 56: 335. 1913« Hiaalaya (Siikim), 
China (Yunnan, Szechwan, Shensi). 

Th, levyi (Foum,) Morton, Contrib, U.S. Nat. Herb., 38(2): 37. 
1967. Basionyra: Aapidiua levyi Foum., Bull. Soc. Bot. France, 
19: 255. 1872. Nicaragua. 

Th, ligulata (J. Smith ex Presl) Ching, Bull. Fan Mem. Inst, Biol., 
Bot, 10: 252. 1941. Basionym: Lastrea lipxlata J.Sndth ex Presl, 
Epim. Bot., 35. 1849. Synonym: Dryopteris luerssenii (Harr.) C. 
Chr., Ind. Fil., 276. 1905. Philippine Isls. 

Th, limaensis (Copel.) Reed, comb. noT, Basionym: Dryopteris liaa- 
ensis Copel., Univ. Calif. Publ. Bot., 19: 296. 1941. Peru. 

Th, limbata (Swartz) Proctor, Rhodora, 61: 306. (1959) I960. Basi- 
onym: Aspidium limbatum Gwartz, Schrad. Joum. Bot., 1800(2): 
35. 1801. Synonym: Amauropelta breutelii Kunze, Farnkr., 1: 86, 
109. 1343. West Indies, St. Kitts, Guadeloupe. 

111. limbosporma (Allioni) H.P.Fuchs, Acier, Fern Joum,, 48: 144. 
(195s) 1959c Basionym: Polypodlum limbo spenaim Allioni, Auct. 
Fl. Pedemont., 49. 1789 (Prior to Apr. 1). Synonym: Polypodiua 
oreopteris Shrh. in Willd„, Prodr., 292. 17S7. Europe, >fadeira, 
Japan, Pac. N. Amer, (Alaska - Wash.). 

Th, X lindheimeri (A.Br, ex C.Chr.) Wherry, Araer. Fern Joum., 54: 
145. 1964. Basionya: Dryopteris normalis var, lindheimeri A.Br, 
ex C.Chr,, Dansk. Vid. Selsk, Skr. , VII, 10: 182. 1913 (pro 
33m.), SS United States, 

Tho lindijgii (CChr.) Alston, Joum, Wash. Acad, Sci«, 48(7): 233. 
1958. Basionym: Dryopteris lindijdi C.Chr., Ind. Fil,, 275. 
1905. * Th. deflexa (Presl) Tryon. 

Th. lindmanii (C.Chr,) Reed, corab. nor. Basionym: Dryopteris lind- 
aanli C.Chr,, Vid, Selsk. Skr,, VII, 4: 281, f. 9. 1907 ( lind- 
mani ). S. Brazil, 

Tho lineaU (Blume) K.Ivrats., Mem. Coll. Sci., Ser. B, 31(1): 34. 
1964. Basionym: Aspidium lineatum Bluine, Qium, PI. Jar., 144. 
1828. Synonym: Dryopteris tenompokensls C.Ghr., Gardens Bull* 
Straits Settlements, 7: 248, 1934. Jara, Borneo, Malaya. 

Th. lingulata (C.Chr.) Morton, Contrib. U. S. Nat. Herb., 38(2): 
43. 1967. Basionym: Dryopteris lingulata C.Chr., Dansk Vid. 
Selsk. Skr., VII, Naturv. Afd., 10(2): 271. 1913. Costa Riea. 



1968 Reed, Index Thelypteridis 289 

Thelypteris linkiana (Presl) Tryon, Rhodora, 69: 6. 1967. BasionTmi 

Graromitls linkian a Presl, Tent. Pterid., 209. 1836; Gfeymnogramna 

polypodioides Link, Hort. Berol., 2: 50. 1833. Synonym: Gymnogram- 

ma diplazioides Desv., Mem. Soc. Linn, Paris, 6: 214. 1627. West 

Indies, Hispaniola, 
Th, linnaeana (Cooel.) Reed, comb. nor. Basionym: Laatrea linnae- 

ana Copel., Gen. Fil., 139. 1947. Synonym: Dryopteris sancta 

Kuntze, Rev. Gen. PI., 2: 813. 1891. West Indies, Gent. Amer, 
Th, lithcphylla (Copel.) Reed, comb, nov, Basionya: Dryopteris 

lithophylla Copel., Hiilip. Journ. Sci,, 12C: 57. 1917. Borneo. 
The liukiuensis (Christ in Mat sun, ) K.Iwats., Mem. Coll. Sci,, UniVo 

Kyoto, Ser. B, 31(3): 191o 1965. Basionym: Meniscium liukiuense 

Christ in Matsum. , Bot. Mag. Tokyo, 24: 240. 1910. Synonym: Aba- 

copteria triphylla var. siniplicifolia Chlng, Bull. Fan Mem. Inst. 

Biol., Bot. 8: 243. 1938; Abacopteria samp son! (Baker) Ching, 

Bull. Fan Mem. Inst, Biol,, Bot. 8: 244. 1938 (p.p.). I^yukyus, 

Okinawa; Taiwan, 
Th, lobangensis (C.Chr. ex C.Chr. et Holtt.) Reed, comb, nor, 

Basionym: Dryopteris lobangensis C.Chr. ex C.Chr. et Holtt., 

Card. Bull. Straits Settlements, 7: 245. 1934. Borneoo 
Th, lobata (Copel.) Reed, comb. nov. BasionjTn: Cyclosorus loba- 

tus Copel., Philip. Journ. Sci., 81; 33. 1952. Philippine Isls. 
Th. loheriana (Christ) Reed, comb. nor. B&sionym: Aspidium loheri- 

anum Christ, Bull. Boiss., 6: 191. 1898. Kiilippine Isls. (Luzon). 
Th. lomatosora (Copel.) Reed, comb, nov, Basionym: Dryopteris 

loraatosora Copel., Univ. Calif. Publ. Bot., 19: 298. 1941. Peru. 
Tho lonchodes (D.C.Eaton) Ching, Bull. Fan Mem. Inst, Biol,, Bot, 

10; 252. 1941. Basionym; Aspidiua lonchodes D.C.Eaton, Mera, 

Amer, Acad., n.s., 8: 210, 186o*. Cuba, 
Th, longlcaulis (Baker) Reed, comb, nov, Basionym: Nephrodiua 

longicau le Baker, Journ. Bot., 1881: 204. 1881, New Grenada, 
Th, longicuspis (Baker) Schelpe, Journ. S. Afr. Bot,, 3lU): 262. 

1965. Basionym: Nephrod ixun longlcuspe Baker, Journ. Linn. Soc., 

16: 202, 1877. Synonym: Nephrodiua zambesiacum Baker, Ann, Bot,, 

5: 318, 1391, Madagascar, Nyasaland. 
Th, longifoli* (Dcsy,) Tryon, Rhodora, 69: 7. 1967. Basionym: Me- 

niscium lon^ i folium Desv. . Mem, Soc. Linn. Paris, 6: 223. 1827. 

Synonyms: Dryopteris longifoli a (Fee) Hieron,, 1907, illegit.; 

Dryopteris desvauxli Maxon et torton. Bull. Torr, Bot, Club, 

65: 369. 1938. Brazil, 
Th, longifolia forma glandulosa (Maxon et torton) ttorton, Contrib, 

U.S. Nat. Herb,, 38(2): 52. 1967, Basionym: Dryopteris des- 

vauxii forma glandulosa Maxon et Morton, Bull, Torr, Bot, Club, 

65: 372. 1938. Brazils 
Th, longipes (Blume) Reed, comb, nov, Basionym: Aspidium longipes 

Blume, Enum. PI. Jav., 155, 1828, Java, Celebes. 
Tho longipetiolata (K.Iwats/) K.Iwats,, Mem, Coll, Sci,, Univ. Kyoto, 

Ser, B, 31(3): 194. 1965. Basionym: Abacopteris longipetiolata 

K.Iwats., Acta Phytotax. Geobot., 18: 11. 1959. Taiwan, 



290 P H y T L G I A Vol. 17, no. I4 

Thftlypterle longipilosa (Godiro) Reed, comb, nor, BaaionjrB: N»~ 
phrod ium lon/^ipilo .sum Sodiro, 3ert. Fl, Ecoad,, II: ^6, 1908, 
Ecuador, 

Tho longiasima (brack.) Reod, comb, nor, Baaionya: ItoxiloptTJa 
longiasima Brack,, ^pl. lixp., I6: 29, t. 5. Ii5k. Tahiti. 

Th. lorentzii (Hieron.) Abbiatti, Darvdniana, 13(2-A): 566. 196^. 
Basionym: Aspidiun lorent zii Hieron., Qigl. Bot. Jahrb., 22: 
368, (1896) 1897. Arcentina. 

Th, 3.ucida (Baker) Reed, comb. nov. Basionrra: Neph ro dia-B lucidm n 
Baker, Gard. Chron., n.s., 8: 456. 1877. Madag.iac--*r. 

Th, lugubrifonois (Rosenst.) Tryon, Hhodori, c';9: 7. 1967. BasionjTi: 
Dryopterls lugubriformls Rosenato, Fedde Rapert,, 7: 299. 1909. 
Peru. 

Th. lurida (Underw. et Maxcn ex Slosson) Proctor, Bull. Inst. Ja- 
maica, Scl. Ser. , No. 5: 6I. 1953. Basionym: Dryopteris lurida 
Underw. et Maxon ex Slosson, Bull. Torr. Bot. CLub, 40: 183, 
t. 3, f. 1» 1913. ( Nephrodium luridura Jerun. , nom, )o Jamaica, 

Th. lurida forma leucochaete (Slosson) Proctor, Bull. Inst, Jam., 
Sci. Ser., No, 5' 6I. 1953. Basionym: Dryopteris leucochaeta 
Slosson, Bull. Torr. Bot. Club, 40: 184, t. 3, f. 2. 1913. 
Jamaica, 

Th, luzonica (Christ) Reed, comb. nov. Basionym: DryoptexT.3 lu- 
zonica Christ, Philip. Joum. Sci. Bot., 2: 196. 1907, Philip- 
pine I sis, 

Th, macbridei (Maxon) Tryon, Rhodora, 69: 7. 1967, Basionym: Dryo- 
pteris macbridei Maxon. Joum. Wash. Acad. Sci„, 34: 25. 1944<, 
Peru, 

Th, macgregorii (Baker) Ching, Bull, Fan Mem, Inst. Biol,, Bot. 
10: 252, I94I0 Basionym: Nephrodium macgregorii Baiter, Ann, 
Bot,, 5: 320. 1892. Nexv Guinea, 

Th. macilenta E.P.St, John, Amer, Fern Journ. , 26: 50, 52, t, 5, 
1936. (Florida), = Dryopteris, 

Th, macradenia (Sodiro) Morton, Amer. Fern Joum., 51: 3^. 1961, 
Basionj'm: Nephrodivua macrodenium Sodiro, Rec, 47. 1383, Ecuador, 

Tho macroptera (Copel.) Reed, corpb. nov, Basionym: Dry opt e iris macro - 
ptera Copel., Univ. Calif. Publ. Bot., 12: 392. 1931. Tonga. 

Th, macropus (Mett,) Reed, comb, nov, Basionym: Aspidium Tnacro- 
pus Mett.. Fil. Lechl., 2: 20, 1859. Brazil. 

Th, macrorhizoma E.P.St, John, Amer. Fern Joum., 32: 146-147. 
(1942) 1943. Florida. 

Th, macrotis (Hook.) Tryon, Rhodora, 69: 7, 1967. Basionym: Nephro- 
dium macrotig Hook., Sp. Fil., 4: 86, 1862. Peru, 

Th, madagascariensis (Fee) Schelpe, Joum. S. Afr. Bot., 31(4): 267. 
1965. Basionym: Goniopteris madagascariensis Fee, Mem. Foug, 
5®: 251. 1852, Synonym: Goniopteris patens Fee, Mem, Foug. 5®: 
253. 1852; Gymno gramma unita Kunze, Linnaea, 18: 115. 1844; 
Goniopteris silvatica Pappe et Raws,, Syn. Fil., Afr. Austr., 
39, 1858 (nom. illegit.); Nephrodium costulare Baker, Joum. 
Linn. Soc. Lond., Bot. 16: 203. 1877; Dryopteris gladiata C.Chr., 
Ark. f. Bot., 14(19): 4, t. 1. 1916, Madagascar, S. Afr. (Natal). 



1968 Reed, Index Thelypteridis 291 

Thelypteris maemonensis (VJagner et Grether) B.C.Stone, Micronesica, 
2:3. 1966, Basionym: Cyclogorus maemonensis Wagner et Grether, 
Occ. Paper Bishop Mus., 19(2): 54, f. 5. 1948. Marianas (Guam). 
Th. magna (Copel.) Reed, comb. nov. Basionym: Cyclosorus aas^us 
Copel,, Philip. Joum. Sci., 81: 30. 1952. Philippine Isls. 
(Negros)o 
Th. magnifica (Copel.) Reed, comb. nov. Basionym: Dryopteris mag- 

nlfica Copel., Bishop Mus. Bull., 59: 11. 1929o Fiji. 
Th. malangaa (C.Chr.) Morton, Amer. Fern Joum., 53: 66.*1963. 
Basionym: Dryopteris malangae C.Chr., Svensk Vet. iUtad. Handl., 
Ill, 16(2): 21. 1931. Hispaniola, Jamaica, Haiti, Cuba. 
Th. malayensis (C.Chr.) Reed, comb. nov. Basionym: Dryopteris ma- 
layensis C.Chr., Vid. Selsk. Skr., VII, 10: 171. 1913. Synon^s: 
Aspidiujn /^landulosum Blume, Enum. PI. Jav., lAA. 1828, non Poly- 
Eodiuffl glandulosum Desv., 1811; Dryopteris excrescens Copel,, 
Univ. Calif. Publ. Bot., 14: 374. 1929. Perak, W. Malaysia, 
Sumatra, Philippine I sis. 
Th. malodora (Copel.) Reed, comb, nov, Basionym: Dryopteris maio- 
dora Copel., Philip. Joum. Sci., 60: 108, t. 13. 1936. SolSi^n 
Isls, 
Th. mapirensis (Rosenst.) Alston, Joum, Wash, Acad. Sci., 48(7): 
234. 1958, Synonym: DryopteriB mapirensis Rosenst,, Fedde Re- 
pert,, 6: 313. 1909. Colombia, Bolivia, 
Th. margaretae ( S.Brown in E. et F. Brovm) Ching, Bull. Fan Mem. 
Inst, Biol., Bot. 10: 252. 1941. Basionym: Dryopteris margaretae 
E.Brovfli in S. et F.Brown, Bernice P. Bishop Mus. Bull . 8o- 29 
pl. 3. 1931. Rapa. ^ ^ 7, 

Th. marginalis (L.) Nieuwl., Amer. Mdl. Mat., 1: 226. 1910. Basio- 
nym: Polypodium mar^inale L., 3p. PI., 1091. 1753. = Dryopteris, 
Til. marginalis forma davenportii (Floyd) L.B.Smith, Rhodora, 30: 17, 
1928. Basionjon: Nephrodium marginale forma davenportii Floyd, 
Rhodora, 4: 245. 19U2. = Dryopteris. 
Th, marginal is forma elegans (Robinson) Weatherby ex Hoffm,, Proc. 
Bost. Soc. Nat. Hist., 36: 197. 1922. Basionym: Aspidium mar -' 
ginale var. elegans J. Robinson, Bull. Essex Inst. (Salem, Mass.). 
8: No. 3, 151. 1875. = Dryopteris. 
Th. mascarensis (Baker) Reed, comb. nov. Basionym: Polypodium mas- 
c^rense Baker in Hook, et Baker, Syn. Fil., ed. 1, 455. 1868^ 
Madagascar. 
Tho matutumensis (Copel.) Reed, comb. nov. Basionym: Dryopteris 
matutumensis Copel., Philip. Joum. Sci., 40: 299, t. 3 1929 
Philippine Isls, (Mindanao). 
Th. raauensis (C.Chr, j Reed, comb. nov. Basionym: Polypodium 
clarkei Baker, Ann. Bot., 5: 457. 1891; Dryopteris raauensis 
C.Chr., Ind. Fil., 277. 1905. Hawaiian Isls. (Maui J^ 
Th, mauritiana (Fee) Reed, comb, nov, Basionym: Nephrodium mauri- 
tianum Fee, Gen, Fil., 308, 1850-1852, Ifedagascar, Masca?^^. 
Th. meeboldii (Rosenst.) Reed, comb. nov. Basionym: Dryop teris 
meeboldii Rosenst,, Fedde Repert., 12: 247. 1913. S IndiT, 



292 PHYTOLOGIA Vol. 17, no. U 

Thelypteris ■egacuspis (Baker) Reed, corah, nov. BaeionyrD: Poly- 
podi'x m mcKacuope Baker. Joum. EJot., 1H90: ,?66. IBVO. Tonkin, 

Th, ■e/?alocarpa (v.A.v.R. ) Ching, Bail, Pan Mem. Inst. Biol,, Bot. 
10: 252. 19U.. Basionym: Dryopteris me/^alocarpa v.A.v.R., Bull. 
Buit., Ill, 5: 199. 1922. Sumatra. 

Th . wefTalodus (Schkuhr) Proctor, Bull, Inst,. Jamaica, Sci, Ser., 
No. 5: 6l. 1953. Basionym: Polypodlua ae/>alodua Schkuhr, Krypt, 
Gew., 1: 2k, t. 19B, 1806, - Th. pennata. 

Th, jnegaphylla (Hett.) K.Iwate., Hem. Coll, Sci., Univ. Kyoto, 31(1): 
3^. 1964, Basionym: Aspidiua ae^aphyllun Mett., Ann, Lugd, Bat,, 
1: 233o I864. N. India, Ceylon, Malecia, Ins. CoiK)ra«, Wert Afr. 

Th, megaphylloides (Rosenst.) Reed, comb, nov, Basion^'ra: Dryo- 
pteris mep.aphylloide 3 Rosenst,, Fedde Repert,, 12: 174. 1913. 
New Guinea, 

Th, nelanochlaena (CChr.) Reed, comb, nov, Basionym: Dryopteris 
melanochlaena C.Chr. , Smiths. Muse, Coll., 52: 324. 1909. Guate- 
mala. 

Th. nelanophlebia (Copel,) Reed, comb, nov, Basionym: Dryopteria 
melanophlebia Copel., Philip, Journ. Sci., Bot. 6: 147. 1911, 
Philippine Isls. (Negros), 

Tho membranacea (Mett.) Tryon, Rhodora, 69: 7. 1967o Basionym: 

PheKOpteris membranacea Mett.. Fil. Lechl., 2: 22. 1859. Synonym: 
Nephrodiua lechlerl Hieron,, Sngl, Bot, Jahrb., 34: 4^*8. 1904. 
Peru, 

Th, nembranif era (C.Chr, in Bonap.) Reed, comb, nov, Basionym: Dryo- 
literis membranifera C.Chr, in Bonap,, Notes Ptertd,, 16: 170, t. 
2. 1925. Madagascar, 

Tho menisciicarpa (Blume) K.Iwats,, Acta Hiytotax, Geobot., 21(5-6): 
171, 1965c Basionym: Aspidium menisci! carpon Blune, Enum, Plo 
Jav,, 142, 1828« Synonyms: Qv'^closorus roenisciicarpus (falume) 
Holtt., Danak Bot. Ark., 25(2): 39. 1967; Phegopteria cordifolia 
v.A.v.R., Tiull. Jard. Bot. Buit., Ser. II, 11: 19, t. 5. 1913; 
Dryopteris mlrabilis Copel., Philip. Joum. Sci., 6C: 137, t, 
19. I9II; Dryopteris verrec^ilosa v.A.v.R., Bull. Jard, Bot. Buit., 
Ser. II, 11: 12. 1913; Polypodium holophyllum Baker, Joum. Bot., 
1888: 325. 1888, non P, holophyllum Baker, 1879= Malaya, Suma- 
tra, Borneo, Java, Philippine Isls. (Palawan). 

Th, meniscioides (Liebm, ) Reed, comb, nov, Basionym: Polypodiua 
meniscioides Liebm., Vid. Selsk. Skr., V, 1: 211. 1849. Synonym: 
Goniopteris rostrata Fee, 11 Mem., 64, t, 17, f. 3. 1866. Kexico- 
Panama , Guad eloupe , 

Th, aercurii (A.Braun ex Hieron,) Reed, comb. noVo Easionym: Dryo- 
pteris aercurii A.Braun ex Hieron,, Hedwigia, 46: 335, t. 5, f. 
9. 1907; Aspidiua mercurii A.Br., mcs., Christ I906, nomen. Mexi- 
co - Costa Rica - Ecuador. 

Th. merrillii (Christ) Reed, comb, nov, Basionym: Dryopteris aer- 
rillii Christ, Philip. Journ. Sci., Bot. 2: 201. 1907. Philippines. 

Th, mertensioides (C.Chr.) Reed, comb. nov. Basionym: Dryopteris 
mertensioides C.Chr., Vid. Selsk, Skr., VII, 4: 328, f. 50, 1907. 
Costa Rica, 



1968 Reed, Index Thelypteridis 293 

Thelypteris metcalfei (Baker) Reed, comb. nov. Basionyra; Poly - 
podium met calf ei Baker, Ann. Hot,, 5: 461. 1?^1. New Hebrides. 

The mesocarpa (Copel,) Reed, corrb. nov. Basionyrc: Dryopteris mfcso - 
carpa Gopel., Bishop Mus. Bull., 93: 9, t. 7A. 1932. Society Isls. 

Th, raetteniana Ching, Bull. Fan Mera, Inst. Biol., Bot, 10: 252. 

1941. Basionym: Nephrodium palustra Baker in Hook, et Bak,, Syn, 
Fil,, 270. 1827, non Thelypteris palustris Schott, 1834o Brazil. 

Th. mettenii (Copel,) Abbiatti, Darwiniana, 13(2-4): 566. 1964. 
Basionjrm: Lastrea mettenii Copel., Gen, Fil., 139 (nomo nov,). 
1947. Synonyms: Dryopteris palustris (Mett. ex Fee) Kuntze, Rev, 
Gen. PI., 2: 813. 1891; Aspidium palustre Mett. ex Fee, Crypt, 
Vase, Bresil, 2: 76. 1872-1873. Brazil. 

Th, mexiae (C.Chr, ex Copel.) Ching, Bull, Fan Mem. Inst. Biol,, 

Bot. 10: 252. 1941. Basionym: Dryopteris mexiae C.Chr. ex Copel., 
Univ. Calif, Publ, Bot., 17: 32, t. 6. 1932. Brazil. 

*Th, meyeri (Heer) Reed, comb, nov, Basionym: Aspidium meyeri Heer, 
Tertiarfl. der Helv., 1: 36, t. 11, f. 2. 1855. Tertiary, Mio- 
cene: Switzerland, Oeningen, Lausanne. Aff, Th, DoUis, 

Th. micans (Brause) Reed, comb, nov, Basionym: Dryopteris rolcans 
Brause, Fjigl. Bot. Jahrb,, 56: 98. 1920, New Guinea. 

Th. raicrobasis (Bak. in Hook, et Bak.) Tard,, Not. Syst., 14: 344. 
1952 (1953)| M«m„ IFAN, 28: 117, t. 20, f. 1. 1953o Basionym: 
Nephrodium microbasis Bak. in Hook, et Bak., Syn. Fil,, 496, 
1874. Synonym: Diyopteris adenochlamys C.Chr,, Fedde Repert,, 
9: 370. 1911 Africa (Senegal, Sudan, Guinea, Fr, Guinea, Da- 
homey, Ivory Coast, Ghana, Nigeria), 

Th, micr^carpa (v,A,v,R, ) Ching, Bull, Fan Mem, Inst. Biol,, Bote 
10: 252. 1941. Basionym: Dryopteris microcarpa v.A,v,R, , 
Bull, Buit., Ill, 2: 146. 1920. Sumatra. 

TTi, Joicroloncha (Christ) Reed, comb. noVo Basionym: Dryopteris 

microloncha Christ, F^ilip. Journ. Sci., 2: 202. 1907o Philippines. 

Th. microsora Reed, nom. nov. Basionym: Dryopteris microsora Copel., 
Bishop Mus. Bull., 59: 12. 1929 (homonym) , non Dryopteris micro- 
sora (Hook.) Kuntze, 1891, Fiji. 

Th, mil lei (C.Chr.) Reed, comb, nov, Basionym: Dryopteris millei 
CChr., Vid. Selsk. Skr., VII, 10: 138. 1913. Ecuador. 

Th, mindanaensis (Christ) Reed, comb, nov, Basionyiri: Dryopteris 
mindanaensis Christ, Philip. Journ, Sci,, Bot, 2: 194. 1907.. 
Philippine Isls (Fandanao). 

Th, rainensis Abbiatti, Darvdniana, 13(2-4): 563-564, f. 8, t. 6, 
1964,. Brazil, 

Th. minuscula (M^xon) Morton, Contrib, U.S. Nat. Herb,, 38(2): 43, 
1967. Basionym: Dryopteris minuscula Maxon, Kew Bull, Misc, 
Inf., 1932: 135o 1932, Colombia. 

Th, ndnutula Morton. Amer, Fern Journ,, 43: 173-174. 1953. Ecuador, 

Th, mixta (Rosenst,) Reed, comb, nov. Basionym: Dryopteris mixta 
Bosenst., Fedde Repert., 12: 172. 1913. New Guinea, 

Th. mollicella (Maxon) Ching, Bull. Fan Mem, Inst. Biol,, Bot. 10: 
252. 1941, Basionym: Dryopt eris mollicella Haxon, Proc, Biol. 
Soc, Wash., 36: 49. 1923. Dominica. 



29l PHYTOLOGIA Vol. 17, no. I; 

TtialyiBtfli'lB mclllcula (Kunze ex Link) Chinf,, Bull. Pan I-tem. Inst. 

Biol,, Bot. 10: 252. 19^1, Basionyin: Polypodlum nolllc'ilujo Kunze 

ex Link, Fil. 3?., 130. 1B41. Trop. Aiaer. 
Tho molllB (Mett.) Tryon, Rhodora, 69: 7. 1967. Basionym: Phegojgteriji 

inollis'Mett.. Ann, 5c^ . "Jat., V. Bot., 2: 2U2. 186/*, non Dryo- 

pteris mollia (Jacq.) HJ.eron., 1907. Synonyn: Oryopteria per- 

mollia Maxon et Morton, Bull, Torr, Bot. Club, 65: 3^2, 1938, - 

Th. arborescens (Hu«b. et Bonpl.) Morton. 
Th. adlliuscula (Kuhn) K.Iwats,, Fl. Eastern Himalaya, hSU. 1966, 

Basionym: Aspidiura molliusculua Kuhn, Bot, Zeit,, 26: 41. 1368. 

N. India. 
Th. monilifonnlB (Tagawa et K.Iwats.) K.Iwats., Mem. Coll, Sci,, 

Univ. Kyoto, Ser. B, 31(1): 36. I964. Basionym: DimorphopteriB 

moniliforadf Tagawa ot K.Iwats. ex K.Iwats,, Acta Phytotax, .ieo- 

bot., 19: e, 1961, Molucca Isls. 
Tho morobensis (Copel.) Reed, comb, nov, Baaionya: Dryopteris rnoro - 

bensis Copel., Univ. Calif. Publ. Bot., IS: 221. 1942. New Guinea. 
Th. raosenii (CChr,) Reed, comb. noVo Basionym: Dryopteria mosenii. 

C.Chr,, Vid. Selsk. Skr., \r[i, i,: 300, f. 27. 1907. Brazil. 
Th. motleyana (Christ) Holtt., Rev. Fl. Mai,, 2: 247, f. 140. (1954) 

]955o Basionym: Aspidium motleyaLnum Christ, Ann, Jard. Buit., 

II, 5: 105. 1905; Nephrodium n^otleyanum Hook, ex Bakep in Hook, 

et Baker, Sjnio Fil., 266. 1867 (nomen). (Mesoneuron). Malaya 

(Java, Sumatra), 
Th, multiauricolata (Copel.) Reed, comb, nov, Basionym: Drropteris 

multiauriculata Copel., Univ. Calif, Publ, Bot., 18: 221, 1942. 

New Guinea, 
Th, multifonnis (C.Chr,) Reed, comb, nov, Basionym: Drropteris 

multiformis C.Chr., Vid. Selsk. Skr., VII, 10: 154, f. 17. 1913. 

(Th. multiformis (C.Chr.) Murillo, Cat. Illus, Plantas de Cundi- 

naraarca, 2: 107 (nom. illegit.). 1966). 
Th, multifrons (C.Chr. in Bonapo) Ching, Bull, Fan Mem, Inst, Biol., 

Bot. 10: 252. 1941. Basionym: Dryopteris multifrons C.Chr. in 

Bonap., Notes Pfcerid., 16: 172, t. 2B. 1925. MadagascAr, 
Th, multijugis (Eak, in Hook, et Bak.) Reed, Comb, nov, Basionym: 

Nephrod ium multiiu/yum Bak. in Hook, et Bak., Syn, Fil., 291. 

1867. J^ay Peninsula. 
Th, multilineata (V/all, ex Hook.) Morton, Amer, Fern Joum,, 49: 

113o 1959. Basionym: Polypodlum multilineatuB V.'all, ex Hook., 

Sp, Fil., 5: 11.(1863) IS64, Synonyms: Abacopteris taultilineata 

Ching, 1938; Nephrodium moulmeinense Bedd. . Handl. 275. 1883; 

Dryopteris moulmeinensis (Redd.) CChr. , 1905. Malay Peninsul,, 

Selangcr, Ceylon, N, India - S, China, Sumatra, Philippines, 
Th, niultilineata var, malayensis Reed, nom, nov, ( Abacopteris aul- 

tilineata var, malayensis Holtt., Rev. Mai, Fl., 2: 297. 1954 

(tiom. illegit,) Jo Laminae tenuiores, pinnae latiores cum non 

densatis marginibus, vix dentatis, Malaysia 
Tho multiseta (Bak,) Ching, Bull. Fan Mem. Inst, Biol,, Bot, 6: 

347 (err. raultisectua ), 1936, Basionym: Nephrodium multisetum 

Bak,, Joum. Linn. Soc. , 22: 226-227, 1886, Borneo, Java, 



1968 Reed, Index Thelypteridis 295 

Thelypteris multisora (C.Chr. ex C.Chr, et Holtt.) Reed, comb, nov, 
Basionym: Dryopteris multi sora C.Chro ex C.Chr. et Koltt,, Gard, 
Bull, Straits Settlements, 7: 241. l^JU, BorneOc 

Th„ raunda (Rosenst.) Chinp, Bull. Fan Mem, Inst, Biol., Bet. 10: 

252. I94I0 Basionym: Dryopteris mainda Rosenst.. Med. Rijks Herb., 
No. 31: 5o 1917. New Guinea. 

Tho muricata (Brause) Reed, comb. nov. Basionym: Drropteris muri- 
cata Brause. Sngl. Bot. Jahrb^ , 56: IO6, 1920. New Guinea. 

Th, muscicola Proctor, Rhodora, 63: 33, 1961, West Indies, NeirLs, 

Th, nrutabilis (Brause) Reed, comb, nov, Basionym: Dryopteris muta- 
bilis Brause, 5iigl. Bot. Jahrb., 56: 97. 1920. New Guineao 

Tho rauzensis (Hieron,) Alston, Joum. V;asho Acaa, Sci,, 48(7): 233. 
1958. Basionym: Dnropteris muzensis Hieron., Hedwigia, 46: 331> 
t, k, f. 6. 1907. Colombia, Ecuador. 

Tho myrioscra (Copel.) Reed, comb, nov, Basionym: Dryopteris igyrio- 
80 ra Copel,, Riilip. Joum. Sci,, 60: 108, t. 14. 1936, Solomon 
Isla, 

Th, namburensis (Bedd.) Reed, comb, novo Basionym: Nephrodium nam - 
burense BedJ . . Handb, Suppl, 1892, p, 69. Assam, 

Tho navarrensis (Christ) Proctor, Bull. Inst, Jamaica, Sci, Ser,, 
No, 5: 61. 1953. Basionym: Aapidium navarrense Christ, Bull. 
Boiss., II. 7: 262. 1907. Trop. Amer., Costa Rica, Jamaica, Haiti. 

Th, neglecta (Brade et Rosenst.) Ching, Bull. Fan Mem, Inst, Biol,, 
Bot, 10: 253. 1941 o Basionym: Dryopteris neglecta Brade et Rosen- 
st., Bol. Mus. Nac, Rio Janeiro, 7: 142, t. 7. 1931c Brazils 

Tho nemoralis Ching, Bxill. Fan Mem, Inst, Biol., Bot. 6: 338, 1936. 
(China: Kiangsi, Anwbei), = Th, hattorii var, 

^0 nemora lis (Sodiro) Tryon, Rhodora, 69: 7. 1967. Basionym: Neph- 
rodiu m n emorale Sod., Crypt. Vase, Quit,, 267, 1893. (Ecuador), 
- Th.'sodiroi. 

Th, neo-auriculata (Ching) Ching, Bull, Fan Mem, Inst. Biol,, Bot. 
6: 281. 1936, Basionym: Dryopteris neo-auricul ata Ching, Bull, 
Fan Mem. Inst, Biol., Eot. 2: I96, t. 10. 1931." China (Yunnan), 

Th. nephrodioidea (Klotsch.) Proctor, Bull. Inst. Jamaica, Sci. 
Ser., No, 5: 61. 1953. Basionym: AspiAium nephrodioides Klotz- 
seh,, Linnaea, 20: 370. 1847, Synonym; Nephrodium guadala- 
pense Fee. 11*^ Mem, Foug, : 89, t. 24, f. 3. 1866, Trop, Amer. 
(Colombia, Jamaica). 

Th, nephrolepioides (C.Chr.) Reed, conb. nov, Basionym; Dryopteris 
nephrolepioides CChr., Brittonia, 2: 295, f. 1 c-d. 1937. New 
Guinea. 

Th, nervosa (Klotzsch.) Tryon, Rhodora, 69: 7. 1967. Basionym; 
Polypodium nervosum Klotzsch,, Linnaea, 30: 386, 1847, non Poly- 
podium nervoeum Boj., 1837, nom. nud. Guiana. 

Th, nesiotica (Maxon et Morton) Morton, Contrib, U.S. Nat, Herb,, 
38(2); 43. 1967, Basionym: Dryopteris nesiotica Haxon et Merton, 
Bull. Torr, Bot. Club, 65: 362, t. 12, 1938, Trinidad. 

Th, nevadensis (Baker) Clute, ex Morton, Amer. Fern Joum,, 48; 139o 
1958. Basionym: Nephrodium nevadense Baker, Ann, Bot., 5; 320. 
I89I, non Aspidium nevadense D.C.iiaton, Ferns N. Amer,, 1; 73, 



296 P H y T L G I A Vol. 17, no. U 

t. 10. 1878, non. illegit. Synonym: Dryopteris or»i^.«jitt C.Chr., 
C.Chr,, Ind. Kil., 281. 190^). Western United Statee. (Callf., 
Oregon). 

Thelypterj e niwiraguensia (Fourn.) Morton, Gontrib. U.3. Nat, Herb., 
38(2): 55. 196?. Basionym: Phef.optbria nicaraffjensia Fourti., 
Bull. Soc. Bet. France, 19: 252. 1872. Centr. Ara-er., Nicaragua, 

Tho nigrescentia (Jenin. ) Reed, comb, ncv, Basionym: Polypodiua nl- 
/treacentium Jonra. , Gard. Chron., Ill, 17: 100. 1895. Jamaica. 

Th, pipponica (Franch. et Sav. ) Ching, Bull. Fein f.'^m, Inst. Biol., 
Bot. 6: 309. 1936. Basionym: Aapidlum nipponiciun Frauich. et 3av., 
Enura. PI. Jap., 2: 2A2. 1867, p. 636. 1869. Cent, and W. China, 
Japan. 

Th, nipponica var. borealls (Hara) Hlyaraa in Yas«, 3: I30. 1937. 
Basicnym: Dryopteria nipponica var. borealia Hara, Bote Mag, 
Tokyo, 48: 695» 193A. Japan (Hokkaido, , Honshu, Yezo, Saghalien). 

Th, nipponica var. typica; H.Ito, Wova Flora Jap., 1: 133. 1929. 

Th, nltena (Desv.) Tryon, Rhodora, 69: 7. 1967. Basionym: Poly- 
podium nitena Deav., Mem. Soc. Linn. Paris, 6: 240. 1827. Andes, 
Peru- Ecuador. 

Th, nitidula (Preal) Reed, comb, noVo Basionym: Nephrodiua niti- 
dulum Presl, Epim. Hot., i+6. 1849. Synonyms: Nephrodium philip - 
pinense Baker, Ann. Bot., 5: 327. 1891; Nephrodium basU.are 
Presl, Epira. Bot., 258 (nomen). 1849; Dryopteria basilaris (Pre si) 
CChr,, Ind, Fil., 254. 1905 (nom. illegit.); Cyclosorua aer- 
ratus Copel., Riilip. Joum. Sci., 81: 36. 1952; Fern Fl. Philip,, 
2: 365-366. 1958. Philippine I sis. (Luzon, Bohol). 

Th, nockiana (Jenm,) Proctor, Bull„ Inst. Jamaica, Sci. Ser., No. 
61. 1953. Basionym; Nephrodium nockianum Jenm,, Joum. Bot., 
1886: 270, 1886. Jamaica, 

*Th, nodosa (Goepp, ) Reed, comb, nov. Basionym: Aspidites nodosus 
Goepp,, Syst, Fil. Foss., 372, t. 23, f. 1. I836, Synonyms: 
Pe copter! 3 nodosa Goepp., Ubersicht d. Foss. Flora Schlesiena, 
2: 215. 1844; Aspidites leptorrhachis Goepp., Syst, Fil. Foss., 
373, t. 23, f. 2. IS36; Aspidium nodosum (Goepp.) Ett., Die 
Famkr, der Jetztwelt., 198, I865. Upper Carboniferous: Sileaia 
(Waldenburg), Aff. Tho noveboracensis. 

Th, normalis (C.Chr.) Moxley, Bull. So. Calif. Acad. Sci., 19: 57. 
1920, Basionym: Dryopteris normalis C.Chr., Ark. f. Bot,, 9(11): 
31, 1910. - Th. kunthii. 

Th, normalis var. harperi (C.Chr.) Wherry, Araer. Fern Joxim., 54: 
146. 1964. Basionym: Dryopteris normalis var. harperi . C.Chr., . 
Dansk. Vid. Selsk. Skr. , VII, 10: 182. 1913. = Th. k^onthii. 

Th, notabilis (Brause) Ching, Bull. Fan Mem. Inst. Biol., Bet. 10: 
253, 1941. Basionym: Dryopteris notabilia Brause, Engl, Bot. 
Jahrb,, 56: 91. 1920, New 'iiinea, 

Th. novae-hibemiae Holtt., Dansk Bot, .Ark., 25(2): 50. 1907. New 
Ireland, New Hebrides. 

Th, noveboracensis (L.) Nieuwl., Amer. Midi. Nat., 1: 225. 1910. 
Basionym: Polypodium no vebora cense L., Sp. PI., 1091. 1753. 
Newfoundl.and to Minnesota, south to Georgia and Arkansas. 



1968 Reed, Index Thelypteridis 297 

Thelypteris noveboracensis forma excurrens Neidorf , Amer, Fern 
Journ., 39: 101. 1%9. New York. 

Til. noveboracensis forma fragrans (Peck) Reed, comb, nov, Basio- 
nym: Aspidiiun novaboracense var. fragrans Peck, New YoiSc State 
Mus, Kept., 28: SA. 1676, Synonym: Aspidium noveboracense 
forma suaveolens D.C.Eaton, Ferns N. Amer., 1: 50. 1878, New York. 

Th, novoguineensis (Brause) Reed, comb. nov. Basionym: Drropterls 
novoguinpensis Brause. Engl. Bot, Jahrb., A-9: 21. 1912„ New 
Guinea, 

Th. nuna (J.W.Moore) Ching, Bull. Fan Jiem. Inst, Biol., Bot, 10: 
253. 1941. Basionym: Dryopteris nuna J.W.Moore, Bemice P. Bis- 
hop Mus. Bull., 102: 7. 1933. Society Isls., Raiatea. 

Th, nymphalis (G.Forst,) Reed, comb, nov, Basionym: Folypodium 
nymphale G.Forst., Prodr., 81. 1786, New Zealand, Polynesia. 

Th. obliquata (Mctt,) Ching, Bull. Fan Mem, Inst, Biol., Bot, 10: 
253. 19A1. Basionjrm: Aspidium obliquatum Mett., Ann, Sci, Nat,, 
IV, 15: 75. 1861, New Caledonia, 

Th, obliterata (Swartz) Proctor, B'lll, Inst. Jamaica, Sci, Ser,, 

No, 5: 62. 1953. Basionym: Polypodiuja obliteratum Swai-tz, Prodr,, 
132. 1788, V/est Indies, Jamaica, Mexico. 

Th, oblonga (Brause) Reed, combo nov. Basionym: Dryopteris oblonga 
Brause, Engl. Bot. Jahrb,, 56: 109. 1920, New Guinea, 

The obstructa (Copel,) Reed, comb, nov, Basionym: Dryopteris ob- 
structa Copel., Univ. Calif. Publ, Bot., 12: 378. 1931. Rarotonga. 

Th. obtusata (v.A.v.R, ) Ching, Bull. Fan Men, Inst, Biol,, Bot. 10: 
253. 1941. Basionym: Dryopteris obtusata v.A,v,R. . Bull, Buit,,, 
IT, No, 28: 22. 1918, Sumatra. 

Th. obtusifolia (Rosen^t.) Reed, comb. noVc Basionym: Dryopteris 
obtusifolia Rosenst,, Fedde Repert., 10: 336, 1912^ New Guinea, 

Th, occulta T^ope) Reed, comb, nov, Basionym: Nephrodium occultum 
Hope, Journ, Bomb. Nat. Hist. Soc, 12: 627, t. 13. 1899c N.India. 

Th, ochropteroides (Baker) Proctor, Bullo Inst, Jamaica, Sci. Ser., 
No. 5: 62o 1953. Basionym: Nephrodium ochropteroides Baker, 
Ann, Bot., 5: 325. 1891. Jamaica, Panama - Colorabiao 

Th. ochthodes (Kunze) Ching, Bull. Fan Mem, Inst. Biol,, Bot, 6: 
300, 1936, Basionym: Aspidium ochthodes Kunze, Linnaea, 24: 282, 
1851. S. China (Yunnan), Tomkin, Japan, S, India, 

Th, odontosora (Bonap.) Ching, Bull, Fan Mem, Inst. Biol., Bot, 
10: 253. 1941; Alston, Bol, Soc. Broter., Ser. II, 30: 26, 1956, 
Basionym: Dryopteris odontosora Bonap,, Notes Pterid,, 4: 17. 
1917. Guiana, 

•*Th, oeningensis (A.Braun) Reed, comb. nov. Basionym: Goniopteris 
oeningensis A.Braun in Bruckmann, Jahresb. Ver. Vaterl, Naturk. 
kHrtt,, 6: 226, 1850, Synonyms: Lastrea oeningensis (A.Braun) 
Heer, Fl. Tert, Helv. , 1: 32, t. 6, f. 3, 1855j Dikstra, Fobs, 
Cat, 68: 3905. 1968; Phegopteris oeningensis (A,Braun) Ett,, Die 
Fsxnkr. der Jetztwelt., 196, 1865. Upper Miocene: Rumania, 
Baden (Oeningen), Hungary, 

Th, ogasawarensis (Nakai) H.It'o, ex Honda, Nom, PI. Jap,, 520, 1939. 
Basionym: Dryopteris ogasawarensis Nakai, Rigakkwai 26, Apr. 10, 
1928 (noraen); Bot. Mag, Tokyo, 43: 2. 1929. Bonin I si. 



298 P II Y T L G I A Vol. 17, no. h 

Thelypteris oligocArpa (Huob. et Bonpl. ex VVilld.) Chlng, Bull, 
Fan Mfim. Inst. Biol., 3ot. 10: 253. 19U.. Baoionym: Polyrxxiiuni 
oll^'^ocanjum Humb, et Bonpl. ex Willd. in L, , Sp. PI., ci. U, 5: 
201. 1810. West Indies (St. Kitts, Haiti, Jamaica, Cuba), M»?xico - 
Co?ta Ric<i, Trop. Ainer, 

Th. ollj^ocarpa var. crassistipitata (Hieron.) Reed, comb. nor. Baoi- 
onym: Aspidiiun oligocarruTi var. crassiatlpitatiun Hieron., ^jjl, 
Eot. Jahrb., 22: 367.(1?^6) 1^7. A.rsentina (Tucuraan). 

Th, oli^odictya (Baker) Reed, comb, nov, Basionym: Acrcstichum 
oli^odlctyon Baker, Joum. Linn. Soc, 24: 261, 1887, Synonym: 
Syn^ramma angusta Copel., Philip. Joum. Sci., 3C: 3A8. 1909. 
(Holtt'un, Blvunea, 11: 530. 1962). Sarawak, Brunei. 

Th. oligolepia (v.A.v.R. ) Ching, Bull. Fan Mem. Inst. Biol., Bot. 
10: 253. 19/*1. Basionym: Dryopteris oligolepia v. A.v.R,, Nora 
Guinea, 14: 17. 1924. New Guinea. 

Th. olij^ophlabia (Baker) Ching, Bull Fan He'll, Inst, Biol., Bot. 6: 
339. 193^. Baaionym: tJephrodium oli^ophlebiura Baker, Journ. Bot,, 
291. 1S75. = Th. torresiana, 

Th . oli^oph-lebj a var. eleg.ans (Koidz.) Ching, Bull. Fan Mem, Inst, 
Biol., Bot. 6: 339, 341. 1936. Basionym: Dryopteris elegans 
Koidz., Bot. Mag. Tokyo, 38: 108. 1924. = Tr., torresiana. 

Th . cli;;ophl'tbia var. eleganr forma lophaea (Ogata) Namegata et 
Kurata, Snum. Japo Pterid,, 344, 196l. ^Basionym: Drropfetris 
oljgophlebia var. lophaea Ogata, Icon, Fil, Jap,, 5: t, 227. 
1933. (Japan). 

Th . oligophlebia var. lasiocarpa (Hayata) H.Ito, Bot. Mag, Tokyo, 
52: 539, 1938; Nakai et Honda, 'slova Flora Ja?., 1: 144. 1939o 
Basionym: Dryopteris lasiocarpa Hayata, Joum. Coll. Sci., Univ. 
Tokyo, 30: 41'^. 1911. = Th. torresiana, 

Th. oligophlebia var. subtripinnata (Tagavra) H.Ito, Nova Flora Jap,, 
1: 144. 1939. Basionym: DrycTpteria elegans var. subtripinnata 
Tagav;?., Acta Phytotax. Geobot., 2: 193c 1933. = Th, viridifrons. 

Th. oligophylla (Maxon) Proctor, Bull. Inst. Jajnaica, Sci. Ser., 

No. 5: 62, 1953. Basionym: Dryopteris oligophylla M.axon, Contrib, 
U. S. Nat. Herb., 10: 489. 1908. = Th. invisa. 

Th. omeigensis (Ching) Reed, conb. nov, Basionym: Cycloscrus omei- 
gensis Ching, Bull. Fan Mem. Inst. Biol., Bot. Ser. II, 1: 239. 
1949. China (Szechuan). 

Th. cneiensis (Baker) Ching, Bull. Fan Mem. Inst. Biol,, Bot. 6: 
282. 1936, Basionym: Pclypodiuir omeiense Baker, Joum. Bot., 
1838: 229. 1888, Synonyms: Dryopteris izuensis Kodaaa in Mat- 
sa-n. , Icon. PI. Koisikav., II: 7, t. 88. 1914; Dryopteris pseudc- 
africana Makino et Ogata, Joum. Jap. Bot., 4: 140. 1927; Dryo- 
pteris leveiLlei Christ, Bull. Geogr. Bot. Man, 1909: Mem, XX, 
176, 1909, China (Szechuan, Kweichow), Taiwan, Japan (Kyushu, 
Honshu), 

Th, oosora (Baker) Reed, combo nov, Basionym: Nephrodivun oosorua 
Baker, Kew Bull., 1896: 41. 1896. Bom.eo. 

Th, ophiura (Copel.) Reed, comb, pov,, Univ. Call^, Publ, Bot,, 
18: 220, 1942 o New Guinea. 



1968 Reed, Index Thelj-pteridis 299 

Thelypteris opp.ca (Don) Reed, CGnb„ nov. Basionyni: Hemioniti s 

opaca Don, Prodr. Fl. Nepal., 13. 1825. Synonym: GyTiino/;raam& 

obtusata Bluine, Eniun. PI, Jav., U3. 1628. N. India, Jara. 
Tho opposita (Vahl) Ching, Bull. Fan Kern, Inst. Biol,, Bot. 10: 

253. 1941. Basionym: Polypodium oppoaitum Vahl, Ecloque Amer., 

3: 53. 1807o Synonym: Oochlainys revoirej Fee, Gen. Fil., 297. 

1852, Trop, Amer,, Colombia, West Indies. 
The oppositifolia (Hook,) Reed, comb, nov, Basionym: Polypodium 

oppositifcllum Hook. . 3p. Fil., 5: 8. 1863, Guinea, So Thome., 
Th, oppositiformie (C.Chr, in Bonap,) Ching, Bull, Fan Memo Inst, 

Biol., Bot. 10: 253. 1941. Basionym: Dryopteris oppositiformis 

C.Chr. in Bonap,, Motes Pterid,, 16: 173, t, 2. 1925. Madagascar, 
Tho opposltipinna (v,A,v,F.) Ching, Bull, Fan Mem, Inst, Biol,, 

Bot, 6: 268, 19360 Basionym: Phegopteris oppositipinna v,A,VoR,, 
• Bull, Jard, Bot, Buit., Ser. II, 16: 24. 1914.. 3, Himalayas to 

Malaya and Sumatra » 
Tho orbicularis (CChTo) Feed, comb, nov, Basionym: Dryopteris 

orbicularis C.Chr., Ind, Fil., 281. 1905. Synonyms: Aspidivmt 

nephrodioide s Hook., Sp. Fil., 4: 42, t. 235. 1862, non Klotzsch., 

1847; Aspidium hookeri Baker, Syn. Fil., 257. IS67, non VJall. 

1829 nee Klotsch, lft47. Malesia, 
Th. oregana (C.Chr.) H. St. John, Proc, Biol, Soc, vVash., 41: 192. 

1928. Basionym: Dryopteris oref:ana C.Chr., Ind, Filo, 286. 

1905. = Th, nevadensiSo 
Th, oreopteris (Ehrh. in V.'iUd.) Slosson ex Rydb,, Flora Eocky Mts., 

1043. 1917. Basionym: Polypodiom oreopteris Ehrh, (in Willde, 

Prodr,, 292, 178?) Beitro Naturk, Verw. Wiss,, 4: 44 . 1789o - 

Th, limbo sperma, 
Th . oi^opteris (var«) hesperia Slosson ex Rydb., Flora Rocky ^fts,, 

104i4. 1917. - Th. limbosperaao 
The organensis (Rosenst.) Ching, B\ill. Fan Mem„ Inst, Biol,, Boto 

10: 253. 1941. Basionym: Dryopteris organensis Rosenst,, Fedde 

Repert., 20: 91. 1924e Brazil. 
Tho omata (V/all. ex Bedd,) Ching, Bull. Fan Mem. Inst, Biol., Bot. 

6: 346, 3936c Basionym: Polypodium omatum Wal] . ex Bedd,, 

Ferns S. India, t. 171. I864. (= Th. setigera ?). India, Burma, 

Taiwan, 
Th, orthocaulis (K.Iwats.) Reed, comb, nov. Basionym: Cyeloeortta 

orthocaulis K.Iwats,, Araer, Fern Joum., 53: 135p t, 9. 1963. 

Tonga. 
Th. ovata R.P.St John in Small, Ferns S.E, States, 230, tab, 1938b 

Florida, Alabama, Georgia. 
Th. ovata (var,) harperi (C.Chr.) R.P.St, John in Small, Ferns S.E, 

States, 233. 1938, Basionyri: Dryopteris normalis var, harperi 

CChr,, Dansk. Vid, Selsk, Skr,, VII, 10: 182, 1913- = Th. kunthii. 
The oxyoura (Copel.) Reed, comb, nov, Ba3ion3rm: Dryopteris oxyoura 

Ccpel., Philip, Joum. 3ci., 60: 107, t. 12. 1936. Solomon Isls. 
Th, pachyrachis (Kunze ex Mett,) Ching, Bull. Fan l-Iem. Inst. Biol,, 

Bot. 10: 253. 1941; Proctor, Bull. Inst. Jamaica, Sci, Ser., 5: 

62. 1953. Basionym: Aspidium pachyrachis Kunze ex Mett., Phego u. 

Asp., 83, n. 199, 1858, Trop Amer,, Colombia, Veneeuela, 



300 P H Y T L G I A Vol. 17, no. !■ 

Thelypterie pachyrachia var, bogatenais (C.Chr.) Alston, Joum. 

Wash. Acad. Sci., h^.{l)'. 233. 1958. Ba8lor;/in: Dryopteria pachy- 
rachlq var. bogat'inBJa C.Chr., Y.g\. Dansk, Vid. Selak. 3kr., '^I, 
k: 306. 1907. Colombia. 

Th, pachyrachia var,, Jenmanl (3aker) Proctor, Bull, Inat. Januiica, 
Sci, Ser., No. 5: 63. 1953. Ba-sionjna: Nephrodiiun .jemaani Ba>er 
ex Jenm. , Joum, Bot,, 15: 263. 1377. Jamaica. 

Th, pacifica Reed, nora, nov. Basjonyii: Polypodiuir clarkei Baker, 
Ann. Bot., 5: ^57. 1891, non Pleocnemia clarkei Bedd,, 1876. 
Hawaiian Isls. 

Tho palauQnsis (Hosok.) Reed, conb. nov, Ba8ion3ni: Meniscium palau- 
ense Hoaok., Trana. Nat, Hist. Soc. Forrvoaa, 28: !/**». 1938. Palau 
Isl. 

The palawanenais Reed, nora, noVo Baaionyti: CycloBorua aubdL-nor- 
phus Copel., Philip. Joum. Sci., 81: 38. 1952. Philippine IpIs. 
(Palawan). 

Tho paleata (Copel.) Holtt., Rev. Fl. Mai., 2: 249, f. Ul. (1954) 
1955o Basionym: Dryopteris paleata Copel., Philip, Joum. Sci,, 
9C: 228, I9IA. Sumatra, Borneo, Malaya. 

*Tho paleoe.legans Reed, nom, nov, Basionym: Pecopteris art?ita 
Brongn., Hist. Veg, Poss., 1: 303, t. 108, f. 3. 1828, non Aa- 
pidiua arguta Kaulf., 1824, Synonyms: Phegopteria argut a (Brongn.) 
Ett., Die Famkr. der Jetztwelt, 195. 1865; Pecopteris elegan s 
Germar, Petref., 39, t, 15. 1844, non Pecopteris ele^ans Sternb., 
1821; Polypodltes elegan s (Germar) Goepp., ^st. Fil. FosSc, 344, 
t. 15, f. 10. I836; Unger, Gen. et Sp. PI. Foss., 168. 1850. Car- 
boniferous: France, Germany; Rhode Island. Aff. Th. decussata 
et Th, meniscioidea. 

Th, pallescens (Brause) Ching, Bxill. Fan Mem. Inst. Biol., Bot. 10: 
253. 1941. Basionyir: Dryopteris paLlescens Brause, Engl, Bot. 
Jahrb., 56: 88. 1920. New Guinea. 

Th, paludosa (Blurae) K.Iwats., Acta Phytotax. Geobot., 19: 11. 1961, 
Basionym: Aapidiura paludosura Blunie, Enuni. PI. Jav., 163. 1828, 
Synonyms: Polyjodium pyrrhorhachis Kunze, Linnaea, 24: 257. 1851; 
Polypodium distans D.Don, Prodr. Fl. Nepal., 2, 1825, non Kaulf. 
1824; Dryopteris brunnea WaLl. ex C.Chr., Ind. Fil., 255. 1905; 
Dryopteris somai Hayata, Icon. PI. Formos., 5: 287. 1915; Dryo- 
pteris chriatii Lev., Fl, Kouy-Tscheou, 491. 1915; Dryopteris 
hirtirachis C.Chr., Ind. Fil. Suppl., II: 15. 1917. Malaysia, 
N, India - Taiwan, S. China, Tonkin, Luzon, Polynesia, 

Tho palustria Schott, Gen, Fil. in Obs. sub t. 10, 1834. Synonym: 
Acrostichum thelypteris L,, Sp. PI., 2: 1071. 1753; Polypodium 
paluatre Salisb., Prodr., 403 c 1796 (nom, illegit,), Serai- 
coamopolitan (Europe, Algeria, Asia temp,, Hiraalaya, Cento China, 
S, India, Atl. N, Amer., Trop. Africa, New Zealand). (An Th. con- 
fluens (Thunb,) Morton, 1967). 

The palustri s fonM, afurc&ta Clute, Our Ferns, 151, 387. 1938. = Th, 
palustris var. pubescens. 

Th. palustris forma glabra H.Ito in Nakai et Honda, Nova Flora 
Jap,, 1: 127 (nota). 1939. 3urope, Manchuria, Korea. 



1968 Reed, Index Thelypteridis 301 

Thelypteris palustris var. haleana Fsmald, Rhodora, 31i 34. 1929. 
Florida to Louisiana, N to Pennsylvaniaj Bemiuda. 

Th, palustris forma pubescena (Lawson) dute, Our Ferns, 152, 387. 
1938, Basionya: Lastrea thelypteris (^ pubescens Lawson, Edinb, 
New Phil. Journ., n.s., 19: 277. I864. = Th. palustris var. 

Th, palustris var. pubescens (Lawson) Femald, Rhodora, 31: 34, t. 
160. 1929. Basionym: Lastrea thelypteris [' pubescens Lawson, 
Sdinb, New Phil. Journ., n.s,, 19: 277. 1864. Newfoundland to 
Manitoba, S to Georgia and Oklahoma j N.E.Asia (Karttschatka). 

Th, palustris vai". pubescens forna linearis (Farwell) Reed, comb, 
nov. Basionym: Fllix thelypteris var, linearis Farwell, Papers 
Mich. Acad. Sci., 2: 14. 1923. Michigan. 

Th, palustris var, pubescens forma pufferae (A.A.Eaton) Reed, comb, 
ncvo Basion3Tii: Nephrodium thelypteris forma pufferaa A.A,Eaton, 
Fern Bull,, 10: 78. 1902. E, Massachusetts, 

Tho palustrirj var, pubescens fonna suaveolens (Clute) Reed, comba 
nor, Basionj'Tn: Nephrodiura thelyp teris forma su aveolens Clute, 
Fern Bull., 18: 87. 1910. Nora Sc'otia, N«w Haiqjshire. 

Th . palustris forma puffera« (A.A.Iiiaton) L.B.Smith, Hhodora, 30: 
16, 1928, Basionym: Nephrodium thelypteris forma pufferae 
A.A.Saton, Fern Bull., 10: 78, 1902, = Th, palustris var, pubes- 
cens forma. 

Tho palustris var, squamigera (Schlecht.) Weatherby in Johnston, 
Contrib. Gray Herb,, 73: 40. 1924 J Tard., Mem. IFAN, 28: 119, 
t. 20, f , 7-3. 1953. Basionym: Aspidium thelypteris var, sgua- 
mi<;erum Schlecht., Adumbr., 23, t. 11. 1825.. - Th. confluens, 

Th, palustris forma suaveolens (Clute) Femald, Hhodora, 23: l65o 
1921. Basionym: Nephrodium thelypteris forma suaveolens Clute, 
Fern Bull., 18: 87. 1910, «= Th, palustris var, pubescens forma, 

Th, palustris Schott, var. typica; Femald, Rhodora, 31: 33. 1929, 

Th, panamensis (Presl) E.P.St. John, Amer. Fern Journ,, 26: 44, 
1936, Basionym: Nephrodium panamense Presl, Rel. Haenk,, 1: 
35. 1825. (Mexico-Panama, Cuba, Jamaica). ■= Th. resinifera, 

Th, papilio (Hope) K.Iwats,, Mem, Coll. Sci,, Univ. Kyoto, Ser. B, 
31(3): 175, 1965, Basionym: Nephrodium papilio Hope, Joumo 
Bombay Nat, Hist, Soc, 12: 625, t, 12. 1399. Himalaya, Sikkira, 
N, India, Assam, Ceylon, Taiwan, 

Tho papyracea (Bedd.) Reed, corab, nov, Basionym: Wephrx>diuai papy- 
raceum Bedd,, Handb, Suppl,, 69. 1892« N, India, 

Th, paraphysata (Copel,) Reed, comb, nov, Basionym: Diropteris 
paraphysata Copel,, Philip, Journ, Sci., Hot. 6: 74. 1911o New 
Guinea, 

Tho parasitica (L.) Tard, in Tard. et C.Chr,, Not. Syst,, Paris, 
7: 75. 1938; Fosberg, Occ, Pap, Bish. Mus., 23(2): 30, 1962; 
K.Iwats,, Journ, Jap. Bot,, 33: 315. 1963. Basionym: Polypodiua 
parasiticum L, . Sp, PI,, 2: 1090, 1753. Synonym: Dryopteris 
mollis var, subjclabra Hosok,, Trans, Nat. Hist, Soc, Foraos,, 
26: 73, 1936, Southeast Asia, China (Yunnan, Kwangtung), S, 
Japan, Taiwan, Caroline Isls., New Zealand, trop. and subtrop, 
Atlo Isls, 



302 P H Y T L G I A Vol. 17, no. I4 

'flielypteris paraflitica forma boninensla (H.Ito) Re'.-d, comb. noT. 

Basionyw: Cycloaorua parasiticus fonnR bonlnenaia H.Itb, Bot. 

Mag. Tokyo, 51 : 727. 1937. Bonin Isis. 
Th. parasitica var, formosana (Chiny) Reed, conb. noy. Basionyrax 

Cyclosorus parasiticus var. forfaosonus Ching, Bull. Fan Mem, 

Inst, Biol., fiot. 8: 205. 1933. Taiwan. 
Th. parasitica forma latiloba (M.Ito) Reed, comb, nov,, Basicnyn: 

Cycle so rug parasiticus forma latilobus H.Ito, Bot, M^g. Tokyo, 

51: 727. 1937. Bonin Isle. 
The parasitical forma pilosissima (H.Itc) Reed, comb. noTo Basio- 

nyra: Cyclosorua parasiticae fonna pilosissiauB H.Ito, bot. Kag. 

Tokyo, 51: 727. 1937 » Japan (f^jrukyus). 
Th, parasitica forma pubglabra (Hosok.) Reed, comb. nov. Basionya: 

Dryopteri s mollis var, subglabra Hosok., Traxis, Nat, Hist, Soc, 

FomoSo, 26: 78, 1936, Japan (Sikoku, Kyushu, Ryukyus), 
Th, parathelypteris (Christ) Ching, Bull, Fan Mom, Inst, Biol,, 

Bot. 6: 3IU. 1936, Basionym: Aspidium parathelypteris Christ, 

B-oU, Soc, Bot. France, 52 (Men. Ij: 36. 1905. China, 
Ttia parlpinnata (C«pelo) Reed, comb, nov, Basionym: Dryopteris 

paripinnata Copel., Univ. Calif, Publ. Bot., 18: 2:^0. 19^« 

N«w Guinea, 
Th, patens (Sxartz) Small, Ferns S.E. States, 243, 475, tab. 1933, 

Basionym: Polypodium patens Swartz, Prodr,, 133. 1788, West 

Indies, Mexico to Argentina and Paraguay, Galapagos Isls, 
The patens var, dependens (CChr,) Proctor, Bull, Inst. Jamaica, 

3ci, Ser,, No. 5: 63.. 1953. Basionym: Dryopteris patens var. 

dependens CChr., Dansk Vid. Selsk. Skr., 711, 10: 178, 1913. 

Jamaica, 
Th, pauciflora (Hook.) Reed, comb, nov, Basionyra: Meniscium pauci- 

florua Hook., Sp. Fil., 5: I64. 1964. ( Menisorus Alston, 1956). 

Trop, West Africa, 
Th, paucinervata (C.Chr.) Reed, comb, nov, Basionyn: Dryopteris 

pauclnervata C.Chr., Ind. Fil., 283. 1905. Synonym: Pclypodiun 

oli^ophlebium Baker in Hook, et Baker, Syn. Fil,, 506, lL7u, non 

P, oligophlebiua Kunze, 1350, Peru, 
Th, paucipinnata (,Donn,anith) Reed, comb, ncv, Basionym; Uephr»~ 

dium fendleri var, paucipinnatura Donn, Smith, Bot. Gaz., 12: 134. 

1837, Guatemala. Synonym: Dryopteris donnell-amithii Maxon, 

Contrib. U.S. Nat, Herb., 13: 19. 1909. 
Th. pavcniana (Klotzsch) Tryon, Rhodora, 69: 7. (Mai*. 31). 1967; 

Crabbe, Brit. Fern Gaz., 9(8): 3I8, 1967. Basionym: Polypodium 

pavonianum Klotzsch, Linnaea, 20: 386, 1847. Peru, 
Th. pectiniformis (CChr.) Ching, Bull. Fan Mem. Inst. Biol., Bot„ 

10: 253. 1941. Basionyn: Dryopteris pectinifomis C.Chr., Gard, 

Bull. Straits Settlements, 4: 379. 1929, Malaya, Perak. 
Th. pectinifomis var, eglandulosa Reed, now. nov. Piachis et costae 

eglandulosae. Malaya. (Holtt., ReVo Fl, Mai., 2: 254. (1954) 

1955, nom. illegit.), 
Th. pectinifomis var. hirsuta Reed, nom. nov. Pinnae subtus Mr- 

sutae. Malaya, Padang. (Holtt., Pev, Fl, Mai,, 2: 254. (1954) 

1^55, nom. illegit,). 



1968 Reed, Index T^elypteridis 303 

Thelypteris peltata (v.A.v„R. ) Reed, comb. nov. Basionvm: Dryo- 
pteris peltata v.A.v.Ro, Bull. Jard, Hot. Suit., II, No, 16: 12. 
1914« Sumatra. 

The peltociilarays (C.Chr.) Heed, coirb, nov, Basionym: Dryopteris 

peltochlaraya C.Chr., Dansk Bot, Ark., 9: 65. 1937. Sumatra, Java, 
Malaya, 

Th, penanglana (Hook.) Reed, comb, nor, Basionym: Polypodium penan - 
gianum Hook,, Sp. Fil,, 5s 13 ( panangianum ). 1363, China, No 
India, Penaiig. 

Th, pennata (Poir. in Laia. ) Morton, Gontrib. U.S. Nato Herb,, 38(2): 
64, 1967, Basionym: Polypodiuia pennatuia Poir, in Lam,, Encycl, 
Meth, , 5i 535. 1304, Synonyns: Polyp odiua aegalodus Gchkiihr, Kr, 
Gew., 1: 24, t. 19b. 1806j Thel.-'.TDteriB negalodus Proctor, 1953. 
Trop. Amero 

Tho pennigera (G.Forsto) Allan, Fl, jN, Zeal,, 51. 19^1, Basionym: 
Polypodium pennip.erum G.Porst. f., Prodr., 02, 1786, New Zea- 
land, Polynesia, Philippine Isls, 

Th, pentaphylla (Roaenst.) Reed, comb, novg Basionym: Dr:/^opteris 
pentaphylla Rcpenst., Fedde Repert,, 12: 529. 1913. New Guinea. 

Tho perakensis (B«ddo) Reed, comb. noTo Basionym: Aspidium pera- 
kense Bedd., Journ. Bob., 1888: ko 1838, Perak. 

Tho perglandulifera (v.A.v-.R, ) Reed, comb, nov, Basionym: Dryo- 
pteris perglandulifera v.A.v„R., Bull. Buit., Ill, 2: 150. 1920. 
Sumatra, 

Tho peripae (Sodiro) Reed, comb, nov. Basionyra: Nephrodium peripae 
Sodiro, Rec, 52. 1883; Crypt. Vase, (^t., 265. 1893. Ecuador, 

Th, perpilifera (v.A.v.R, ) Reed, comb. nov. Basionym: Diropteria 
perpillfera v.A.v.R,, Bull. Jard, Bot, Buit,, II, No, 11: 12, 
1913. New Guinea, 

Th, perpubescens (Alston) Reed, comb, nov, Basionym: Dryopteris 
perpubescens Alston, Joum. Bot., 78: 227, 1940 (Oct.); Nova 
Guinea, Ser, 2, 4: 111, t, 8. 1940 (Dec,), New G-oinea. 

Th, perstrigosa (J-Iaxon) Ching, Bull, Fan Mem, Inst. Biol., Bot. 
10: 253. 1941 o Basion^nn: Dryopteris perstrigosa M sixon. Kew 
Bull., 1932: 135. 1932. Colombia. 

Tho peiTiviana (Rosenst,) Tryon, Rhodora., 69: 7. 1967, Basionyra: 
Dryopteris peruviana Rosenst., Fedde Repert,, 7: 298. 1909o Peru. 

Tho petelotii Ching, Bull. Fan Mem, Inst. Biol., Bot. 6: 326, 
1936. Tonkin o 

Tho petrophila (Copel.) Reed, comb, nov, Basionym: Dryopteris 
petrophila Copel., Univ. Calif, Publ, Bot,, 10: 220. 1942, 
New Guinea, 

Tho phacelothrix (CChr, et Rosenst, ex Rosenst.) Tryon, Rhodora, 
69: 7. 1967. Basionyra: Dryopteris phacelothrix CChr, et Rosen- 
st, ex Rosenst,, Fedde Repert., H: 56. 1912, Bo3J.via. 

Th, phegopteris (L.) Slosson ex Bydb,, Flora Rocky Mts., 1043o 
1917. Basionym: Polypodium phegopteris 1,^, 3p, PI,, 2: 1089, 
1753. Synonym: niegopteris polypodioides P^e, Gen. Fil., 243. 
I85O-IS52, Newfoundland to Alaska, S to Pennsylvania, in ats, 
to W. N.C. and E. Tenn,, west to Washington; Aleutians; Green- 
land; Iceland, Siberia; N and NW China; N. India; Asia Minor; 
northern Europe. 



30U PHYTOLOGIA Vol. 17, no. h 

Thelypteris phillppina (Presl) Chlng, Bull, Fan Mem. Inst. Biol., 

Bot. 10: 253. 1941. Basionym: Physematlujii phlllpplnwun Presl, 
Epia. Bot., 3U, 18A9. Philippine Isls. 
Th, physeraatioidea (Kuhn et Christ ex Kmig in Urban) Morton, Aaer. 

Fern Joum,, UJi 17/^. 1953o Basionym: Aspidiun phyBematioides 

Kuhn et Christ ex Krug in Urban, Engl, Bot. Jahrb., 24: 115. 

1897c Hispaniola. 
Th. piedrensis (C.Chr.) Morton, Amer. Fern Joum., 53: 69. 1963. 

Basionyn: Dryopteris piedrensis C.Chr., Sndthe. Misc. Coll., 

52: 372, 1909. Cuba, Puerto Rico. 
Tho pilosa (Martens & Gal.) Crawford, Amer, Fern Joum,, /J.: 16, 

t. 3a, I95I0 Basionyn: Qymnogramna pilosa Martens et Gal., 

Mem. Acad. Brux., 15: 27, t. U, f. 1. 1842. Mexico-Ouateaala. 
Th, pilosa rar, aLlabamensis Crawford, Aner, Pern Joum., 41: 19- 

20, t, 3b. 1951. Alabama; Mexico. 
Th, pilosa Tar, najor (Foum,) Crawford, Amer, Fern Joum,, 41: 

19> t, 4. 1951. Basionyn: Qyimograania pilosa *ar, major Foum,, 

Mex, PI., 1:.73» 1872; GymnoKraama prooirrens F^e, Men. Foug, 

8: 78, 1857; G, tctta Tar, procvirrens Baker in Hook, et Baker, 

1874 (illegit,); Dryopteris pilosa rar. procurrens CChr,, 1913 

( ille git . ) , Mexi co-Guat emala , 
Th, pilosissina Morton, Fieldiana, 28: 11-12. 1951. Venezuela. 
Th, pilosiuscula (Zippel ex Racib.) Reed, comb, nov, Basionyn: 

Nephrodium pilosiusculun Zippel ex Racib,, Pterid. Buit,, 189. 

1898. Java, 
Th, piloso-hispida (Hook.) Alston, Joum, Wash, Acad. Sci., 48(7): 

233. 1958. Basionym: Nephrodium pilo so-hi spidun Hook,, Sp, 

Fil., 4: 105c I862. Mexico - Bolivia, Colonbia, 
Th, piloso-squajuata (v.A.y.R, ) Reed, coreb, nov, Basionyn: Dryo- 

pterie piloso-squaaata v.A.v.R. , Bull. Dept. Agric. Ind, Neerl., 

21: 4. 1908. New Guinea, 
Th, pilosula (Mett,) Tryon, Rhodora, 69: 7. 1967. Basionym: A5- 

pidiun pilosulun Mett., Fil, Hort. Bot, Lips., I30. 1856; As- 

pidiun lasiesthes Mett,, I856, non Aspidiun lasiethes Kunze, 

1850 (non. nud,); non Aspidiun pilosulun Wall,, 1829 (non. 

nud, ) , Ifexico-Peruc 
Th, pinnata (Copel.) Ching, Bull, Fan Men. Inst. Biol., Bot. 10: 

253. 1941. Basionyn: Dryopteris pinnata Copel., Univ. Calif c 

Publ. Bot., 14: 373. 1929. Sumatra. 
Th, pittieri (C.Chr.) Reed, conb. nor, Basionym: Dryopteris pit- 

tieri C.Chr., Smiths. Misc. Coll., 52: 393. 1909. Colombia. 
Th, pittsfordensis (Slosson) Victorin, Fil, Quebec, 2: 51. 1933. 

Basionyn: Dryopteris pittsfordensis Slosson, Rhodora, 6: 75. 

1904. (Vermont, Quebec), - Dryopteris, 
Th, platensis Abbiatti, Darwiniana, 13(2-4): 553, f. 5, t. 3. 

19 64 , Argent ina , 
Th. platyptera (Copel.) Reed, comb, nov, Basionym: Dryopteris 

platyptera Copel., Univ. Calif, Publ. Bot,, 18: 219. 1942. 

New Guinea. 



1968 Reed, Index Thelypteridis 305 

ThcljTJteris plunosa (C.Chr.) Reed, comb. noT, Basionym: Dryo- 
pteris pluaosa C.Chr,, Dansk. Bot. Ark., 9(3): 65-66. 1937. 
Borneo. 

Th. plurifolia (v.A.v.R. ) Reed, conb, nor, Basionyn: Dryopteri g 
plu rifolia v.A.v.R., Bull, Buit., Ill, 5: 201. 1922. Sumatra. 

TTi. poecilophlebia (Hook.) Reed, comb, nov, Basionym; Polypodium 
poecilophleblua Hook., Sp. Fil., 5: 14. 1863, Australia. 

Th, poiteana (Bory) Proctor, Bull. Inst. Jamaica, Sci. Ser,, No, 
5: 63. 1953. Basionym: Lastrea poiteana Bory, Diet. Class,, 
9: 233. 1826. Synonym: Polypodium crenatum Swartz, Prodr,, 132, 
1788, non Forsk,, 1775. Trop. Araer,, Jamaica, Galapagos Isls. 

Th. polycarpa (Blune) K.Iwats., Mem. Coll, Sci., Univ. Kyoto, Ser. 
B, 31(1): 32. 1964. Basionym: Aspidium polycarpon Blume, Enua, 
PI. Jav. , 156. 1828, Synonym: Sphaero Stephano s as plenioides J.Sm, , 
in Hook, et Bauer, Gen, Fil,, 21. 1839. l^alaysia, Thailand Pen, 

Th, polyotis (CChr. ex KJellb. et C.Chr.) Reed, comb, nov, Basio- 
nyro: Dryopteris polyotis C.Chr. ex Kjellb, et C.Chr., Engl, Bot, 
Jahrb., 66: 46. 1933. Celebes. 

Th. polyphlebia (C.Chr.) Morton, Amer, Fern Joum., 51: 38. 1961. 
Basionym: Dryopteris polyphlebia C.Chr,, Vid. Selsk, Skr,, VII, 
10: 161, f . 19. 1913. Costa Rica - Andes of Quito, Ecuador. 

Th, polyphylla (Copel.) Reed, comb, nov, Basionym: Dryopteris 
polyphylla Copel,, Univ, Calif. Publ. Bot., 19: 288, t. 37. 
1941, Mexico, 

Th, polypodioides (Raddi) Reed, comb, nov, Basionym: Ceterach 
polypodioides Raddi, Opusc, Sci. Bol,, 3: 284. 1819 j Fl, Bras,, 
1: 10, t. 22. 1825. Brazil. 

Th, polyptera (Copel.) Reed, comb, nov, Basionym: Cyclosorus poly- 
pterus Copel., Philip. Joum. Sci,, 84: I6I, 1955. Philippine 
Isls (Negros). 

Th. ponapeana (Hosak.) Reed, comb, nov, Basionym: Phegopteris pona- 
peana Hosak., Trans. Nat. Hist. Soc, Formosa, 26: 233. 1936. 
Caroline Isls (Ponape). 

Th. porphyricola (Copel.) Ching, Bull. Fan Mem. Inst, Biol., Bot, 
6: 237. 1936. Basionym: Dryopteris porphyricola Copel., Hiilip, 
Joum, Sci,, 7C: 60. 1912, Borneo, Sarawak, Malays. 

Th. porphyrophlebia (Christ) Reed, comb, nov, Basionym: Aspidium 
porphyrophlebium Christ, Bull, Acad, Geogr, Bot. Mans, 1904: 
117. 1904. China, 

Th, pozoi (Lagasca) Morton, Bull. Soc. Bot. France, IO6: 234. 1959, 
Basionym: Hemionitis pozoi Lagasca, Nov. Gen. et Sp., 33. I8I6, 
Synonyms: Polypodium tottum Willd. in L. , Sp, PI,, ed, IV, 5: 
201. 1810, non Thunb., 1800j Acix>stich um pilosiusculum Wikstr, , 
Kgl. Vet. Acad. Handl., 1825: 439. 1826:' Polypodium af ricanun 
Desv., Mem. Soc. Linn. Paris, 6: 239. 1326; Polypodium eliasii 
Sennen et Pau, Bull. Soc. Geogr. Bot. Mans, 1910: 94. 1910. Spain, 
Madeira, Azores, S. Africa, N. India-China, Korea, Japan, Malesia. 

Th. pozoi subsp. hiraalaica (Ching) Morton, Amer. Fern Joum., 56(4): 
179. 1966. Basionym: Leptogramraa himalaica Ching, Sinensia, 7: 
100. 1936. India, Himalaya. 



306 P H Y T L I A Vol. 17, no. U 

Thelypteria pozoi subap, molliaslna (Kunze) Morton, Aaesr. Pern 

Joum., 56(/*): 178. 1966, Basionym: Qyrnno^ raaraa tott.ft yar. nol - 
llsslma Kunze, Linnaea, 2/*: 2A9. 1851; G. mo'lllBsi-na Fischer tx 
Kunze, Linnaea, 23: 255, 310. 1850 (no.-a. nud.). N. India, Oy- 
lon, S, China, S, Korea, Jaoan, F^yukyus. 

Th, prentlcei (Carr. in Seem.) Alston, /jner. Fern Joum., il*5: 120, 
1955. Basionym: Laat.rea prenticei Carr. in Seera. , Fl. Vit,, 
359. 1873. Fiji, Samoa, 

Th, prisraatica (Desv. ) Schelpe, Bol. Sec, Broter., Ser. 2A, 41: 
217. 1967 (1968), Basionym: Nephrodium priamaticua Desv., Mem, 
Soc, Linn, Paris, 6: 256, 1827, Mascarenes, I-ladacascar. 

Th, procurrens (Mett,) Reed, comb, nov, Basionym: Aspidiun pro- 
currens Mett,, Ann. Ludg. Bat., 1: 231. 186/*, Java, Celebes, 
N. India. 

Th, producta (Kaulf.) Pteed, comb, nov, Basionym: Aspidiun pro- 
ductura Kaulf., Enixra. , 237. 1B24. Phj.lippine lals. (LuzonTT 

Th, prolif«ra (Retz.) Reed, comb. nov. Basionym: Hemioniti s 

prol i f erum Retz., Obs., 6: 38. 1791. Synonyms: Menisciua pro- 
lifenin (Retz.) Swartz, Syn. Fil., 19, 207, I8O6; Ampelopteris 
elegans Kunze, Bot. Zeit., 6: 114. 1848; A, prolif era (fletz.) 
Copel., Gen, Fil., 144. 1947. S. China, N. India, Trop. Africa, 
Malesia, Polynesia, Mascarenes, Australia, 

Th. prolixa (Willd. in L. ) Ching, Bull. Fan Mem. Inst. Biol., Bot. 
10: 254. 1941. Basionym: Aspidium prolixum Willd, in L. Sp. 
PI., ed. 4, 5: 251. 1810, India, Trop. Africa, Mascarenes. 

Th, prominula (Christ) Reed, comb, nov, Basionym: Aspidium pro- 
minvilum Christ, Brill, Boiss., 4: 659. 1896, Costa Rica, 

Th, protecta (Copel.) Reed, comb, nov, Basionym: DiTrppteris pro- 
tecta Copel., Univ. Calif. Publ. Bot,, 18: 221. 1942, New Guinea. 

Th, pseudarfakiana (Hosak.) Reed, comb, nov, Basionym: Phegopteris 
p seudarf akiana Hosak,, Trans, Nat. Hist. Soc. Formos., 28: 147. 
1938. Palau fsl. 

Th. pseudogueintziana (R.Bonap.) Alston, Ferns W. Trop. Af r. , 61, 
1959. Basionym: Dryopteris pseudogueintziana R. Bonap., Bull. 
Jard. Bot. Brux. , 4: 4. 1913. S. Africa, Cameroun, Madagascar. 

Th, pseudohirsuta (Rosenst.) Reed, comb, nov, Basionym: Dryopteris 
pseudohirsuta Rosenst., Med. Rijks Herb. No. 31: 7. 1917. 
Philippine I sis. 

Th, pseudoparasitica (v.A.v.R, ) Reed, comb, nov, Basionym: Dryo- 
pteris pseudoparasitica v.A,v,R., Nova Guinea, 14: 19. 1924. 
New Guinea. 

Th, pseudoreptans (C.Chr., Reed, comb, nov, Basionym: Dryopteris 
pseudo rep tans C.Chr., Ind. Fil., 286. 1905. Synonym: Nephrodium 
debile Baker, Journ. Bot., 1880: 212. 1880. Sumatra. 

Tti, pseudosancta (C.Chr.) Reed, comb, nov, Basionym: Dryopteris 
pseudoeancta C.Chr., Smiths. Misc. Coll., 52: 378. 1909. 
Costa Rica - Guatemala. 

Th, pseudostenobasis (Copel.) Reed, comb. nov. Basionym: Dryo- 
pteris p seudo st enoba si s Copel., Joum. Arnold Arb., 10: 176. 
1929, New Guinea. 



1968 Reed, Index Thelypteridis 307 

Thelypti?i*i8 ptarmica (Kunze ex Mett.) Reed, comb, nov. Basionym: 
Aspidiu m ptarmi cum Kunze ex Mett., Pheg. u. Asp,, 80, n, 191. 
1858. S'. BraziY. 

Th, ptaradcifojrnds (C.Chr. et Rosenst. ex Rosenst.) Reed, comb, nov, 
Basionym: Dryopteris ptar n dcifomria CChr, et Rosenst, ex Rosen- 
st., Fedde Repert., 12: 472. 1913. Bolivia. 

Th. pterifolia (Mett. ex Kuhn) Reed, comb, nov, Basionym: Aspidiua 
pterifol ium Mett, ex Kxihn, Linnaea, 36: 110, I869, ^nonym: As- 
pidium glei chenioide s Christ, Bull, Herb. Boiss,, II, 4: 960, 
1904. Guatemala, Colombia, Bolivia. 

Th. pteroidea (Klotzsch) Tryon, Rhodora, 69: 8, 1967, Basionym: 
Polypodiun pteroideum Klotzsch, Linnaea, 20: 389. 1847. Colom- 
bia-F,cuador-Bra zil , 

Th, pterospora (v,A,v,R. ) Reed, comb, nov, Basionym: Dryopteris 
pterospora v.A,v,R,, Bull, Buit., Ill, 2: 148. 1920. Sumatra, 

Th, puberula (Bak, in Hook, et Bak,) Morton, Amer, Fern Journ, , 48: 
138, (1958) 1959. Basionym: Nephroditia puberulum Bak, in Hook, 
et Bak., Syn, Fil., ed. 2, 495. 1874, non Aspidium puberulum 
Fee, Mem. Foug. 10: 40, 1865 (illegit, nom. ). nee As. puberxilum 
Gaud, in Freyc. Voy. Bot., 342. 1827. Centr, Amer., Mexico. 

Th, pubescens (L.) Proctor, Bull. Inst, Jamaica, Sci. Ser, , No, 5: 
63. 1953. Basionym: Polypodium pubescens L,, Syst, Nat,, ed. 
10, 2: 1327. 1759. West Indies, Jamaica, 

Th. pubirachis (Bak.) Reed, conb. nov, Basionym: Nephrodiun pubi- 
rachis Bak., Journ. Bot., 1876: 344. 1876, Fiji, Samoa. 

Th, pusilla (Mett. in Triana et Planch.) Ching, Bull. Fan Mem. Inst. 
Biol., Bot. 10: 254. 1941. Basionym: Aspidium pusillua Mett. in 
Triana et Planch,, Ann, Sci. Nat,, V, 2: 245. I864. Colombia, 

Th, pustulosa (Copel.) Reed, comb. nov. Basionym: Cyclosorus pus- 
tule sus Copel., Philip. Journ. Sci,, 81: 37. 1952, Philippine 
Isls, (Luzon, Mindoro). 

Th, quadrangularis (Fee) Schelpe, Journ. S. Afr. Bot,, 30(4): 196. 
1964. Basionym: Nephrodiun quadrangulare Fee, Gen, Fil., 308. 
1850-52, Synonym: Dryopteris conti^ a Rosenst., Med, Rijks Herb,, 
No. 3I: 8, 1917, Pantropical: Borneo, Brit. Guiana, Africa (Mo- 
cambique, Zambesia, Ivory Coast, Oubangui). 

Th, quadriaurita (Christ) Reed, comb, nov, Basionsm: Dryopteris 
quadriaurlta Christ, Philip. Journ, Sci., 2C: 209. 1907. New 
Guinea, Philippine Isls (Mindanao). 

Th, quadriquetra (v.A.v.R. ) Ching, Bull. Fan Mem, Inst, Biol,, Bot, 
10: 254. 1941. Basionym: Dryopteris quadriquetra v,A,v,R, , 
Nova Guinea, 14: 16, 1924. New Guinea, 

Th, quaylei (E,Brown in E, et F. Brovm) Ching, Bull, Fan Mem, Inst, 
Biol., Bot, 10: 254. 1941. Basionym: Dryopteris quaylei E.Brown 
in E. et F. Brown, Bemic© P. Bishop Mus, Bull., 89: 28, f, 9, 
1931. Marquesas, 

Th. quelpaertensis (Christ in Lev.) Ching, Bull. Fan Mem. Inst. 
Biol., Bot, 6: 328. 1936, Basionym: Dryopteris quelpaertensis 
Christ in Lev., Bull, Acad. Geogr. Bot. Mans, 7. 1910, Synonyms: 
Nephrodium laontanma var, fauriei Chidst, Bull. Herb. Boiss,, 4: 
671. I896; Dryopteris Christiana Kodaaa ex Koidz., Bot, Mag, 
Tokyo, 38: 107, 1924. Korea, Aleutian Isls., Kamtschatka, 



308 PHYTOLOGIA Vol. 17, no. h 

Thelypteris quelpaertensis var, yakuaontana (Masaa. ) Tagawa, Act* 
Phytotax. Geobot., 5: 196. 1936. Basionym: Dryopteris yaku - 
montana Masam. , Joum. Soc. Trop. Agr. Forraos., U: 76. 1932* 
Taiwan, Kyushu. 

Th, rampana (Bak.) Reed, corab. nov, Basionym: Nephrodium ramp an 8 
Bak., Journ. Bot., 1889: 177. 1889. Centr. China. 

Th, randallii Maxon et Morton, in Itorton, Araer. Fern Joum., 53 J 
69. 1963. Jamaica. 

Th, recurabens (Rosenst.) Reed, comb. nov. Basionym: Dryopteris 
recumbens Rosenst., Hedwigia, i+6: 123. 1906. S. Brazil. 

Th. reducta Small, Ferns S.E. States, 254, illus. 1938. (Florida), 
"» Diyopteris, 

Th, reederi (Copel,) Reed, comb. nov. Basionym: Cyclosorus reed- 
eri Copel., Amer. Fern Joum., 43: 12. 1953. New Guinea, 

Th, refracta (Fisch. et Mey. ex Kunze) Reed, comb, nov, Basionya: 
Polypodium refract um Fisch. et Mey. ex Kunze, Linnaea 23: 283, 
321. 1850. Brazil, Paraguay, Argentina. 

Th, regis (Copel.) Reed, corab. nov, Basionym: Dryopteria regie 
Copel., Univ. CaUf. Publ.. Bot,, 18: 220. 1942. New Guinea. 

*Th, reichiana (Presl in St^mb,) Reed, comb, nov, Basionym: 
Pecopteris reichiana Presl in Stemb., Flora der Vorwe]t, 2: 
155, t. 37, f. 2. I838, Synonyms: Aspidium reichiamum Ett., 
Die Farnkr, der Jetztwelt, 197. 1865; Pecopteris striata Stemb., 
Flora der Yorwelt, 2: 155, t. 37, f. 3-4. 1838. Upper Creta- 
ceous (Senonian): Ba'^aria, Saxony. Aff. Th. ligulata. 

Th, remotipinna (Bonap.) Reed, comb, nov, Basionym: Dryopteris 
reootpinnna Bonap. . Notes Pterid,, 5: 57. 1917. Madagascar, 

Th. remotipinnata (Hajrata) Alston in Koie et Reching., Biol. Skr. 
Danske Vid. Selsk., 10(3): 10, 1959. Basionym: Dryopteris remo- 
tipinnata Hayata, Gen. Ind. Fl. Formosa, 108. 1917. Sachalin- 
Manchuria-Mpngolia, N & E China, 

Th, repandula (v.A.v.R. ) Reed, comb, nov, Basionym: Dryopteris r^ - 
pandula v.A.v.R., Nova Guinea, 14: 20. 1924. New Guinea, 

Th, repens (Hope) Ching, Bxill, Fan Mera. Inst. Biol., Bot. 6: 304. 
1936, Basionym: Nephrodium repens Hope, Joum. Bombay Najf. 
Hist. Soc, 21: 535. 1899, Himayala - Tonkin. 

Th. reptans (J.F.Qnel.) Morton, Fieldiana, 28(1): 12. 1951; Amer, 
Fern Joum,, 41: 87. 1951. Basionym: Polypodium reptans J.F. 
(inel., Syst. Nat., 2(2): 1309. 1791. Florida - Brazil, Vene- 
zuela, Jamaica. ^ 

Th, reptans var. tenera (Fee) Proctor, Rhodora, 61: 306. (1959) 
i960, Basionym: Goniopteris tenera Fee, 11 Mem, Foug., 60, 
t. 15, f. 3. 1866. Guadeloupe, 

Th, resinifera (Desv, ) Proctor, Bull. Inst. Jamaica, Sci, Ser,, 
No. 5: 63. 1953. Basionym: Polypodium resiniferun Desv,, Berl, 
Mag,, 5: 317. 1811. Synonyms: Nephrodium paname nse Presl, Rel, 
Haenk., 1: 35. 1825; Nephrodium caribaeum Jenm. , Joum, Bot,, 
24: 270, 1886; Lastrea normalis Copel., Gen. Fil,, 139. 1947. 
West Indies, Mexico - Panama. 



1968 Reed, Index Thelypteridis 309 

Thelypteris resinifera var, proraixa (C.Chr. ) Reed, comb, nov, Basi- 

onym: Dryopt erl_s panamenaia var. proxima C.Chr., aniths. Misc, 

Coll., 52: 377'. 1909. Mexico. 
Th, resinosofoetida (Hook.) Ghing, Bull. Fan Mem. Inst. Biol,, Bot, 

10: 254. 1941. Basionym: Nephrodium resinosofoetidxim Hook., Sp. 

Fil., 4: 105. 1862. Costa Rica - Bolivia. 
Th, reticulata (L.) Proctor, B\ill. Inst. Jamaica, Sci, Ser., No. 5i 

63. 1953. Basionym: Polypodlum reticxilatum L. , Syst. Nat., ed, 

10, 2: 1325. 1759. Trop, Amer., Jamaica, 
Th, retusa (Swartz) Reed, comb, nov, Basionym: Polypodium retuaum 

Swartz, Vet, Akad, Handl., 1817: 61, 1817; Lindm., Ark, f, Bot,, 

1: 227, t. 10, f. 11. 1903. BrazU, 
Th, rhorabea (Christ) Reed, comb, nov, Basionym: Dryopteria diver- 
si lob a var, acroBtictioldes subvar. rhombea Christ, Philip, Joum. 

Sci., "Bot. 2C; 200. 1907, Philippine Isls,, Celebes. 
Th. rigida (Ridl.) Reed, comb, nov. Ba3ion3nn: Goniopteris rigida 

Ridl., Trans. Linn. Soc, II, Bot,, 9: 258. 1916, Synonym: 

Phegopteris wollastoni_i v.A.v.R., Mai. Ferns Suppl,, 515. 1917. 

New Guinea. 
Th, rigidifolia (v.A.v.R.) Ching, Bull, Fan Mem. Inst. Biol,, Bot, 

10: 254. 1941. Basionym: Dryopteria rigidifolia v,A,v.R, , Nova 

Guinea, 14: 18. 1924. New Guinea, 
Th, rimbachii (Rosenst.) Reed, comb, nov, Basionym: Dryopteria 

rimbachii Rosenst,, Fedde Repert., 7: 147. 1909. Ecuador. 
Th, riograndensis (Lindm,) Reed, comb, nov, Basionym: Polypodium 

riograndense Lindm., Ark. f.Bot., 1: 230, t. 9, f. 6, 1903. 

S. Brazil, Paraguay, Uruguay, 
Th, riopairdensis (Rosenst.) Reed, comb, nov, Basionjrm: Dryopteria 

riopardenais . Ros. .Hedw, .46: 121. 1906, S, Brazil, 
Th, riparia (Copel.) Reed, comb, nov. Basionym: Dryopteria riparia 

Copel,, Univ. Calif. Publ. Bot., 18: 221, 1942. New Guinea, 
Th, rivulariformis (Rosenst,) Reed, comb, nov, Baaionym: Dryow 

pteris ri vularif ormi s Rosenst,, Fedde Repert,, 6: 316, 1909, 

Synonym: Dryopteris stenophylla Rosenst., Fedde Repert,, 5: 

233. 1908. Bolivia. 
Th. rivularioidea (Fee) Abbiatti, Rev. Mus. LaPlata, Ser II, Bot., 

9: 19. 1958, Basionym: Aspidium rivularioides Fee, Crypt, Vase, 

PI,, 1: 148, t. 50, f. 1. 1869. S. Brazil, Paraguay, Uruguay, 

Argentina, 
Th, rivularioides var, arechavaletae (Hieron,) Abbiatti, Darwiniana, 

13(2-4): 566. 1964, Basionym: Aspidium arechavaletae Hieron, . 

Engl. Bot. Jahrb., 22: 370. 1$96 (1897). Uruguay, 
Th. rivularioides var. pseudomontana (Hieron.) Abbiatti, Darwiniana, 

13(2-4): 566. 1964. Basionym: Aspidium pseudomontaum Hieron,, 

Engl, Bot. Jahrb., 22: 373. 1896 (1897). Argentina, S, Brazil. 
Th. robertiana (Hoffm. ) Slosson ex Rydb,, Fl. Rocky Mts., 1044. 

1917. Basionym: Polypodium robertianum Hoffm., Deutsch. Fl., 

2: 20. 1795. ■= (fymnocarpium. 



310 P II Y T L G I A Vol. 17, no. h 

Thelypteris robinaonii (Ridl.) Ching, Bull. Fan I-Ien. Inst. Biol., 

Bot. 10: 25^. 1941. Basionyia: Lastr^a robir'aonll 5tldl., Joum. 

Feu. Mai. 3tat«8 Mus., 10: (1/! 8-156). 1920; Joum. Mai. Br. Roy. 

Ariatic Soc, U: 65. 1926. Malacca. (= Th. v:l.»cosa). 
Th, rodigasiara (Moore) Reed, comb, nov, BasionTm: Nephrodium rodl_- 

Kaaianuji Moore. L'lll. Hort., 29: 27, t. UU2. ieS2. Samoa. 
Th. roeraeriana (Rosenst.) Reed, comb, nov, Sasionym: Dryopteris 

roeme r iana Rosenst., Nova Gruinea, 8: 723. 1912, New '.>iinea, 
Th, rolandii (C.Chr, ) Tryon, Hhodora, 69: 8. 1967. Basionym: Dryo- 
pteris rolandi i C.Chr., Dansk. Vid. Selsk, Skr., VII, 10: 258. 

1913. Ecuador. 
Th, roraimenais (Baker) Reed, comb, nov, Basionym: Polyx odium ro- 

ralcwnae Baker, Timehri, 5: 214, 1886, Brit. Quiana, 
Th, rosei (M^xon) Trji-on, Rhodora, 69: 8. 1967. Basionya: Dryopterj " 

roaoi I-fexon, Smiths. Misc. Coll., 65(8): 10. 1915. Peru. 
Th. rcsenstockii (C.Chr,) Tryon, Rhodora, 69: 8, 1967. Basionym: 

Dryopteris rosenstockii C. >.r.. Dansk, Vid. Selsk. Skr., VII, 

4: 304, 1907. Ecuador. 
Th, rot\unaensis (St. John) Reed, comb, nov, Basionya: Cyclosorus 

rotumaensis St. John, Occ, Papers Bichop Mus., 21: 180, f. 3. 

1954. Fiji Isls, (Rotxina), 
Th, rubicunda (v.A.v.R. ) K.T-vats., Mem, Coll, Sci., Univ. Kyoto, 

Ser. B, 31(3): 196, 1965. Basionym: Phegopteris rubicunda 

v.A.v.R,, Bull. Jard, Bot. Buit., Ill, 2: l62. 1920, Malaya, 

Sumatra , 
Th. rubida ( J, Smith) K.Iwats., Mem. Coll. Sci., Univ. Kyoto, Ser. 

B, 31(3): 195. 1965. Basionym: Ctoniopteris rubida J.Smith, 

Joum. Hot., 3: 395, 1841. Philippine Isls. 
Th, rubinervis (Mett, ex Kuhn) K.Iwats., Mem, Coll, Sci., Univ, 

Kyoto, Ser. B, 31(3): 195. 1965. Basionym: Phegopteris rubiner- 

vis Mett. ex Kuhn, Linnaea, 36: 116. 1869. Polynesia, 
Th. rubra (Ching) K.Iwats., Joum. Jap. Bot., 38: 315. I963, Basi- 
onym: Dryopteris rubra Ching, Bull. Fan Mem. Inst, Biol,, Bot, 

2: 198, t. 12. 1931. N. India - S. China. 
Th. rudis (Kunze) PrDctor, Bull. Inst. Jamaica, Sci. Ser., No, 5: 

64. 1953- Basionym: Polypodium rude Kiinze, Linnaea, 13: 133. 

I839. Synonym: Polypodium ctenoides Jenm., Bull, Bot, Dept. 

JamaiCH, II, 4: 129. 1897. Jamaica, Mexico - Bolivia, Peru. 
Th. rufostraminea (Christ) Ching, Bull. Fan Mem, Inst, Biol., Bot, 

6: 291. 1936, Basionym: Aspidium ruf o stramineum Christ, Bull, 

Soc. Bot, France, 52 (Mem. 1): 36. 1905. S. China, 
Th, rupestris (Klotzsch) Reed, comb, nov. Basionym: Leptograrama 

rupestr e Klotzsch. Linnaea, 20: 415. 1847. Colombia-Venezuela. 
Th. rupicola (C.Chr,; Ching, Bull. Fan Mem. Inst. Biol,, Bot. 10: 

254. 1941. Basionym: Dryopteris rupicola C.Chr., Fedde Repert., 

15: 24. 1917. Hispaniola. 
Th, rurutensis (Copel.) Reed, comb, nov, Basjonyra: Dryopteris ru- 

rutensis Copel,, Occ, Papers Bishop i*j3., 14: 55, t. 7. 1938, 

S, E.Polynesia (Australian Isls.). 



1968 Reed, Index Thelypteridis 311 

Thelypteris rusbyi (C.Chr.) Tiyon, Rhodora, 69: 8, 1V67. Basio- 

nyn: Dryopteria rusbyi C.Chr., Smiths. Misc. Coll., 52: 390. 

1909. Bolivia. 
Th, rustica (F^e) Proctor, Rhodora, 6l: 306. (1959) I960, Basio- 

njmi PheKopteria rustica Fee, 11® Mem, Foug., 55, t. 13, f. 1, 

1866. Synonym: Dryopteris dominicensis C.Chr., Smith. Misc. 

Coll., 52: 384. 1909. Guadeloupe, Dominica, St, Vincent, Costa 

Rica. 
Th, sagittata ( Swart z) Proctor, Bull. Inst. Jamaica, Sci, Ser., 

No. 5: 64. 1953. Basionym: Polypodium sagittatua Swartz, Prodr,, 

132. 1788, West Indies, Jamaica. 
Th, sagittifolia (Blume) Reed, comb, nov, Basionym: Aspidium 

saKitt aefo lium Blume. Enura. Pl..Jav,, 153. 1828, Java, Perak, 
Th, sagittifolioides (Copel.) Reed, comb. nov. Basionym: Cyclo- 

soras sagittlfolioides Ccpel., Philip. Joum, Sci,, 81: 29, t, 

21. 1952, Philippine Isls. (Samar). 
Th, sakayensis (Zeiller) Reed, comb, nov, Basionym: Nephrodium 

sakayense Zeiller, Bull, Soc. Bot. France, 32: 75. 1385. Perak. 
Th, salicifolia (Wall, ex Hook.) Reed, comb, nov, Basionym: Menis- 

ciura salicifolium Wall, ex Hook., Icon. PI., t. 990. 1854. Pe- 

nang, Singapore, Malaya, Sumatra, Borneo. 
Th, salzmannii (Fee) Morton, Los Angeles County Mus. Contrib, Sci,, 

35: 7. i960. Basionym: Meniscium salzm annii Fee. Gen, Fil,, 

223 ( salzmann l ) . 1850-52, Amer, trop, 
111, saraarensis (Copel.) Reed, comb, nov, Basionym: Cyclosoras 

samarensia Copel.. Riilip. Joum. Sci., 81: 35, 1952. Philip- 
pine Isls. (Samar;. 
Th, sanbiranensis (C.Chr,) Reed, comb, nov, Basionym: Dryopteris 

sambiranensis C.Chr., Cat. PI. Hadagas, Pterid,, 26, 1932 (noa.); 

Dansk Bot. Ark., 7: 50, t. 12, f. 11, 1932, Madagascar. 
Th, ssirapsoni (Bak.) K.Iwats., Mem, Coll, Sci., Univ. Kyoto, Ser, 

B, 31(3): 192. 1965. Basionym: Polypodium sanrpsoni Bak,, Ann. 

Bot., 5: 471. 1891. Tonkin. 
Th, sancta (L.) Ching, Bull. Fan Mem. Inst. Biol., Bot, 10: 254. 

I94I; Proctor, Bull. Inst. Jamaica, Sci. Ser., No. 5: 64, 1953. 

Basionym: Aero sti chum sanctum L, , Syst. Nat., ed. 10, 2: 1320, 

1759. West Indies, Jamaica; Guatemala - Quito, Peru, 
Th, sancta var. hirta (Jenm. ) Reed, comb. nov. Basionym: Nephro- 
dium sanctum var, hirtunn Jenra, , Bull. Bot, Dept. Jam,, II, 3: 

20. 1896, Jamaica, 
Th. sancta var. Jamaicensis (Bak, in Jenm,) Proctor, Bull, Inst. 

Jamaica, Sci. Ser., No. 5: 64. 1953. Basionym: Nephrodium ^Jamai- 

cense Bak. in Jenm., Joum. Bot., 15: 264. 1877. Jamaica, 
Th, sancta var, magna (Jenra,) Proctor, Bull, Inst. Jamaica, Sci. 

Ser., No. 5: 6/*. 1953. Basionym: Nephrodium sanctum var. ma/yium 

Jenm., Bull. Bot. Dept. Jamaica, II, 3: 20. 1896. Jaimaica, 
Th, sancta var. portoidcensis (Kuhn) Morton, Amer, Fern Joum., 

53: 64. 1963. Basionym: Aspidium sanctum var. portori cense 

Kuhn, Bot. Jahrb. Ehgl., 24: 115. 1897. Puerto Rico. 



312 P H y T L G I A Vol. 17, no. U 

Thelypteris sancta VcT. strigosn (C.Chr.) Rued, conb. nov, bAisio- 

nym: Dryopteris sancta var. strigosa C.Chr,, Smiths. Misc. "^11,, 

52: 379. 1909. Cuba, Puerto fiico. 
Th, 8anctif«rrais (C.Chr.) Reed, comb. nov. Basionyra: Dryepteris 

sanctifomda C.Chr., Vid. S^lsk. 5kr., VII, 10: 130, f. 12D. 

1913. Panama - Ecuador. 
Th, Bandvdcensis (Hook. «t Am.) Fosborg, 9cc. Papers Bishop Hus., 

23(2): 30. 1962; I.e., 23(8): 129. 1?66. Basjonym: Polypodium 

sandwicense Hook, ot Am., Hot. Beechey Voy., 105. 1832. = Ctcni- 

tis sandvdcennis. 
Th, savaienais (Bak.) Reed, comb. nov. Baeionyra: Nephrodiun aaval - 

ense Bak., Ann. Bot., 5: 318. 1891. oamoa, 
Th . saxatilis R.P.St. John in Snail, Ferns, S.E. States, 236, illus. 

1938. = Dryopteris, 
Th, saxicola (Swartz) Reed, comb, nov, Basionyia: Polyrjodj.iig saxi - 

eola Swartz, Vet. Akad. Handl., 1817: 59, t. 3, f. 5. 1817. 

Costa Rica - Peru. 
Th, scaberula (Ching) Reed, comb, nov, Basionym: Cyclosorus sca- 

berulus Ching. Bull, Fan Mem, Inst, Bi»l., B«t. 8: 223. 1938. 

China (Hainan) . 
Th, scalaris (Christ) Alston, Joum. Wash, Acad, Sci., hS{7): 23i*. 

1958, Basionym: Aspidiun scalare Christ, Bull, B-tiss., II, 6: 

159. 1906, Mexico t» Guatemala and Costa Rica, and Colombia, 
Th. scallanii (Christ in Baronl et Christ) Morton, Amer. Fern Jcum,, 

56(4): 179. 1966, Basionym: Aspidium scallanii Christ in Ba- 

roni et Christ, Bull. See. Bot. Ital., 1901: 296, 1901, Chin* 

(Szechuan), 
Th, scalpturoides (Fee) Reed, comb, nov, Basionyia: Phegopteris 

scalptureidea FeTe, 11^ M^m. F«ug., 51. 1866. Cuba, 
Th, scalpturoides var, jamaicensis (C.Chr.) Reed, comb, nov, Basi- 
onym: Dryopteris scalpturoides var. jamaicensis C.Chr., Kgl, 

Dansk Vid. Se.lsk. Skr,, 7: 299. 1907. Jamaica, 
■Ri, scariosa (Rosenst.) Reed, comb, nov, Basienym; Dryopteris 

a carlo sa Rosenst., Hedwigia, 46: 127, 1906, Brazil, 
Th, schaffneri (Fee) Reed, comb, nov, Basionyia: Nephrodiua schaf- 

fneri F ee, 8 Mem. Foug,, 108. 1857. Mexico, 
Th. sclerophylla (Poepp, ex Spreng,) Morton, Amer. Fern Joum., 

41s 87 (err, "Kunze"). 1951. Basienym: Aspidiun sclerophyllum 

Poepp, ex Spreng. in L. Syst, Veg., ed. I6; 4: 99, 1827; Kunze, 

Linnaea, 9 J 92. 1834. Florida, V/est Indies (Cuba, Janaica, Puerto 

Ri CO , Hi spaniola ) , 
Th, scolopendrioides (L.) Proctor, Bull. Inst. Jamaica, Sci, Ser,, 

Ne, 5: 64, 1953. Basionyra: Polypodium scolopendrioides L. , Sp. 

PI., 2: 1085. 1753, ^monyms: Polypodium incisua Sv/art,z, Prodr, 

Veg, Ind, Occ,, I3I, 1788; Qoniepteris strigosa Fee, 11« Hem, 

Foug,, 59, t, 15, f. 1. 186^^ West Indies, 
Th, seraihastata (Kunze) Ching, Bull. Fan Mem. Inst. Biol,, Bot, 

10: 254, 1941. Baaionym: Aspidium semihastatua Kunze, Linnaea, 

9: 91. 1834. Peru, 



1968 Reed, Index TheljTJteridis 313 

Thelypteris seriacea (Scott in Bedd,) Reed, comb, nevo Basicnyra: 

Lastrea serice a Scott in Bedd,, Ferns Brit, India, t, 3^8, 

1869, nen Dryopteris sericea C.Chr., Bot. Gaz., 56: I36. 1913. 

India, Burma, China ( Yunnan )„ 
Tho serra (Sv/artz) R.P.St, John in Small, Feme S.E. States, 2i*l, 

illns, I93S. Basionym: Polypodium serra Swartz, Prodr,, 132o 

17^8, West Indies, Jainaicao 
Th, serrata (Gav.) Alston, Kevr Bull. 1932: 309. 1932. Basionyn: 

Meniscium serratum Cavo, Descr. PI,, 5^*8, 1803. Florida, West 

Indies; Mexico - Bolivia, Brazilo 
Tho serrulata (Swartz) Proctor, Bull. Inst. Jamaica, Sci, Ser,, 

No, 5: 65, 1953.. Basionym: Polypodluro serrulatum Swartz, Schrad, 

Joum. Bot., 1800(2): 25. I8OI0 Jamaica. 
Th, aerrutula Ching, Bull, Fan Kern. Inst.^iol,, Bot. 6: 319. 1936. 

China (Szechuan), 
Tho sessilipinna (Copel.) Reed, comb, nov, Basionym: Dryopteris 

seasilipinna Ccpel., Philip, Journ, Sci., Bot. 6: 145. 1911. 

Philippine Isls, 
Th. setigera (Blume) Ching, Bull, '^an Nera. Inst. Biol., Bot. 6: 

345, 1936, Basionym: Cheilanthee 3eti/i;era Blume, Enum, PI, Jav,, 

138, 1828. Continental S. Asia, 1-ialaysia, Philippines - Poly- 
nesia, Taiwan, 
Th. setosula Reed, nom, nov, E^onym: Nephrodiuia angusti folium 

Presl, Spia,, Bot., 48. 1849e Philippine Isls, (Luzon), 
Th, sevlllana Reed, noTi. nov. Synonym: Cyclosorus glaber Copel,, 

Philip. Journ, Sci., 81: 34. 1952„ Philippine Isls (Bo hoi, 

Se villa River), 
Th, siaabonensis (Hieron.) Abbiatti, Danviniana, 13(2-4): 566, 

1964. Basionym: Aspidium siambonense Hieron,, Engl. Bot^, Jahrb., 

22: 372. (I896) 1?97, Argentina (Tucuinan). 
Th, sikkiraensis (Bak.) Reed, comb, nov, Basionya: Aspidittm sik- 

kimense B ak. in Hook, et Bak., Sjm, Fil,, 256, 1867. Sikkim. 
Th. silvatica (Pappe et Rav;son) Reed, comb. nov. Basionym: Gonio- 

pteris silvatica Pappe et Rawson, Syn. Fil. Afr, Austr., 30. 

1858, Synonyms: Goniopteris patens Fe'e, Gen. Fil,, 253. 1852, 

non Polypodium patens Swartz, 1788; Gyrmo/Traa'ina unita Kunze, 

Linnaea, IS: 115. 1844, ncn Polypcdiura unitum L. , 1759. Giiana, 

Liberia, S, Africa, 
Th. sindllinia (C.Chr.) K.Iwats., Acta Phytotax. Geobot., 21(5-6): 

169. 1965. Basionym: Dryopteris simill iina C.Chr., Ind. Fil., 

292. 1905. Synonym: Nephrodium simulans Bak., Journ. Bot,, 

1888: 325. 1888, non Bak., 1874. Borneo, 
Tho simozawae Tagawa, Acta Phytotax, Geobot., 6; 157. 1937. (Tai- 

wan). = Th. angularilobao 
Th. simplex (Hook.) K.Iwats., Mem. Coll, Sci., Univ. Kyoto, 31(3): 

l?9o 1965o Basionym: Meniscium rimplex Hook., Lond. Joum, 

Bot., 1: 294, to 11, 1842. So China, Tonkin, Indochina, 

Taiwan, ??yukyu3o 
Th, ?iiriplex var. trifoliata (Ching) Reed, corabc nov, Basionym: Ab^- 

eopter is simplex var. trifoliata Ching, Bull. Fan Mem, Inst., 

Biol., 'Bot. 10: 10. 1940, China (Fukien). 



"il^ P }( y T L G I A Vol. 17, no. h 

Thelypteris aimplicifolia (J.'>dt.h ex Hook.) Reed, comb. nov. Basi- 
onyjTi: AsplHiu m slm^jj^ cifoli^im J.Smith ex Hook., Icon, PI., t. 
919. 1B5/*. ITiilipplne Ifln,, Fiji. 

Th, si mulana Chinp, Bul\. Fan Mom, Inst. Biol., Bet. 6: 230. 1936. 
(Taiwan). = Th. auriculata, 

Tho sinaulata (Davenp.) Nieuwl., Amer. I'ldl. Nat., ].: 226, 1910. 
Baaionym: Aspidiu m simulatuji Davenp., Bot, Gaz., 19: i*95. 1894. 
Car.idT (P.E.I, to 3. C^uebec), south to li.Z. Alabama, New York 
and Wept Virginia. 

Th. finf^alanenpis (Bak.) Ching, Bull, Fan Mem. Inst. Biol., Bot. 
6: 334. 19360 Basionjni; Nephr o djum sin/^^alan'gnse Bak., Joum, 
Bot., 1880: 212. 1880 „ Synonym: Nephr odiu/n d'ayi Bedd., Joum. 
Bot., 1?87: 323« 188?. Malaya, Sureatra, Borneo", PeraK, Taiwan, 

Th, sintenisii (Kuhn et Christ ex Krug in Urban) Reed, corab, nov. 
Basionyra: Asp idium sintenisii Kuhn et Christ ex Krug in Urban, 
Engl, Bot. Jahrb.," 24: il9o 1897; Urban, Syab. Ant,, 4: 19. 1903. 
Puerto Picoo 

Th. skinneri (Hook.) Reed, comb, nov, Basionym: Aspidium akinneri 
Hook., Icon. PI., t, 924. 1854. Guatemala, Ecuador. 

Tho aodiroi Reed, nom. nov. Synonjin: .Jephrodiux* nenorale Sodiro, 

Crypt. Vasco Quit., 267. 1893; Thelypterig nercoralis ("^od.) Tryon, 
Rhodora, 69: 7. 1967, non Th. nemoralis Ching, 193^. Ecuador, 

Th, sogerensis (Gepp) Reed, comb. nov. Basionyi": Dryopteris so^er- 
ensi 3 Gepp, Joum. Dot., 1923 (Suppl.): 61. 1923a Nev; Guinea. 

Th. spekei (Bak, in Hook, et Bak.) Ching, Bull. Fan Mem. Inst. Biol., 
Bot. 10: 254. 1941. Basionym: NephrodiuTi spekei Bak. in Hook, 
et Bak., 5yn. Fil., 263. 1867o West Africa, iComores Isls. 

Th, spenceri (Copel. ex Christ) Reed, corab. noVo Basionym: Dryo- 
pteris spenceri Copel, ex Cl-irist, Philip. Joum. 5ci., Bot. 2: 
290. 1907. Philippine Isls. 

Th, spinosa (Copel.) Reed, comb, nov, Basionym: Dryo pteris spinosa 
Copel., Univ. Calif. Publ^ Bot., 18: 219. 1942. "Synonyn: Las- 
trea amiata Copel,, Gen, Fil., 138. 1947o Nev Guinea, 

Th." ''Plnulos a (O.F.Muell.) Hieuwl., Aner. Mdl. Nat., 1: 226. 1910. 
BasionjTa: Polyp odium spinu l osum O.F.Muell,, Fl. Fridr., 113, 
f. 2. 1767. =■ DryopteriB. 

Th, spiniilosa Tar, americana (Fisch, ex Kunze) '^eatherby, Rhodora, 
21: 178. 1919. Basionym: Aspidium spinulosum (var,) airericanum 
Fisch. ex Kunze, Amer. Joum, Sci,, II, 6: 84. 1848. = Dryopteris, 

Th o spinulosa var, concordiana (Davenp,) Weatherby, Rhodora, 21: 
178, 1919o Basionym: Mephrod ium spinulosiua var, ccnccrdianua 
Davenp,, Rhodcra, 6: 33. 1904= = Dryopteris intermedia var, 

Th. spinulosa var, dilatata (Hoffni.) St„ John et Warren, Pr-lim, 
List nV xKaniksu Nat, For., 1: 1. 1925. Basionym: Polypodium 
dilatatun Hoffra., Deutsch. Fl., 2: 7. 1795. = Dryopteris. 

Th , spin ulosa varc fructu osa (Gilbert) Femald, Rhodora, 28: I46. 
19260 BasionjTfn: Mephrodium gpinulosuro (var.) fructuosum Gil- 
bert^ List N. Amer. Pterid., 37- 1901. = Dryopteris intermedia. 

The spinulosa var, intermedia (Muhl.) Nieuwl,, Amer. Midlo Nat., 
2: 278. 1912i Weatherby, Rhodora, 21; 178. 1919 o Basionjnn: 



1968 Reed, Index Thelypteridis 315 

Folypodium vf;l Aspidium i ntermedium Kuhl. ex "..'illd. in Lo Sp. 

PI., ed, U, 5: 262. 1810, = Drj-'opteris intermedia. 
Th. sprengelii (Kaulf.) Prcctor, Pull. Inst, Jamaica, Sci, Ser„, 

No. 5: 65, 1953. Basionym: Aspidium sprenfielii Kaulf,, Flora, 

1823(1): 365. 1823, = 7h, balbisiio 
Tho sprucei (Bak, in Hook, et 3ak,) Ching, Bull, ''an Mera. Inst. 

Biol., Bot. 10: 254. 1941. BasionyTi: Nephrodiujn sprucei Bak, in 

Hook, et Bak., Syn, Fil., 269, 1H67. Ecuadoro 
The squamae stipes (Clarke) Ching, Bull. Fan Kem, Inst. Bid., Bot, 

6: 281, 1936, Basionyro: Pcl.-^niodium appendiculatum varo squamae- 
stipes Clarke, Trans. Linn. Soc., II, Bot. 1: 543, t. 79, f. 2. 

1880, Hiraalaya, Sikkira, N, India, 
Dlo 3ouandf:era (Schlecht„) Ching, Hull. Fan Mem, Inst, Biol,, Bot, 

6: 329. 1936 (pro errn . " squamulos a" ) . Basiomin: Aspidium thely> - 

pteris vara squami ^enim Schlecht,, Adiunbr, , 23, t, 11, 1825o 

(Trop. et Austr. Africa, lladagascar, S, India, New Zealand) = 

Th, palustris var, vel Th. confluens, 
Th„ squandpes (Copel.) Reed, comb, noVo BasJonym: Dry o pterin 

squamj-pes Copel., Philip, Joum, Sci., 56: 99, t, 5. 1935. 

Philippine Isls. (Mindanao) „ 
Tho squainulopa (Presl) Ching, Bullc Fan Kera„ Inst, Biolo, Boto 6: 

5, 329. 19360 Basionym: Lastrea squamulosa Pre si. Tent, Pterid,, 

76 (nomo nud.), 183b; Nephrodiun squainulosuiri ("Presl") Hook, f., 

T?] , New Zealand, 2: 39, 1855. = Th, confluens, 
Th, standleyi (Maxon et Morton) Tryon, Rhodora, 69: S, 1967, Basi~ 

onjnn: Dryopteris standleyj. Maxon et Morton, Bull, Torr. Bot, 

Club, 65": 368, 193^5, Guatemala, 
Th, stegnogramnoides (Bako) Fosberg, Occ. Papers Bishop Mus,, 23s 

30. 1962o Basionym: Polypodiuir ste/TioRrammoides Bak,, Syn, Fil,, 

317. 1867, Hawaiian Isls, 
Th, stellato-pilosa (Brause) Reed, comb, nov, Basionyn: Dryopteris 

stellatc-pilosa Brause, Fiigl, Bot. Jahrb., 56: 96, 1920, New 

Guinea „ 
Th, stenobasis (C.Chr,) Ching, Bull. Fan Mem, Inst, Biol,, Bot, 10: 

254« 19i+lo Basionym: Dryopteris stenobasis C,Chr„, Ind, Fil,, 

294 o 1905. = Th. attenuate. 
Th, stenodonta (Copel.) Reed, comb, nov, Basionym: Cyclosorus 

stenodontus Copel,, Philip, Journ, Scio, Bl: 28, t, 20, 1952, 

Philippine Isls. (Panay). 
Tho stenolepis (Bako) Reed, comb. noVo Basionym; Polypodium steno- 

lepis Bak., Kew Bull,, 1893: 231, 1898, Synonjrm: Aspidi um yun- 

nan ense Christ, Bull. Boiss„, 6: 965o 1898, 3. China, 
The stenophylla (Bak,) Reed, comb, noVo Basionym: H enis c ium gteno~ 

p hyllum Bako, Journ, Bot., 1891: 108, 1891, Synonym: Drr/opterlr 

brevlpinna C.Chr,, Ind„ Fil., ?55, 1905, Bomeoo 
Th, stereophylla (v.A,v,R,) Ching, Bull, Fan Mem, Inst. Biol,, Bot. 

10: 254. I94I0 Basionym: Dryopteris stereophylla v.A.v.P,, Nova 

Guinea. 14: 17. 1924, New Guinea, 



316 PHYTOLOGIA Vol. 17, no. U 

Thelypteris stierii (Roaenst.) Reed, comb. nov. Basionym: G^mno- 
p.ramna atierii Rosenst., Fostschr. Alb, v. Bamberg, 6h. 1905. 
S. Brazil. 

Th. stipellata (Eluroe) K.Iv.'Ats., Acta Phytctio , Geobot., 21(5-6): 
16C. 1965» Basionym: Aapidlum ^tipellati un Blume, Snun. PI, Jav,, 
152, 1828, '..'. Malaysia, Johore, Pahang, Trenggam, Perak, Ladrones 
Isls, 

♦Th« gt.iriaca (Unger) Roed, oonb, nov. Basjonyrn: ?clypodit*;a stiri - 
acua Unger, Cblor. Prot., 212, t. 36, f. 1-6. IBh?. Synonyms: 
Goniopteris stiriaca (Unger) A,Braun,(Ueber Foss. Goniopteris- 
Arten) Zeitachr, Geol. Ges., U: 553, 556. Ifc51; Krftusel. Palao- 
botanische Notizen, VIIT. 1927; Lactrea stiriaca (Unger; Veer, 
Fl. Tert. Helv., 1: 31 ( styriaca ). t. 7-8. 1«55; I.e., 3: 151« 
1859; Dotzler, Palaeontographica, 83B: U, t. 1, f. 1, t. 2, fo 
1-2. 1938; Phegopterlg stiriaca (Unger) Ktt,, Die Famkr, der 
Jetztwelt, 195. 186$; Dryopteris stiriaca (Unger) Palibin, 1937; 
Cyclosorua gtiriacua (Unger) Granbast, Ann, Paleont., 48: lo6o 
1962; Ghing et Takht, in Takht., Paleobotanika, U: 195, t. 2, f. 
1-3. iy63; Lastrea helvetica Heer, Fl. Tert. Helv., 1: 33, t, ^, 
f. 2a-2c, 1855. Upper Tertiary (Oligocene-Kiocene): Swi.tzerland, 
France; (Neogene): Transcaucasia Goderdzi Pasa, S.W. Georgia, SSR, 

Tho stokeaii (E.Brown in E. et T. Brovvn) Reed, comb, nov, Basionym: 
Dryopteris stokesii E.Bro'/m in K. et F. Brown, Bishop >fas. Bull., 
89: 20, f. 6. 1931. Polynesia (Rapa Isl.). 

Th, straminea (Bak, in Hook, et Bak.) Reed, comb, nov, Basionym: 
Polypodium stramineum Bak, in Hook, et Bak., 3yn. Fil., 316. 
1867. Venezuela . 

Th, striata (Schuia.) Schelpo, Joum, S« Afr. Bot,, 21(A): 268,, 
1965. Basionym: Aapidium striatum Schum. , Kgl, Dansk Vide 
Selsko, Afdo 4: 230, 1829. Synonyms: Polypodium pallidivenium 
Hook,, Sp. Fil., 5: 8. 1863; Dryopteris hemitelioides Christ, 
Ann, Mus. Congo, 5: 26. 1909. Trop, Africa (Guinea, "Canerouns, 
Congo, Senegal - Angola). 

Th, striata var, molundensis (Brause) Reed, comb, nov. Basionym: 
DryopterJB molundensia Brause, Engl. Bot. Jahrb,, 53: 378o 1915. 
Trop. Africa (Oubangui, Cameroun), 

Th, strigosa (Willd,) Tard. in Humbert, Fl. Madagas, Fara, 5> 1: 27/^, 
f . 38(1-15). 1958, Basionym: Aspidjum strigosum Willd o in L. 
Sp, PI,, ed. Uf 5: 249. I8IO0 Madagascar, Mauritius, Mascarenes, 
Reunion, S. Rhodesiao 

Tho strigosissima (Copel.) Reed, comb, nov. Basionym: Dryopteris 
strigosiasima Copel., Univ. Calif, Publ, Bot., 18: 221. 1942, 
New Guinea o 

The atruthiopteroides (C,Chr«) Reed, comb, nov, Basionym: Dryo- 
pteris struthlopteroides CoChro, Smiths. Misc. Coll,, 52: 3S8. 
1909 Guatemalao 

Th . stuebelil (Hieron.); I'iurillo, Cat, Illus, Plantes de Cundina- 
marca, 2: 110 ( sttlbelii ) . nomen„ I966. (Colonbia), = Th, thoro- 
sonii. 



1968 Reed, Index Thelypteridis 317 

Thelypteris subalpina (v.A.v.R.) Reed, comb, nov, Basionym: Drjo- 

pteris subalpina v.A.v.R., Bull. Buit., III. 5: 200. 1922, TernatCo 
Th, subandina (C.Chr, et Roaenst. ex Rosenst.) Tryon, Rhodora, 69: 

8. 1967» Basionym: Dryop terls aubandln a C.Chr. et Rosenst. ex 

Rosenst o, Fedde Repert., 12: 472. 1913. Boliviao 
Thg Eubapppndioulata (Copel,) Reed, comb, nov, Basionym: Dryopteris 

subappendlcvilata Copel., Univ. Calif. Publ. Bot., 18: 220. 19k2, 

New Guineao 
Tho subarida (Tatew« et Tagawa ex Tagavra) Reed, cotnb. nov, Basio- 
nym: Cycloporus subaridu s Tatew. et Tagawa ex Tagawa, Acta Phyto- 

tax. Geobot., 7: 77. 1938, Philippines, Taiwan, China (Kwang- 

tung, Fukien, Chekiang, Kwangsi), 
Th. pubattenuata (Rosenst.) Reed, combo nov, Basionym: Dryopterl p 

subattenuata Rosenst., Fedde Repert,, 10: 332. 1912. New Guinea, 
Th, subaurita (Tagawa) Ching, Bull. Fan Mem, Inst. Biol., Bot, 6: 

276. 1936, Basionym: Dryopterls subaurita Tagawa, Acta Phyto- 

tax. Geobot., 1: 157. 1932, Taiwan, Japan, Ryukyus, 
Th„ subcuneata (Bak„) Reed, comb, nov, Basionym: Nephr-odium sub- 

cuneatum Bak., in Mart, Fl. Bras., 1(2): 487, I87O0 CayennCe 
Th, subdlraorpha (Copel,) Reed, comb, noVo Basionym: Dryopterls 

subdimorpha Copel., Univ. Calif. Publ, Bot., 18: 220. 1942, 

New Guinea, 
Th, subelata (Bak.) Reed, comb, nov, Basionym: Nephrcdium sube- 

latum Bako, Kew Bull., I9O6: 11. I906. China (Yunnan). 
Th, subfalcinella (v.A.v.R.) Reed, comb, nov, Basionym: Dryopterls 

subfacinella v.A.v.R., Bull. Buit., Ill, 2: 151. 1920. Sumatra. 
Th, subglanduligera Ching, Bull. Fan Mem, Inst. Biol., Bot, 6: 323« 

1936. Malay Penino (Perak). 
Th. subimmersa Ching, Bull. Fan Mem. Inst, Biol., Bot. 6: 306. 

1936. China (Hainan). An Th. iramersa (Blume) Ching, ace, Holtto, 

1954. 
Th, subintegra (Bak,) Reed, comb, nov, Basionym: Polypodlum subin~ 

tegrum Bak^ , Joum, 3ot,, 1877: I64. 1877o Ecuador. 
Tho submarginalis (Langsd. et Fisch,) Small, Ferns S,E, States, 

258, illus. 1938. Basionym: Polypodiam submarginal* Langsd, et 

Fisch., Icon. Fil., 12: t. I3. 1810, = Dryopterls^ 
Thi subnigra (Brause) Ching, Bull. Fan Mem, Inst, Biol., Bot. 10: 

254. I94I9 Basionym: Dryopterls subintegra Brause, Engl. Bot, 

Jahrb,, 56: 82. 1920, New Guinea. 
Th, subobliquata (Hook.) Ching, Bull. Fan Memo Inst, Biol., Bot, 

10: 254. 1941, Basionym: Polypodium subobliquatum Hook,, Sp, 

Fil., 4: 240. 1862, Brazil, Guiana," Colombia. 
Th o subochthodes Ching, Bull, Fan Mem. Inst, Biol., Bot. 6: 305. 

1936. (China, Hongkong, Japan, Taiwan, Korea). = Th. esqui- 

rolii var. glabrata, 
Th . subochthodes foniia lacinlata Kurata, Joum. Geobot,, 11(2): 39. 

1962. (Japan, KyushuJ^ 
Th. subpectinata (Copel.) Reed, comb, nov, Basionym: Dryopteris 

aubpectinata Copel,, Bishop Mus. Bull., 93: 9, t. 70, 1932, 

Tahiti. 



318 P H Y T L G I A Vol. 17, no. li 

Thelypteris subpennigera (C.Chr.) Reed, comb. nov. Basionym: Dryo - 
pteria subp8nnlf?:erii C.Chr., Cat. PI. Madagaa. Pterid., 26. l^y^ 
(nomen); Dansk Bot, Ark., 7: 52, t. 12, f. 1-2. 1932. Madagascar. 

Th. subpubeecens (Blume) K.Iwats., Mem. Coll. Sci., Univ. Kyoto, 

Ser. B, 31(3): 173. 1965. Basionym: Aflpldiua aubpubeecenB Bltuae, 
Enum. PI. Jav,, 149. 1^28. Synonym: Aapldjug Jaculoaug Christ, 
Bull. Boi>iB., II, Z»: 615. 190A, Java to S. China, Taiwan, Luzon, 
Ceylon, Okinawa, LiuKlu, Queensland. 

Th, subsimilis (Hook.) Reed, corab. nov, Basionym: Gyanograama sub - 
aiBdlis Hook., Sp. Fil., 5: U2, t. 293, leSU, Fernando Po. 

Th , mbtetragona (Link) E.P.St, John, Amer, Fern Jourr»,, 26: UU, 
1936, Basionym: Pol ypodium subtetraf^onua Link, Hort, Berol., 
2: 105. 1833, = Thelypteris tetragona, 

Th, subulifolia (v.A,v.it.) Ching, Bull, Fan Mem. Inst. Biol,, Bot, 
10: 25i*. 19A1, Basionym: Dryopteris subulifolia v.A.v.R.. Bull. 
Buit., II, 28: 22. 1918, Sumatra. 

Th, ambvillosa Ching, Bull. Fan Mem, Inst, Biol., Bot, 6: 270. 

1936 (err. "(Moore)"). Based on Polyp odium auriculatum Wall, ex 
Hook., Sp, Fil,, Ui 237. 1862; Polypodium subvillosua Moore, 
Ind. Fil., 3O8 (noraen), 1861, - Th. auriculata, 

Th, sulfui^a (E.Brovn in E. et F. Brovm) Reed, comb, nov, Basio- 
nym: Dryopteiris sulfurea E, Brown in E, et F. Brown, Bishop Mus, 
Bull,, 89: 23, t. 2. 1931. Marquesas, 

Th, sumatrana (v,A,v,R. ) Reed, comb, nov, Basionym: Dryopteris 

sumatrana v.A.v.R., Handb, Mai. Ferns, 227. (1908) 1909, Sumatra, 
Annam, New Guinea, Singapore, Malacca, Selanger, Penang, 

Th, superba (Brause) Heed, comb, nov, Basionym: Dryoptftris amperba 
Brause, Eogl, Bot. Jahrb,, 56: 105. 1920. New Guinea. 

Th, aupernitens (Christ) Reed, comb, nov, Basionym: Dryoptei-js 
supemitens Christ, Fedde Repert,, 8: 19, 1910, Costa Rica, 
Panama. 

Th, supraspinigera (Sosenst,) Reed, comb, nov, Basionym: Dryo- 
pteris supraspinigera Rosenat., Hedwigia, 56: 353. 1915. N«w 
Guinea, 

Th. suprastrigosa (Rosenst,) Reed, comb, nov, Basionym: Dryopteris 
atiprastrigpsa Rosenst,, Fedde Repert., 10: 335. 1912 <> New Guinea, 

Th, tablaziensis (C.Chr, ex Christ) Alston, Joum. Wash, Acad, Sci., 
48(7): 234. 1958, Basionym: Dryopteris tablaziensis C,Chr, ex 
Christ, Bull, Boiss,, II, 7: 262, 1907. Colombia, Panama, Costa 
Rica. 

Th, taiwanensis (C.Chr,) K.Iwats,, Mem. Coll, Sci., Univ. Kyoto, 
Ser. B, 31(3): 183, 1965, Basionym: Dryopteris taiwanensis 
CChr,, Ind, Fil,, 297. 1905, Synonyms: Aspidiun lobulatua Christ, 
Bull. Herb. Boiss., II, 4: 614, 1904, non Blume, 1828; Dryo- 
pteris subhispidula Rosenst.. Hedwigia, 56: 3k3<, 1915. Taiwan, 
I^yukyus, S, China (Kwangtung), Micronesia (Palau), 

*Th, taka Shimon sis Reed, nom. nov. Based on Lastrea .laponica Krysh- 
tofovich, Joum, Gool. Soc, Tokyo, 25: 26, t. 15, f. 1-la, 
1918; Nagao, Proc, 3rd Pan-Pacif. Sci. Congr., 2(1926): 1552, 
1928; Jongnans, Foss, Cat., 43: 1437. I960; Takahasi, Jap. 



1968 Reed, Index TheljTjteridis 319 

Journ, Geol. & Geogr., 33: 194. 1962; Dijkstra, Foss, Cat., 68: 

3905. 1968. Tertiary (Paleogene): Japan, Kyushu Isl., Taka- 

shlma Coal Mine, Hlzan. 
Thelypteris tannensis (C.Chr.) Reed, comb. noT, Basionym: Dryo- 

pteris tannensis C.Chr., Ind. Fil., 297. 1905. Synonym: Poly~ 

podium excel sum Bak. in Hook, at Bak., gyn. Fil., 505, 1874, 

non Dear., 1827. New Hebrides, 
Th, t&t«i (Maxon at Morton) Mortan, Aner, Fern Jaum., 51: 38. 1961. 

Basionym: Dryopteris tatai Maxon at Msrton in Moirton, Jaum. 

Wash. Acad. Sci., 28: 529. 1938, Bali via. 
Th. tenebrica (Jenm.) Proctor, Bull. Inst. Jamaica, Sci. Ser., No. 

5: 65. 1953. Basianym: Nephrodium tenebricum Jenm«, Jaum. Bot,, 

1882: 326. 1882. Jamaica. An hybrid Th. sagittata X Th. serrul- 

ata ? 
Th. tanarlfrans (Christ) Ching, Bull. Fan Mem. Inst, Bial., Bot. 

10: 254, 1941. Basionym: ^ypalepis tenerifi*on8 Christ, Philip. 

Joum. Sci., Bot. 3: 274. 1908, Philippine Isls, 
Di. tanarrima (F^e) Reed, comb. nov. Basionym: Aspidium taiier- 

rifflua Fee, Crypt. Vase. Br., 1: 134, t. 43, f. 1. I869. Brazil. 
Th. tephraphylla (Copel.) Reed, camb. nov, Basionym: Dryopteris 

taphra»hylla Capel., Riilip. Jaum, Sci,, 40: 296. 1929. Philip- 

pine Isls. (Mindanao ) . 
Th, terrestris (Copel.) Reed, comb. nov. Basionym: Dryopteris 

terrestris Copal,, Univ. Calif. Publ. Bot., 18: 221. 1942. Naw 

Guinea, 
Th, tatragona (Swartz) Small, Ferns S.E. States, 256, 476, 1938, 

Basionym: Polyp odium tetragonum Swartz, Prodr., 132, 1788, 

Florida, Jamaica - Brazil, Peru, Galapagos Isls. 
Th, tatragona subsp, aberrans Morton, Leaflets of Western Hot,, 

8(8): 194. 1957, Galapagos Isls.^ 
Th, tatragona var. guadalupensis (Fee) Kramer, Acta Bot. Neerl,, 

9:^ 298. i960, Basionym: Goniopteris guadalupensis Fee, 11® 

Mem, Foug,, 64, t, 17, f. 2. 1866, ^Saba, 
Th, tauscharl (v,A,v.R,) Reed, comb, nov, Basionym: Dryopteris 

teuscherl v,A,v.R,, Bull, Dapt, Agr, Ind, Neerl,, 18: 6, 1908, 

Borneo, 
I^ thelypteris (L.) Nieuwl., Amer. Midi. Nat., 1: 226. I9IO. 

Basionym: Acrostichum thelypteris L., Sp. PI., 2: 1071, 1753. 

> Th. palustris. 
Th. thomsonii (Jenm.) Proctor, Bull. Inst. Jamaica, Sci. Ser., 

No. 5: 65. 1953* Basionym: Polypodlum thomsonii Jenm., Joum. 

Bot., 1886: 272. 1886. Synonym: Dryopteris stuebelii Hieron., 

Hedwigia, 46: 340, t. 6, f. 13. 1907. Jamaica, Hispaniola, 

Colombia. 
Th. thwaitesii (Hook.) Reed, comb. nov. Basionym: Meni solum 

thwaitesii Hook., Fil. Eicot., t. 83. 1859. S. India, Caylon. 
Th, todayensis (Christ) Reed, comb. nov. Basionym: Dryopteris 

today ensis Christ, Riilip. Joum. Sci., Bot. 2: 193. 1907. 

Philippine Isls. 



320 P H Y T L D T A Vol. 17, no. h 

Th«lypt«ri8 tomentosa (ThouAro) Chine, Bull. Pan Meia. Inst. Biol., 
Bot. 10: 255. 19i+l. Basionym: Polypodlum toaentoBua ThouArs, 
Fl. Triet. d'Acimha, 32, t. 3- 1804. Bourbon, Tristan d'/.cunha, 
Madagascar, Mascarenes. 

Th, tonkinensis (C.Chr.) Ching, Bull. Fan Mem, Inst. Biol., Bot. 
6: 292. 1936, Baaionym: Dryoptgria tonkinanaia C.Chr., Bull. 
Mu8. Paris, II, 6: 102. 1934. Tonkin, China (Kwangsi). 

Th. toppingii (Copel.) K.Iwats., Acta Phytotax. G«obot., 21(5-6): 
168. 1965. Baslonym: Dryoptaris toppingii Copel., Philip. Joum. 
Sci., 120: 56. 1917. Synonym: Nephrodiuc indicua Hidloy, Joum. 
Mai. Branch Roy. Asiat. Soc, U: 73. 1926. Borneo, Malacca. 

Th, torresiana (Gaudich.) Alston, Lilloa, 30: 11. I960. Basionyn: 
Poly8tich^un torresianum Gaudich, in Froyc. Voy. Bot., 333. 1828. 
Synonyms: Aspidiua uli^nosum Kunze, Llnnaea, 20: 6, 1847; Poly- 
podium tenericaule Wall, ex Hook., Kew Joum. Bot., 9: 353. 1857; 
Nephrodium setigerua rar. calvatua Bak., Joum. Bot., 1875: 201, 
1875; NephrodiuM oligophlebium Bak,, Joum. Bot,, 1875: 291. 
1875; Dryopteris laalocarpa Hayata, Mat. Fl. Fonaos., 417. 1911; 
Dryopteris elegane Koidx., Bot. Mag. Tokyo, 38: 108. 1924; Poly- 
podiun trichodee Beinw. ex J.Snith, Joum. Bot., 3: 394. 1841. 
Trop. Asia, Fiji, Bisnarck Archipelago, India - Polynesia, Phil- 
ippine Isla., Taiwan, Assam, China (Kiangsi), Japan (Kyushu); 
introd, in Trinidad, Brazil, Argentina, 

771. torresiana var. calrata (Bak.) K.Iwats., Mem. Coll, Sci,, Univ, 
Kyoto, Ser, B, 31(3): 154. 1965, Basionya: Nephr odium setigerum 
Tar, calvatum Bak,, Joum, Bot., 1875: 201. 1875. E. 4 Cent, 
China, Korea, Japan (Honshu, Shikoku, Kyushu), 

Th, totta (Thunb,) Schelpe, Joum. S. Afr. Bot., 29: 91. 1963. 
Basionym: Polypodlun tottum Thunb., Prodr. PI. Cap., 172, 1800, 
Synonyms: Aapidium gogfd-lodus Schkuhr, Krypt, Gew, , 1: 193, t. 
33c, 1809; Aspldium ecklonli Kunze. Linnaea, 10: 546. 1936; 
NephrodiuM plantianum Pappe et Raws., Syn. Fil, Afr, Austr,, 139. 
1868, Trop, South Amer,, West Indies, S, Africa, Taiwan, Japan, 
S.E.China, Australia, New Zealand, Hawaiian lals. 

The totta yar, glabra (Mett, apud H,Ito) Reed, e<mb, nor, Basio- 
nym: Dryopteris gong7lodeB Tar» glabra Mett, apud H,It$, Joum, 
Jap, Bot., 11: 786. 1935. Japan, ^ 

Th. totta var. glabra forma glaberriaus (H.Ito) Reed, comb, noY, 
Baslonym: Cyclosorus goggilodus Tar, glaber forma glaberrimis 
H.ItS, Bot, Mag. Tokyo, 51: 714. 1937, Japan. 

Th, totta Tar, glabra forma paucipilosa (H.ItS) Reed, comb, nov, 
Basionym: Cyclosorus go^ilodus rar. glaber fonaa paucipilosus 
H.ItS, Bot, Mag, Tokyo, 51: 714. 1937. Japan, Taiwan, 

Th, totta Tar, hirsuta (Mett,) Morton, Contrib, U,S, Nat, Herb,, 
38(2): 73, 1967. Basionym: Aspldium unitum rar, hirautum Mett,, 
Ann, Lugd, Bat,, 1: 230. I864. Synonym: Pteris polypodioides 
Poir, in Lam,, Bicyd, Meth,, 5: 716, 1804, Brazil, Australia, 
New Zealand, Msxico, Hawaiian Isls, 

Th, totta Tar. longipinna (CChr, ) Morton, Contrib, O.S, Nat, Herb., 
38(2): 74. 1967, Basionya: Dryopteris gongylodes Tar, longi- 



1968 Reed, Index Thelj'pteridis 321 

pinna C.Chr., Dansk. Vid. Selsk. Skr., VII, 10: 194. 1913o 
Brazil, Uruguay, Paraguay, 

Thelypteris tottoides (H.It6) Morton, Amer. Fern Joum., 56(4) s 
179. 1966. Baaionym: Leptogreumna tottoides H.Ito, Hot. Mag. 
Tokyo, 49: 434. 1935. Taiwan, S.E.China (Fukien). 

Th. transversaria (Brack.) Reed, comb. nov. Basionym: Nephrodium 
transversarium Brack., E>q>l. Biqp., 16: 187. 1854. Samoa, ?New 
Guinea. 

Th. triphylla (Swartz) K.Iwits., Mem. Coll. Sci., Unir. Kyoto, Ser. 
B, 31(3): 190. 1965. Basionym: MenJBCium triphyllum Svfartz, 
Schrad. Joum. Bot., 1800(2): 16. 1801. Synonym: Meniscium tri- 
phyllum forma cristatum K.Sato, Joum, Jap. Bot,, 12: 824. 1936. 
Trop. Asia, Australia (Queensland), Malesia, Philippine Isls,, 
N. India, S. China, Taiwan, I^yukyus. 

Th. triphylla Tar. parishii (Bedd.) K.Iwats,, Mem. Coll., Sci,, 
Univ. Kyoto, Ser. B, 31(3): 191. 1965. Basionym: Meniscium 
parishii Bedd., Ferns Brit. India, t. 184. 1866. N, India, 
Burma to Indochina, S to Malaya, I^yukyus, Taiwan. 

Th, tristis (Kunze) Tryon, Rhodora, 69: 8. 1967j Morton, Contrib, 
U.S. Nat. Herb., 38(2): 65-66. 1967. Basionym: Polypodium triste 
Kunze, Linnaea, 9: 47. 1834. Venezuela, Peru. 

Th, truncata (Poir, in Lam,) K.Iwats., Mem. Coll, Sci., Unir, Kyoto, 
Ser, B, 31(1)! 33. 1964. Basionym: Polypodium truncatum Poir» 
in Lam., Kncycl., 5: 534. 1804. Synonym: Dryopteris sublaevi - 
frons Tagawa, Acta Phytotax. Geobot,, 5 J 192. 1936. Pantropio: 
N, India - Malesia, trop. Australia, Polynesia, Madagascar, 
Mascarenes, Taiwan, I^yv^cyus, Bismarck Arciiipelago, Brazil. 

Th« truncata forma kwashotensis (Kayata) Reed, comb, noVo Basio- 
nym: Dryopteris kwashotensis Hayata, Icon. PI, Formos,, 5s 278, 
1915. Taiwan. 

Th, truncata forma laevifrons (Hayata) Reed, comb, nov, Basionym: 
Dryopteris laevifrons Hayata, Icon. PI. Formos., 4: 158. 1914. 
Taiwan, Japan (l^aikyus). 

Th. truncata forma sublaevifrons (Tagawa) Reed, comb, nov, Basio- 
nym: Dryopteris sublaevifrons Tagawa. Acta Phytotax. Geobot., 
5: 192, 1936, LluKiu, Taiwan, Trop. Asia, New Guinea. 

Th, tsaratananensis (C.Chr. ) Ching, Biill, Fan Mem. Inst. Biol., 
Bot. 10: 255. 1941; Tard., Pollen et Spores, 7(2): 334. 1965. 
Basionym: Dryopteris tsaratananensis C,Chr,, Cat. PI. Madagas. 
Pterid., 45. 1932 (noraen); Dansk. Bot, Ark., 7: 45, t. 9, f. 
1-5. 1932. Madagascar. 

Th. tuberculata (Ces,) Ching, Bull. Fan Mem. Inst. Biol,, Bot. 10: 
255. 1941. Basionym: Nephrodium tuberculatum Ces., Rend. Acad, 
Napoli, 16: 26, 29. 1877. New Guinea, 

Th, tuberculifera (C.Chr.) Ching, Bull. Fan Mem. Inst. Biol., Bot. 
6: 295. 1936, Basionym: Dryopteris tuberculifera C.Chr., Contr. 
U.S. Nat. Herb., 26: 275. 1931, Assam, Sikkira, Yunnan. 

Th, tuerckheimii (Donn. -Smith) Reed, comb, nov, Basionym: Nephro- 
dium tuerckheiaii Donn. -Smith, Bot. Gaz., 12: 133, t, 11, 1887, 
Guatemala. 



32? P 1! Y T L G I A Vol. 17, no. h 

Thelypteris turrlalbae (Rooenst.) Korton, Contrib. U.S. Nat. Herb., 
38(2): UU. 1967. Baaionym: Dryoc^teris turrlalbae Roeenst., 
Fedde Repert., 22: 10. 1925. Costa Rica. 

Th, ugoensis Reed, nora. noT. Baaed on Gyclosorua rijtidua Copel., 
Philip. Joum. Sci., 31: 27. 1952, non C. rigidus (Ridl.) Copel., 
Philip. Joum. Sci., 78: /♦52. 1951. Philippine Isle. (Luzon, 
Mt. Ugo). 

Th . viliKlnoaa (Kunze) Chlng, Bull, Fan Mea. Inst. Biol., Bot. 6: 
3Z*2. 193<5. Baaionya: Aapidiua uliginoaun Kunze, Linnaea, 20: 6, 
18A7. - Th. torresiana, 

Th . ulijginosa var. calTata (Bak.) K.Iwats,, Acta Fhytotax. Geobot., 
18: 158. i960. Baaionya: Nephrodium aetigerum rar, calvatua 
Bak., Joum. Bot., 1875: 201. 1875. - Th, torreaiana vax. 

Th . uliginosa var. elegana (Koidz.) K.Iwata., Acta Phytotax. Geo- 
bot., 18: 158. i960, Baaionya: Dryopteris elegans Koidz., Bot, 
Mag. Tokyo, 38: 108. 192/*. - Th. torreaiana. 

Th. unca R.P.St. John ex Small, Ferns S.E. States, 2U>, illus, 1938. 
Florida, 

Th, underwoodiana (Maxon) Ching, Bull, Fan Mea. Inst. Biol., Bot. 
10: 255. 19A1; Proctor, Bull. Inst. Janaica, Sci. Ser., No. 5: 
66. 1953. Baaionya: Dryopteris underwoodiana Maxon, Aaer. Fern 
Joum., 18: 49. 1928. Jamaica. 

Th, uniauricxilata (Copel.) Reed, coab. nov. Baaionya: Dryopteris 
unlauriculata Copel., Philip, Joum. Sci., Bot. 9: 3, 191A. 
New Guinea, 

Th, unidentata (Bedd,) Holtt., Rev. Fl. Mai., 2: 251, f. U3. 1955. 
Baaionya: Laatrea unidentata Bedd., Handb. Suppl., 53. 1892. 
Synonya: Dryppteria monodonta C.Chr., Ind. Fil., 278, 1905, Perak, 

Th, unita (L.) Morton, Amer. Fern Joum., 49: 113, 1959. Baaionya: 
Polypodiua unitum L., Syst. Nat., ed. 10, 2: 1326. 1759, Syno- 
nya: Nephrodiua cucullatum Bak., Syn. Fil., 209. 1867. Trop. Aaia, 
- Polynesia, Mascarenes, Seychelles, Ceylon, Philippine lala., 
Micronesia, Bisaarck Archipelago. 

Th, uraiensis (Rosenst.) Ching, Bull, Fan Mea. Inst. Biol., Bot, 
6; 336. 1936, Baaionym: Dryopteris uraiensia Rosenst,, Hedwigia, 
56: 3iltl. 1915. Synonym: Dryopteria hir suti squama ta Hayata, Icon, 
Fl, Formos., 5: 277, f, 105. 1915. Assam, Taiwan, S. China, 
Japan (Kyushu). 

Th, urdanetensis (Copel.) Heed, coab, nov, Basionya: Dryopteris 
urdanetensis Copel,, Leaflets Philip. Bot., 5: 1682. 1913. 
Philippine I sis. (Mindanao). 

Th, urens (Rosenst.) Reed, coab. nov, Basionya: Dryopteris urens 
Rosenst,, Fedde Repert,, 4: 5. 1907; C.Chr,, Vid. Selsk. Skr,, 
VII, 4: 332, f, 521, 1907, Uruguay. 

Th. urophylla (Wall, ex Hook.) K.Iwats,, The Southeast Asian Stud- 
ies., 3(3): 81. 1965; Acta Phytotax, Geobot,, 22(3): 94. 1966. 
Basionya: Polypodiua urophyllum Wall, ex Hook,, ^, Fil,, 5: 9, 
1363. Synonya: Polypodium pinwillii Bak., Ann. Bot,, 5: 77, 1892, 
N. India - Ceylon, S, Qiina, Malesia-Polynesia, Trop, Australia, 
Sumatra, Malaya, Thailand, Penang^ Borneo? 



1968 Reed, Index Theli-pteridis 323 

ThelTpteris urophylla var. nitida (Holtt.) K.Iwats,, Acta Phytotax. 

Geobot., 22(3): %• 1966; (I.e., 21(5-6): 171 (illegit.)- 1965). 

Basionym: Dryopteris urophylla var, nitida Holtt,, Gard, Bull, 

Straits Settlements, 7: 249. 1923, Malay Penin., Borneo. 
Th, utanagensis (Hieron*) Reed, comb, nov, Basionym: Dryopteris 

utanajtensis Hieron., 46: 333, t. 5> f. 8. 1907, Colombia, Ecuador. 

(Murillo, Cat. Illus, Plantas de Cundinamarca, 2: 110 (nomen). 

1966). 
ni. valdepilosa (Bak.) Reed, comb. nov. Basionym: Nephrodlum val- 

depilosum Bak., Joum. Bot., 1879: 26l. 1879. Colombia. 
Th, valida (Christ) Reed, comb, nov, Basionym: Dryopteris vallda 

Christ, Joum. de Bot,, 21: 230, 261, 1908, Annaau 
Th, varians (Fee) Reed, comb, nov, Basionym: Nephrodlum varians 

Fee, 11® Mem. Foug,, 88, t, 24, f. 2, 1866, Trinidad, Amazonas. 
Th, varievestita (C.Chr, ) Reed, comb, nov, Basionym: Dryopteris 

atrispora var. varievestita C.Chr,, Brittonia, 2: 296. 1937, 

New Guinea, Papua. 
Th, vattuonei (Hicken) Abbiatti, Darwiniana, 13(2-4): 566. I964, 

Basionym: Dryopteris vattixonel Hicken, Dar>dniana, 1: 100, 

1924, Argentina, 
Th, velata (Kunze ex Mett.) Reed, comb, nov, Basionym: Aspidium 

velatum Kunze ex Mett., Pheg, u. Asp., 79, n. 190, 1858, Cuba, 
Th, venuloss (Hook.) Reed, comb. nov. Basionym: Nephrodium venu- 

losum Hook., Sp, Fil., 4: 71. 1862, Synonym: Aspidium elatuai 

Mett. in Kuhn, Fil. Afr., 130. 1868, Guinea, Ivory Coast. 
Th, venusta (Hew.) Proctor, Bull. Inst. Jamaica, Sci, Ser,, No, 5s 

66, 1953, Basionym: Aspidium venustum Hew., Mag, Nat, Hist,, 

II, 2: 464. 1838. Jamaica, 
Th, venusta var. usitate (Jenm. ) Proctor, Bull. Inst. Jamaica, Sci. 

Ser., No. 5: 66, 1953, Basionym: Nephrodium u si tat urn Jenm., 

Joxim. Bot., 17: 26l. 1879, Synonym: Nephrodimn calcareum Jena,, 

Joum. Bot,, 24: 271. 1886. Jamaica. 
Tti, verrucosa (J. Smith ex Presl) Ching, Bull. Fan Mem, Inst, Biol,, 

Bot, 6: 308. 1936, Basionym: Lastrea verrucosa J, Smith ex 

Presl, Epia. Bot., 36, 1849. FWJJLppine I si 8,^ Malaya, Borneo, 

Malacca, Loyalty Isls. 
Th, vermiculosa (v.A.v.R, ) Reed, comb, nov, Basionym: Dryopteris 

▼erruculosa v.A.v.R,, Bull. Jard. Bot. Buit., II, No, 11: 12. 

1913, Java, 
Th, versicolor R.P.St. John ex Small, Ferns S.E.States, 250, illus, 

1938. Florida - Texas, 
Th. vestigiata (Copel,) Reed, comb, nov, Basionym: Dryopteria 

vesti^gata Copel., Univ, Calif, Publ, Bot., 18: 220. 1942, 

New Guinea, 
Th. vlllosa (Link) Reed, comb, nov, Basionym: Gymnograaaa villosa 

Link, Hort. Berol,, 2: 51. 1833. Synonym: Dryopteris dasyphylla 

C.Chr., Ind. FU., 260. 1905. Brazil. 
Th. villosipes (Gepp) Ching, Bull. Fan Mem. Inst, Biol., Bot. 10: 

255, 1941. Basionym: Dryopteris villosipes Gepp in C^bba, 

Dutch N.W.New Guinea, 70. 1917. New Guinea. 



32U P H Y T L G T A Vol. 17, no. U 

ThelTptoris rinosicarpa (t.A.v.R, ) Ching, Bull. Fan Mem, Inst, 

Biol., Bot. 10: 255. 19^+1. BaBionym: DnropterlB vinoalcArpa 

v.A.v.R., Bull. Buit., Ill, 5: 198. 1922. Sunatra. 
Th, rlridifronB Tagawa, Joum. Jap. Bot,, 12: 747. 1936, Based on 

Dryopterls alegana var, aubtrlpinnat^ Tagawa, Acta Phytotax* 

G«obot., 2: 193. 1933. Japan (Honahu, KyuBhu). 
Th, Tiridis (Copel.) Reed, comb, nor, Baeionym: QyclosoruB virld - 

is Copel., Philip. Joum. Sci., 81: 35, t. 24. 1952. Luaon. 
Th, TlBCosa (Bak. in Hook, et Bak.) Ching, Bull. Pan Ken. Inst. 

Biol,, Bot. 10: 253. 1941 (err., "J. Smith"). Basiony«i( U8trea 

TJBCosa J, Smith, Joum. Bot., 3s 412 (nom. nud.). 1841). Nephro- 

diua rlBCoaum Bak. in Hook, et Bak., 3yn, Fil., 264. 1867. Fiji, 

Borneo, Malaya, Philippine I sis. 
Th, Tivipara (Raddi) Reed, conb, nov, Basionym: PolyrxxiiuH viyj - 

para Raddi, PI. Bras., 1: 22, t. 32, 1825. Colombia, Brazil, 

Ecuador, Peru. (Murillo, Cat. Illus. Plantas de Cundinamarca, 

2: 110 (nomen), f. 50, 1966). 
Th* Tulcanlca (Bak.) Reed, comb. nov. Basionym: NephrodiuM vul- 

eanicum Bak., Ann. Bot., 8: 127. 1894. Java. 
Th, wantotensis (Copel.) Reed, comb. nov. Basionym: Dryopteris 

wantotensis Copel., Univ. Calif. Publ. Bot., 18: 220. 1942. 

New Guinea. 
th, warburgii (Kuhn et Christ ex Warb. ) B.C.Stone, Micronesica, 

2: 3. 1966. Basionym: Aspidium warburgLi Kuhn et Christ ex 

Warb., Monsimia, 1: 81, 1900, Nev Guinea. 
Wio wariensis (Copel.) Reed, comb. nov. Basionym: Dryopteris 

warienais Copel., Philip. Joum. Sci., 6: 73. 1911. Mew Guinea. 
Th, warmingii (C.Chr.) Tryon, Rhodora, 69: 8. I967. Synonym: Dryo~ 

pteris warmingii C.Chr., Dansk, Vid. Selsk, Skr,, VII, 10: 227. 

1913. Brazil, 
Th, weberi (Copel.) Reed, comb, nov, Basionym: Qrclosorus weberi 

Copel., Philip. Journ. Sci., 81: 36, t, 25. 1952, Mindanao. 
Th. williamsii (Copel.) Ching, Bull. Fan Mem. Inst. Biol., Bot. 

10: 255. 1941. Basionym: Dryopteris williamsii Copel., Britto- 

nia, 1: 6?, t. 1. 1931. Mindanao. 
Tho wollastonii (v.A.v.R.) Ching, Bull. Fan Mem. Inst. Biol,, Bot. 

10: 255. 1941. Basionym: Phegopteris wollastonii v.A.v.R., 

Mai. Ferns Suppl,, 515. 1917. New Guinea. 
Th. wri^tii (Hett. ex D.C.Eaton) Reed, comb. nov. Basionym: 

Aspidium wrirfitii Mett, ex D.C.Eaton, Mem. Amer, Acad., n.s., 

8: 210. I860. JMsaica^ Cuba. 
Th. xiphioldes (Christ) Reed, comb. nov. Basionym: Dryopteris 

xiphioides Christ, Philip. Joum. Sci., Bot, 2: 201, 1907. 

Philippine I sis (Mindanao), 
Th. xylodes (Kunze) Ching, Bull. Fan Mem. Inst. Biol., Bot. 6} 

296. 1936. Basionym: Aspidium xylodes Kunze, Linnaea, 24: 283. 

1851. India, S. China (Yunnan), Burma, Luzon. 
Th, yunkweiensis Ching, Bull. Fan Mem. Inst. Biol., Bot. 6: 274. 

1936. China (Yunnan, Kweichow), Tonkin. 



1968 Reed, Index Thelypteridis 325 

Thelypterls zambeaiaca (Bak.) Tard., Not, Syst., 14: 3A4o 1952, 

Basionym: Nephrodliim zambeslanum Bak., Ann, Bot,, 5: 318, 1891, 

' Th, longicuspis, 
Th. zamboangensls Reed, nom. nov. Based on Cyclosorus ohrlgtii 

Copel., Fern Fl. Philippines, 2: 362, I960, Philippine Isls, 

(Mindanao, Negro 3, Luzon). 
Th, zeylanica Ching, Bull, Fan Mem. Inst, Biol,, Bot, 6: 28?. 1936. 

Ceylon, 



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New combinations in Thelypterie . Bhodora, 6l: 305-306. (1959). 

I960. 

Notes on Lesser Antillean Ferns. Hhodora, 63: 31-35. 1961. 

Notes on Lesser Antillean Ferns, II. Rhodora, 68: 466-469. 

1966. 
Rauschert, Stephan Die LegitiaitJlt des Gattungsnaoans Thelypteris 

Schmidel. Taxon, 15: 180-184. 1966. 
Scbelpe, Z.A.C.L.E. A review of the Southern African Species of 

Thelypteris . Joum. S. Afr, Bot., 21(4): 259-269. Oct. 1965. 
Soo, R. Ton A magyar Flora es reget^cio rendsMrtani-novenyfold- 

rajzi Kerzikonye, I, I964. Budapest. 
StAfleu, Frans A. Adanson, Michel. Families 4es Plantes, Paris. 

1763 (-1764), 2 Tols. Oct, Taxonondc Literature, Regn. Veg., 

52: 1-2. 1967. 
Tagawa, Motozi Index Pterid„ .nytoinua Japonicorua. 1959. 
Tardieu-Blot, M.L, In: Aubreville, Flore du Gameroun, Vol, 3: 

Pteridophytes, I964, 
— — et CChristensen. In: Lecomte's Flore Generale de I'Indochine, 

7(2), fasc. 8: 355-372. 19a, 
Tryon, RoUa Taxonomic Fern Notes, V, New combinations in Perurian 

species of Thelypteris . Rhodora, 69: 5-8o 1967. 
Walker, Trevor G, A cytotaxonomic study of the Pteridophytes of 

Jamaica. Trans, Roy, See, Edinb., 66: 9): 169-237, 5 pl«. 

(1964-65), 1966, 
Woynar, H, Zur Momenklatur elniger Famgattungen, II, Filix, 

Hedwigia, 56: 381-387. (1915) 1916. 



TWO NEW PINyON VARIETIES FROM ARIZONA 
Elbert L. Little, Jr. 



The pinyon (nut pine) of central Arizona with 1 slender leaf 
or needle in a fascicle, commonly referred to Finus monophylla 
Torr. & Frdm., is named here as a nei-; variety of P. edulis 
Engelm. Another pinyon of the international border of south- 
eastern Arizona, southwestern New Mexico, and adjacent northern 
Mexico, is distinguished as a new variety of its species, P. cetn - 
broides Zucc. ( sens , strict .) . 

Many species of Pinus have broad geographic ranges distributed 
over widely varying climates, altitudes, and soils. Careful tax- 
onomic examination of a widespread species often reveals the pre- 
sence of geographic races and varieties. Like other pines, the 
pinyons exhibit similar geographic variations. 

The 8 species of pinyons (nut pines) in southwestern United 
States and Mexico are grouped together as Pinus L. subsect. Cem - 
broides Engelm. (St. Louis Acad. Sci. Trans, h: 176, 178. l880) . 
Distribution maps have been published by William B. Critchfield 
and Little (U.S. Dept. Agr. Misc. Pub. 991, maps I5-I8. I966) . 
Four species are native in the United States, thoiagh treated by 
some authors as varieties under the oldest name Pinus cembroides 
( sens , lat . ) . Morphological differences in seeds and other char- 
acters are sufficient not only for the retention of these ^i- 
species, but also for the further recognition of additional geo- 
graphical varieties and unnamed races. 

From 1937 to 19^1 I did research on Pinus edulis Engelm., the 
common species of pinyon, in Arizona and Ne\^) !lexico as part of 
the research project of management of pinyon- juniper woodlands by 
the United States Forest Service (U.S. Dept. Agr., Agr. Handb. 
27I: 398-^03, illus. 1965). The large edible seeds of that 
species are knovm as pinyon nuts (from Spanish piffon, plural 
pifTones), pine nuts, and Indis.n nuts. They provide an annu^.l 
harvest of about a million pounds or more. A taxonomic study of 
pinyons iras begun and an abstract was published (Amer. Joixr. Bot. 
27 (10) sup. 2ij-s. 19^4-0 ) . However, work on the project wps dis- 
continued during World War II. 

One nev; variety from the Edwards Plateau in southwestern Texas 
\r3.s published, Texas pinyon, Pinus cembroides Zucc. var. remota 
Little (Wrightia 3: I83. I966) . ^Owing to delay in completing 
the taxonomic study, two additional varieties al].uded to in that 
article are published here. A field trip to Arizona in May I968 
provided the opportunity for further study and collections of the 
tvjo new varieties. 

329 



330 PHYTOLOGIA Vol. 17, no, U 

PINU3 MONOPHYLIA Torr. & Fr4m. cingleleaf pinyon 

Plnus nuDnophylla Torr. & Fr^. Ln FV^. , F^t. Explor. Exped. 
Rocky MtB. 319 > t. h l&^3', " nionophyllug ." 

This is the pinyon of the Great Basin recion, mountains from 
southern Idaho and northern and western Utah to Nevada, central 
and southern California, and northvectern Arizona, also northern 
Lower California, Mexico. Leaves 1 (rarely 2) in a fascicle, 2-6 
cm. long, terete, 1.5-2 mm. wide, stout, rigid, sharp-pointed, 
dull light gray green, with 20- 36 whitish lines or rows of sto- 
mata, and 3-9 (l^) external resjji-ducts. Cones ovoid, 5-7 cm. 
long; seeds narro;-;ly ovoid, acuminate, 15-22 mn. long, very thin- 
walled (0. 1-0.2 mm.), high in carbohydrate content and with mealy 
taste . 

The type specimen i/as collected near Pine Nut Mts., 3E. of 
Gardnerville, Douglas Co., Nevada, in iShk by J. B. Frfeont ( 307 , 
NY) . No varieties of Pinus monophylla are recognized here. How- 
ever, a variation has leaves pairtly or noEtly 2 in a fascicle. 

PINUS EIXJLIS Engelm. pinyon 

Pinus edulis Engelm. in V/isliz., Mem. Totir North. Mex- 88. 
I8if8. 

The commonest species of pinyon -i-ras najned from a specimen col- 
lected in lQh6 by A. Wislizenus (MO) near Santa Fe, lle^-r Mexico. 
It is widespread in foothills and moimtains of Colorado, Utah, 
Arizona, and lle\-T Mexico and has outposts in adjacent states. "Hie 
typical variety, P. edulis va.r. edulis , has 2 leaves in a fasci- 
cle, 2-5 cm. long, stout, rigid, green, with whitish lines of 
stomata on all surfaces, and 2 external dorsal resin-ducts. 
Cones ovoid or subglobose, 3-5 cm. long; seeds oblong, obtuse, 
10-15 inm* long, thin-iralled (0.3-0.4 ran.), high in fat content 
and with oily taste. 

Variation in number of needles in a fascicle has long been 
observed among the pinyons. For example, the t;,'pe specimen and 
plate of Pinus monophylla Torr. & Fr^m. both have rare 2- needle 
fascicles. P. fremontiana Endlicher (S^.m. Conif. I83. 1847) was 
a renaming of P. nonoph:,'lls- ? apparently because the needles were 
thought to be paired and cohering rather than single. P. ed^glis 
var. monophyllus Torr. (in Ives Rpt. Colo. R. pt. 4: 28. i860; 
nom . nud . ; Cebat Mts., J. S. Ngirberr:/ in I858 , US) appsirently \7B.s 
intended to imite both species, though under the later binomial. 
Among those giving additional reasons for combining the two -were 
J. 0. Ne^/berry and Thomas Meehan (Torrey Bot. Club Bui. 12: 50, 
81-82. 1885). 

The legitimate trinomial Pinus monophylla var. edulis M. E. 
Jones (Zoe 2: 25I. I891) ira.s ma,de with the reoarl-: that both 
single and paired leaves were found frequently on the same indi- 
vidual tree. As no basionym i^as cited, the name i/as a nevr 



1968 Little, New pinyon varieties 331 

variety, not a nev combination. 

Tidestrom (Fl. Utah Nev. 53' 1925) used number of resin-ducts 
in identification as follows: " Pinus monoph^.lla is distinguished 
from Pinus edulis by the number of resin ducts in the leaves. In 
the former the number is normally eight (sometimes less), in the 
latter two in each leaf. Tx-ro- leaved forms of Pinus monophylla 
occur in western and southern Utah; these are recognized by three 
or four ducts in each leaf. Occasionally one- leaved forms of 
Pinus edulis are found, but these can readily be distinguished 
from Pinus monophylla by the number of ducts . " 

The 1-leaf or 1-needle variation of Pinus edulis described be- 
lo\^ ims mentioned by me in five publications. Southwestern Trees 
(U.S. Dept. Agr., Agr. Handb. 9: 12. 1950) stated: ''A form with 
the needles single as in singleleaf pinyon but relatively more 
slender and shorter occurs in central Arizona along the lower 
limit of the woodland zone north to Grand Canyon." The other 
references are: Key to southwestern trees (Southwest. Forest 
and Ra.nge Expt. Sta. Res. Rpt. 8: h. I951) . Seminar and study 
tour of Latin- American conifers (Mex. Inst. Nac. Invest. Forest. 
English Ed. No. 1: 90. I962) . Pinyon ( Pinus edulis Engelm.) (in 
Fowells, H. A., comp. Silvics of forest trees of the United 
States. U.S. Dept. Agr., Agr. Handb. 2T1: ^02. I965) . Critch- 
field, William B., and Little, Geographic distribution of the 
pines of the irorld (U.S. Dept. Agr. Misc. Pub. 991: 8. I966) . 

PHTUS EDULIS Engelm. var. FAUJ^X Little, var. nov. pinyon 
Pinus monoph:;"lla var. tenuis Tidestrom in Tidestrom & Kittell, 
Fl. Ariz. Ne^r Mex. 2. 19^1; without La.tin diagnosis. 

A varietate t;'/pica differt foliis solitariis (l in fasciculo, 
non 2), saepe etiam strobilis leviter maioribus 3.5-5-5 cm. 
longis seminibus paulo maioribus lU-lT ram. longis. 

Arbor parva corona aperta rotunda extensa, ramulis tenuibus 
griseis glabris; gemmae c;;'lindricae, acutae, leviter resinosa.e, 
squamis acutis fulvis; folia acerosa, 1 (raro 2) in fasciculo, 
brevia, plerumque "^-k (2.5-5) cm. longa, 1.0-1. i| mm. lata, 
tereta, leviter curva, paulo flexilia, acuminata, integra, sor- 
dido-flavovirentia stomatibus II-I5 seriebus inconspicuis albi- 
dis; ductis resiniferis 2 (3) externalibus dorsalibus; vagina 
paulatim decidua; strobili subterminales, solitarii vel bini, 
fere sessiles, ovoidei, dehiscentes deciduique, parvi, 3.5-5.5 
cm. longi, fulvi; squamae multae, apophysis rhomboidea, crassa 
umbone dorsali piano inermi; semina plura vel multa, gemina vel 
solitaria, obovoidea vel ellipsoidalia, magna, lU-lT mm. longa, 
9-10 ram. lata, 7-8 mm. crassa, base obtuso, apice rotundo, 
apt era, testa tenue (O. 3-0.4 mm.). 

Small resinous tree 4-9 m. high, with trunk I5-3O cm. in diam- 
eter and open rounded spreading cro^-m of nearly horizontal 
branches. Bark dark gray, rough, thick, furrowed into long scaly 



332 P H y T L G 1 A Vol. 17, no. U 

plates. Twins slender, licrht r:ra,v, nmoothish, hairlesc. Buds 
cylindric, acute, slip;htly resinous; bud-scales acute, li^ht 
brovm. Leaves needlelike, 1 (rarely ?) in fascicle. Short, 
mostly 'i-h (?.5-5) cm. lonpj, 1.0-l.U mm. vide, terete, slirjhtly 
curved, slip^htly flexible, acuminate, entire, dull li.^ht erreen 
with 11-15 inconspicuous whitish to\ts of stomata; resin-ducts 2 
(-U) external dorsal; sheath of light brovm membranous scales 
gradually deciduous. 

Male strobili numerous, crowded, elliptic, 6-7 mm. Iohp;, li/^^t 
yellow. Year-old conelets on scaly stalk k mm. lonp;, sub^lobose, 
about 1 cm. lonr;, lij^ht broiim, umbo rhomboidal with week hori- 
zontal keel and short prickle. 

Cones subterrainal, single or paired, almost stalkless, cr/oid, 
dehiscent and deciduous, small 3-5-5'5 cm. loa/;, 2-^ cm. in diam- 
eter when closed and ^^-6 cm. when open, yellav/ brown. Cone-scales 
many, apophysis rhomboidal, thick, keeled, the dorsal umbo flat, 
without prickle, apical and basal cone-scales reduced and sterile. 
Seeds several to many, paired or sin^^le, obovoid or ellipsoidal, 
large, 1^-17 mm. long, 9-10 ram. wide, 7-8 mm. thick, dark broa^n, 
obtuse at base, rounded at apex, wingless, thin-'vra.lled (O.3-O.U 
mm.), high in fat content and with oily taste. 

TYPE COLLECTION, ARIZONA: Gila Co., Tonto National Forest, 
Sierra Ancha Expt. Forest, near Natural Drainage D, Sec. lU, T. 5 
N., R. 13 E., ait. ^4-700 ft., July h, I96I, E. L. Little, Jr. 
18581 (holotj^pe, US; isotj^pes. A, ARIZ, NY, OKL, TEX, UC, UNM, 
USFS) . 

Additional specimens distributed, ARIZONA: Coconino Co., 
Coconino National Forest, 9 mi. SE. of Sedona., Sec. 25, T. I6 N., 
R. 6 E,, alt. 5200 ft., May I5, I968, E. L. Little, Jr. 23002 , 
23003 (seedlings). 

DISTRIBUTION: Mountains at U5OO-55OO (6OOO) ft. altitude in 
central and eastern Arizona. Also local in Grand Canyon, Coco- 
nino Co., and in Florida Mts., Luna Co., Nei7 Mexico. 

New Mexico: Luna Co., Florida Mts. J. S. Find ley Jan. 3I, 
i960 (UNM). 

Arizona: Mountains in central and eastern parts mostly along 
southern slopes of Mogollon Rim and adjacent mountains southward. 
From Oak Creek Can;/-on south and east to upper tributaries of 
Verde, Salt, and Gila Rivers. Coconino Co., Grand Canyon, Oak 
Creek Canyon, etc.; Yavapai Co., near Camp Verde, near Prescott, 
S. to Bradshaw Mts.; Gila Co., near Pine and Payson, Mazatzal 
Mts., Sierra Ancha, Pinal Mts., Apache Mts., etc.; Pinal Co., 
Superstition Mts. and Devils Canyon near Superior; Graham Co., 
Galiuro Hts., Pinaleflo Mts. (Mt. Graham); Greenlee Ct)., mts. N. 
and E. of Clifton. Kaibab, Coconino, Prescott, Tonto, Coronado, 



1968 Little, New pinj'-on varieties 333 

and Apache National Forests. San Carlos Indian Reservation. 

This variety is found mostly southward and at lower altitudes 
than the typical variety. In the lower part of Oak Creek Canyon 
around Sedona it is common. However, it does not form extensive 
woodlands. It is scattered in the Pinyon- Juniper T:.Tie (SAF No. 
239) J associated with Juniperus osteosperma. and J. monosperma, 
also in the chaparral tj'pe of evergreen shrubs such as Que re us 
turbine 11a . 

In 1935 I first observed this 1-needle variation while in 
watershed management research at the Sierra Ancha Experimental 
Forest on the Tonto National Forest, about 30 miles north of 
Globe, Gila Coujity, Arizona. The trees, referred to Pinus mono - 
nhylla by local foresters, ■<.-Tere scattered in the lower part of 
Parker Creek Canyon in the chaparral zone at an altitude of about 
iLYOO ft. Three miles upstream in the s3,me canyon in the oak 
woodland at 58OO ft. were a fei-r trees of P. edulis, the t^/pical 
2- needle variation. 

Soon after beginning work on the pinyon pro.ject about two 
years later, I concluded that this variation belonged with Pinus 
edulis . At that time, being reluctant to distinguish minor vari- 
ations by name, I designated this one as '' Pinus edulis Enr^elm. 1- 
leaf form.' In 1939> I so annotated specimens in several eastern 
herbaria, using a rubber stamp. Likewise, in 19^+1, I checked 
specimens in California herbaria. 

VTiile in Washington, D.C., in 1939? L mentioned this variation 
of Pinus edulis to the late Ivar Tidestrom, then retired and 
working on his flora of Arizona and Nei-j- Mexico. However, he 
changed from his earlier interpretation and named it Pinus mono - 
phylla var. tenuis Tidestrom (in Tidestrom and Kittell, Fl. Ariz. 
New Mex. ?. 19^-^-1: without Latin diagnosis). His brief English 
description >/as: "Distinguished from the type by its more 
slender leaves and the number of resinducts, the latter usually 
2, marginal. VJest-central Arizona and westirard." I cited this 
variet;; as a sjTiom-m of P. edulis in the Forest Service Check 
List (U.S. Dept. Agr., Agr. Handb. kl: 26k. 1953). 

Kearney and Peebles (Flowering Plants Ferns Ariz. 6I. 19^2; 
Ariz. Fl. 52. 1951) also were aira.re of this variation and re- 
marked ujider Pinus monophylla ; ''As it occurs in Arizona, this 
pine scarcely differs from the ordinary pin\'on (Pinus edulis) 
except in its solitary leaves, and may be only a variant of that 
species. Presumably tipical P. monophylla , in California and 
Nevada, has thicker a.nd more rigid leaves and larger cones than 
the Arizona form.'' 

Trees and specimens of this nei-j variety generally have been 
referred to Pinus monophylla on the basis of needle number. The 
varietal epithet fallax , deceptive, refers to this character. 



33lt PHYTOLOGIA Vol. 17, no. U 

However, herbarium specimenc of the new variety are readily die- 
tinp;uiched by the slender narrow r;reen needles only l.O-lJi im. 
wide. P. monophylla. has relatively stout, rir;id, sharp-pointed 
needles var^inr; r';reetly in lern^th, P-6 cm. lonp;, 1.5-2 ™n- vide, 
straight or sli'3htly curved, and a different color, Rray preen 
(grayer than in P. edulis ) or sometimee pale olive p;reen. 

There are also slight differences in needle anatonr/ in cross 
section. Along with smaller diameter, the new variety has fewer 
(11-15) rows of stomata, usually thinner h^.TJOdermis, and fewer 
resin-ducts, lisually 2 {-k) , as observed by Tidestrom (Fl. Utah 
Nev. 53. 1925). Pinus monophylla has 2O-36 rows of stomata, 
hj'podermis of I-3 layers of thick-•^^ra,lled cells, and 3-9 (16) 
resin-ducts. Of course, differences in needle anatomy are useful 
to the extent they are correlated with other morphological char- 
acters. For example, number of resin-ducts alone would not merit 
separation of populations into species. 

Seeds of Pinus monophylla are easily distinguished from seeds 
of P. edulis (including the ne^-r variety) by their larger size, 
15-2 2 mm. long, thinner shells or seed coats (0.1-0.2 ram.), and 
chemical composition and taste. The shape is slightly different, 
narroiiTl;/ ovoid, relatively less broad and gradually taperiag to 
base. The seeds are so thin-shelled that the;/ can be crushed and 
cracked with the fingers, between thumb and forefinger, ivhile 
seeds of P. edulis must be cracked with the teeth. 

Pinyon nuts of Pinus edulis , including the nei7 variety, have 
an oily flavor, while those of P. monophylla are mealy. These 
taste differences have been confirmed by chemical analyses, for 
example, by C. \h Botkin and L. B. Shires (The composition and 
value of pinon nuts. N. Mex. Agr. Expt. Sta. Bull. 2^k, Ik pp., 
illus. 19^8). Seeds of P. edulis average more than 60 percent 
fat, less than 20 percent carbohydrate, and less than I5 percent 
protein. Seeds of P. monophylla average less than 25 percent 
fat, more than 5O percent carbohydrate, and less than 10 percent 
protein. Most persons prefer the oil;- flavor over mealy. Nearly 
all the pinyon nuts or Indian nuts sold commercially belong to P. 
edulis. However, nuts of P. monophylla are harvested and con- 
sumed locall;-. Botkin and Shires reported a chemical analysis of 
one sample of P. edulis 1-leaf variety from southeast of Kingman, 
Ariz., oily lil^e the tj-pical variety but with slightly higher fat 
content (65.66^^). 

The two varieties of Pinus edulis have separate natural ranges 
and altitudinal zones but meet in a fev^r places. Plants of the 
new, 1- needle variety bear 2-needle fascicles rarely. Plants of 
the tj/pical, 2-needle variety produce 3-n6e<31e fascicles rarely 
but not 1-needle fascicles. 

Fiu'ther field work in early autvmin to collect specimens with 
filled seeds would be desirable in northi^re stern Arizona and 



1968 Little, New pinyon varieties 33? 

southwestern Utah. In those regions, trees with both 1- and 2- 
needle fascicles have been found. Pinus monophylLa is known in 
Arizona only from the northwest comer, in Virgin Mts. and Huaia- 
pai Mts. and vicinity in Mohave County. Easti^reird, for example, 
near Peach Springs, this species meets and may intergrade mth 
the tj'pical 2-needle variety of Pinus e dulis . Also, P. edulis 
var. fallax may extend northi^rest to Hualepei Mts. 

Pinus edulis ybx. fallaLX has no outstanding characters of 
economic value. Its nuts are not of commercial importance be- 
cause of poor seed crops. No bumper pinyon nut crops have been 
reported within its range. Trees at low altitudes seldom mature 
seeds in quantities, and many full-size seeds are empty (blighted 
or bla.sted) . This variety is located in a \-7armer climate than 
the tj'pical variety and sheds pollen up to a month earlier. Be- 
cause of the longer growing season, it might grow less slowly. 
This variety is hardy in a semiarid ^Tarm temperate climate and is 
classed doubtfully in Zone J, while the typical variety is hardy 
in Zone h. Like the latter, the ne^^- variety may have possibili- 
ties for planting for shelterbelts, timber, pulpwood, erosion 
control, wildlife cover and food, and Christmas trees- In future 
tree breeding programs for pinyon nut production, this large-seed 
variety of a lower altitudinal zone should be tested. 

Number of needles in a fascicle has been reduced to 1 indepen- 
dently in 3 species of pinyon, P. monophylla , P. edulis var. fal- 
laix, and P. nelsonii Shaw (Little in Seminar and study toiir of 
Latin- American conifers. Mex. Inst. Nac. Invest. Forest. English 
Ed. No. 1: 90. 1962). In P. nelsonii of Mexico 3 ver^r slender, 
weak needles cohere functionally/ as 1. Reduction of needle num- 
ber in a fascicle occurs in semiarid regions with low rainfall 
and may be assumed to be an adaptation for reduced leaf sirrface 
with less transpiration. These illustrations with 1-needle fas- 
cicles are interesting examples of reversible evolution from 
alternate leaves to whorled and back to alternate. The spur 
shoot in Pinus with mostly 2-5 needles may have developed from an 
ancestral t^'pe like Cedrus or Larix with needles both alternate 
on leading twigs and whorled on spur shoots. The 1-needle pin- 
yons approach the original, primitive tj'pe with alternate leaves 
as in several related genera such as Picea, Tsuga , and Abies . 

PINUS CEMBROinES Zucc. Mexican pinyon 

Pinus cembroides Zucc, K. Bayer. Akad. Wiss. Munchen, 

Abhandl. Math.-Phys. 1: 392. I832; Flora [Jena] I5 (2), 
Beibl. 93, 1832. 

Mexican pinyon ^'/as named from a specimen collected by Wilhelm 
F. Kar-^^rinski in central Mexico, apparently near Zinapdn, Hidalgo 
(Endlicher, Sjmops, Conif. I83. 18^T) . The tj^pical variety of 
this species is widely distributed in mountains of northern and 
central Mexico and extends northward into the United States only 
in Trans-Pecos Texas. It has fascicles of ^ slender leaves 



336 P H Y T L G I A Vol. 17, no. U 

tnoctly 3-5-T' (^m- Ion,, \rith ?-!? liner, of 'forr.al ctonati, aloe 
thick-vralled needc (O. 5-1.0 ran.) 

In the restricted sense, Finns cembroides has tvo additional 
varieties. P. cenbroides var. remotg. Little (V/rir^htia 3: 133- 
1955), Texas pinyon, rare and local in the EdiTardc Plateau and 
Trans-Pecos Texas, is characterized by leaves mor,tly ? in a fas- 
cicle (vii:h dorsal stomata) and by thin-iralled seeds (O.l-O.^ 
mm.). That variety mii^ht be of hybrid ori,3in betveen that 
species and P. edulis . The other variety is described belor/. 

PINUS CH^IBROIEES Zucc. var. BICOLOR Little, var. no-/. 

Kiexican pinyon 

A varietate tj^pica differt foliis bicoloribus sine stotnatibus 
Qorsalibus, sutjerficie dorsali atro-virenti et supejTf iciebus ven- 
tralibus albis atque -laucis: etian strooilis miaoribus 2-2 cm. 
lonrjis seminibus paucioribus minoribus (8) IO-I3 mm. lon^is. 

Arbor par^/a corona, aperta extensa irreyulari vel rotunda, 
ramulis tenuibus .'jriseis r^labris; f:eminae cylindricae, acutae, 
leviter resinosae, squamis elon2;ato-ac'jminatis, apice atro-rubro; 
folia acerosa, 3 (raro 2 vel h) in fasciculo, brevia, plerur.que 
2.^-k (2-5) cm. lonf>a, 0.8-1.0 mm. lata, tenua, flexilia, acumin- 
ata, Integra, bicoloria sine stoms.tibuj5 dorsalibios, stomatibus 
ventralibus 2-3 seriebus, duct is res inif eras 2 externalibus dor- 
salibus; vagina paulatim decidua; strobili subterminales, soli- 
tarii vel bini, brevi-pedunculati, subjlobosi, dehiscentes et 
decidui, perparvi, 2-3 cm. longi, fulvi; squamae plurae vel mul- 
tae, apophysis rhomboidea, crassa ujabone dorsali piano inermi; 
-semina pauca, solitaria vel ^emina, ellipsoidalia, parva, (3) 10- 
13 ram. longa, 7-10 ram. lata, 7-8 mm. crassa, base atque apice ro- 
tundo, aptera, testa crassa (0.7-1.0 ran.). 

Differs from the t;;T)ical variety in the tvo-colored leaves 
without dorsal stomata, dorsal sujrface dark r-,reen, and ventral 
surfaces vhite and glaucous; also in the smaller cones- 2-3 cm. 
long with feiN'er, smaller seeds (8) IO-I3 mm. lon^;. 

Small resinous tree 4-9 (I5) ri. high, vith trunA 12-^5 cm. in 
diameter and open irregular or rounded spreading cravm of nearly 
horizontal branches. Barl-: blackish or dark gray, rough, thick, 
furroved into long scaly ridges or plates, often exposing orange 
brovTi or reddish brcn.m inner bark, on large branches gray and 
smooth. T'.vTigs slender, light gray, smoothish, hairless. Buds 
cylindric, acute, slightly resinous; bud-scales long acuminate, 
light bro'^im, dark red at apex. Leaves needlelike, 3 (rarely 2 or 
h) in fascicle, short, mostly 2.5-^ (2-5) cm. long, 0.3-1.0 ram. 
wide, slightly spreading, slender, straight, flexible, acuminate, 
entire, dorsal surface dark green without dorsal stomata, ventral 
surfaces white and glaucous vith 2-3 inconspicuo'jis rows of sto- 
mata; resin-ducts 2 external dorsal; with sheath of light brown 



1968 Little, Hew pinyon varieties 337 

membranous scales gradually deciduous. 

Year-old conelets on scaly stalk 5-8 mm. long, subglobose, 8- 
10 mm. long, light ■braim, umbo rhomboidal with weak horizontal 
keel and no prickle. 

Cones subterminal, single or paired, short- stalled, subglo- 
bose, dehiscent and deciduous, very small, 2-3 cm. long, 2-2.5 
cm. in diameter when closed and 3-^ cm. when open, yellow brcnm, 
often slightly reddish tinged. Cone-scales several to many, the 
apophysis rhomboidal, thiclc, keeled, the dorsal umbo flat, with- 
out prickle, apical and basal cone-scales reduced a.nd sterile. 
Seeds few (sometimes only l), single or paired, ellipsoidal, 
small, (8) 10-13 mm. long, 7-10 mm. wide, T-8 mm. thick, dark 
broi-m, rounded at base and apex, w3.ngless, thick-Tra.lled (0. 7-1.0 
mm.), edible, high in fat content and with oily taste. 

TYPE COLLECTION, ARIZONA: Santa Cruz Co., Coronado National 
Forest, Santa Rita Mts., Madera Canyon, Sec. 13, T. 20 S., R. ih 
E., alt. 6500 ft.. May 20, I968, E. L. Little, Jr. 23OII (female 
plant, cones and seeds under several trees; holotj'i)e, US; iso- 
types. A, ARIZ, NY, OKL, TEX, UC, UIM, USFS), 23010 (male plant), 
23012 (seedlings). 

DISTRIBUTION: Mountains at 5OOO-7OOO (8OOO) ft. altitude in 
southwestern Ner-r Mexico, southeastern Arizona, northeastern 
Sonora, and western Chihuahija. Also local in mountains of Coa- 
huila, Nuevo Le6n, Tamaulipas, San Luis Potosi, and Zacatecas. 

Neiv Mexico: Mountains of southwestern corner; Hidalgo Co., 
Peloncillo Mts. on Coronado National Forest, Animas Mts., Big 
Hatchet Mts.; Grant Co., mts. near Pinos Altos, Mule Creek, and 
Burro Mts. all on Gila National Forest. 

Arizona: Mountains of southeastern part including nearly all 
divisions of Coronado National Forest; Greenlee Co., mts. N. and 
E. of Clifton on Apache National Forest; Graham Co., Pinaleflo 
Mts. (Mt. Graham); Pima Co., Santa Catalina Mts. (Mt. Lemmon) , 
Rincon Mts., Baboquivari Mts., Coyote Mts.; Santa Cruz Co., Santa 
Rita Mts., Ruby Mts. NV/. of Nogales, Patagonia Mts.; Cochise Co., 
Huachuca Mts., Mule Mts., V/hetstone Mts., Chiricahua Mts., Pelon- 
cillo Mts. Probably on other peaks reaching 7000 ft. altitude. 

Sonora: Mountains in northeastern part and along interna- 
tional boundary west toiNfard Nogales. 

Chihuahua.: San Luis Mts. in northwestern corner along inter- 
national boundary. Apparently less common in Sierra Madre Occi- 
dental in v^estern Chihuahua than typical variety; recorded from 
near El Vergel ( Little 18919 ) in southwestern part, also an 
intermediate specimen with 2 rows of dorsal stomata en leaves 
collected 25 mi. W. of La Junta (Little 18907) . 



338 PHYTOLOGIA Vol. 17, no. U 

Specimens of Pinus cembroldes without dorcal stomata were 
noted from other statec of Mexico, as follovc: 

TAI^ULIPAS: 3 mi. N. of Miquihuana, July 11, 19i«9, Stanford , 
Lauber, Taylor 2397 (UTC) . 

COAHUILA: Mt. Jimulco, 13 km. E. of Jinulco, alt. 3100 m., 
June 29, I9U1, Stanford, Retherford, North craft 110 (CAL, UTC) : 
Sierra de la Madera, Caflon del A[^ua, Mun. de Cuatro Ci^nef^aE, 
Sept. 8, 1939, C. H. Muller 3229 (CAL). 

SAN LUIS POTOSI: I3 mi. SV/. of San Luis Potosl, alt. 78OO- 
8000 ft., July 28, 1958, R. M. Straw & M. Forman IU38 (US) . 

ZACATECAS: Concepci6n del Oro, Sierra M^dre Oriental, alt. 
2500-2700 m., July 18-19, 193^> F. W. Pennell 17^33 (US) ; Ja^uey, 
Cedros, alt. 8OOO ft.. May I908, F. E. Lloyd Ch (US, CAL). 

NUEVO LE6N: Sierra de la Cebolla, Mun. de Montemorelos y 
Rayones, Aug. 21, 1939, C. H. Muller 2916 (CAL). 

All trees and specimens of Pinus cembroides Zucc. ( sens . 
strict . ) native in southeastern Arizona and southvrestem New Mex- 
ico are referred here to var. bicolor , having tvo- colored leaves 
without dorsal stomata. This nei^ variety of Mexican pinyon is 
not abundant within its range and does not form extensive A-rood- 
lands as does Pinus edulis north■^^^ard . Instead, Mexican pinyon is 
scattered in evergreen woodlands of junipers and oaks in the Pin- 
yon- Juniper Type (SAF No. 239) and Interior Live Oak Type (SAF 
No. 2i+l) . It is less common than the associated tree species: 
Juniperus deppeana , Quercus emoryi , Q. h?/T3cleucoides , Q. reticu - 
lata , Q. arizonica , Pinus ponderosa var. arizonica. 

Jack McCormick and John V7. Andresen (A subdioecious population 
of Pinus cembroides in southeast Arizona. Ohio Jour. Sci. 63: 
159-163. 1963) studied Pinus cembroides in Chiricahua Mountains, 
Cochise County, Arizona, pinyon trees included here within the 
nei^ variety. They reported that onl^/^ about 2 per cent of the 
individuals \rere monoecious and the others either male or female. 
When the type collection for this ne\<r variety ira.s made, most 
trees were readily separable into male or female, and a second 
collection ■\ra,s from a male tree. Madera Canyon, Santa Rita Moun- 
tains, is about 90 miles ■(•rest of Chiricahua Mountains. In Madera 
Canyon pinyon trees are scattered ajid not abundant. 

The needle anatomy has been described and illustrated under 
Pinus cembroides by \h M. Harlow (The identification of the pines 
of the United States, native and introduced, by needle structure. 
N.Y. State Coll. Forestry Syracuse Univ. Tech. Pub. 32, 21 pp., 
illus. 1931). His description of the position of stomata, "ven- 
tral, rarely if ever dorsal as well," clearly refers to the new 
variety . 



1968 Little, New pinyon varieties 339 

Absence of dorsal stoina,ta in leaves \ra.s used to distinguish 
Pinus cembroides from P. edulis by Max^'^ell T. Masters ( A general 
view of the genus Pinus . Linn. Soc. London Jour. Bot. 35: 586, 
588. 1904). Hoi-rever, George Russell Shaw (The pines of Mexico. 
Arnold Arboretum Pubs. 1: 6. 1909) added that this character 
fails in Mexican specimens. Obviously Masters was observing this 
ne^.r variety from Arizona. 

Pinus cembroides var. bicolor is easily recognized by the 
slender leaves in a fascicle of contrasting colors, the outer 
surfaces dark green without rows of stomata and the inner sur- 
faces white. The two-colored leaves are conspicuous in herbarium 
specimens though less so in those dried by artificial heat. The 
whitish lines or rows of stomata are present on the dorsal leaf 
surfaces of other pinyons with two exceptions of limited distri- 
bution. P. quadrifolia Pari., Parr;^"- pinyon, of southern Caxi- 
fornia and northern Lo^^rer California, has ti/o- colored leaves but 
stout and mostly 4 in a fascicle, rarely with dorsal stomata. P. 
culminicola Andresen & Beaman, kno-im only from Cerro Potosi, 
Nuevo Le6n, has 5 leaves in a fascicle. 

Pinus cembroides var. bicolor , like P. edulis , could become 
a popular Christmas tree in the Southwest. The two-color foliage 
of slender dark green and ■\^;hite needles is especially attractive. 
Nearly all plants of this variety in the United States are within 
the national forests. Accordingly, the U.S. Forest Service would 
supervise the harvesting of Christmas trees on a sustained yield 
basis. 

The nei-7 variety of Mexican pinyon is of no commercial inrpor- 
tance for pinyon nuts because of its scattered occurrence, gener- 
ally poor seed production with no bumper crops, and pajrticularly 
the small thick-shelled seeds. The wood should be suitable for 
the same uses as in other pinyon species, for example, mine tim- 
bers and pulpwood. The trees should be hardy in semiarid ■^^ra,rm 
temperate regions and could be planted experimentally in marginal 
areas near the latter limits of trees. Hwrever, groijth probably 
would be very slow. The new variety of Mexican pinyon is adapted 
to a mild winter climate and in hardiness is classed doubtfully 
in Zone T, while the typical variety of Pinus edulis is in Zonei^-. 

Pinyon trees of three different taxonomic groups were col- 
lected by me in 1956 growing together where their ranges meet in 
central Greenlee County, Arizona. The easily accessible locality 
is along U.S. High-ira.y 666 about lU miles north of Clifton on the 
Apache National Forest (T. 3 S., R. 29 E.). Here, side by side, 
in a shrub tj-pe at 6200 ft. altitude, were the following: Pinus 
edulis var. edulis to 20 ft. high and 8 in. d.b.h.,near its 
southwestern border and lo-\rer altitudinal limit; Pinus edulis 
var. fallax near its southeastern limit; and Pinu^ cembroides 
var. bicolor at its northernmost limit. There were no intermed- 
iate plants except that those of the 1-leaf variety had some 2- 



3U0 PHYTOLOGIA Vol. 17, no. h 

leaf fascicles. 

Trees of Pinui; edulls var. edulJs and P. cembroldes var. bi- 
color grov toeether also aloniz /urizona Ctate Hi^lTv/ay 78 in moun- 
tains about 15 miles east of Clifton on Apache National Forest 
(T. k S., R. 32 E.). Even though branches of both cpeciec were 
touching; one another, I saw no intermediate or hybrid trees. 

PinuG cembroides var. cembroides may interp;rade in Trans-Pecos 
Texas with P. edulis var. edulic, vhich extends southeast into 
tvo localities there. Also, P. cembroides var. remota of east- 
ward range into the Edi-zards Plateau might be partly of hybrid 
origin or possibly an ancestral intermediate type. Jack McCor- 
mick and John v;. Andresen (Ohio Jour. Sci. G3: l62. I963) men- 
tioned that tferion T. Hall had observed in central !Ie\/ Mexico, 
from the Sacramento Mountains north^ra.rd to the Gandia Range, 
trees considered to be introgressants of the two species. 
Recently George G. Fogg has reported hybridization in the Cem- 
broid pines (Ecol. Soc. .^mer. Bui. h<^: 71. I968) . 

Early in my field work with pinyons in Arizona and Nei^r Mexico, 
I observed the striking characters separating Pinus edulis and P. 
cembroides there. In the meantime I have studied ?. cembroides 
in Texas and Mexico. After field work in April 19^3, I r&'ri&'.red 
the pinyons of Texas (VJrightia 6: iBl-lBT- I966) . '■^niile vrorking 
in Mexico in 19^5 j I observed P. cembroides near the type local- 
ity in Hidalgo. In September-October I96O I made extensive col- 
lections of Mexican pines as the United States representative 
with the Seminar and Study Tour of Latin- American Conifers, lander 
the Food and Agriculture Organization of the United Nations (I'ex. 
Inst. Nac. Invest. Forest. English Ed. No. 1, 209 VV-> illus. 
1962) . Again in March I963 1 collected pinyons and other pines 
in northern and central Mexico. 

The differences between Pinus edulis and P. cembroides in Ari- 
zona and New Mexico were suramarized 30 years ago in a 3-page mim- 
eographed Resea,rch Note (Little, Mexican pinon (Pinus cembroides). 
Southwestern Forest and Range Expt. Sta. Research Note No. kj, 3 
pp. 1938). The essential details, which refer to P. cembroides 
var. bicolor , are quoted below. 

"While the academic question of ran];:ing variations as separate 
species or merely varieties is unimportant in practical forestry, 
it so happens that seed characters of pinons, which several re- 
cent taxonomists have overlooked, aj^e of great economic impor- 
tance. Actually, Pinus cembroides and Pinus edulis, as repre- 
sented in the United States, are so distinct that a single nee- 
dle, a single winter bud, a single seed, a single immature cone, 
and in most cases a single mature cone can be assigned with cer- 
tainty to one of the two species. 'Note added in I968: This 
statement does not hold for some Texas specimens of other var- 
ieties of P. cembroides. j 



1968 



Little, New pinyon varieties 



3ia 



"Differences between the two species which hold true for trees 
growing in Arizona and New Mexico are tabulated below. The char- 
acters of greatest taxonomic value are indicated by asterisks (*) . 



CHARACTER 

Needles 

*Number in cluster 

Shape 
*Width 
*Dorsal stomata 



PINUS CIMBROinES 
; var . bicolor J 

usually 3 

slender 

less than 1 mm 

absent 



Color of outer siorface daxk green 
Color of inner surface white all over 



PINUS EDULIS 



2, rarely 1 or 3 
stout 

greater than 1 mm 
4 to 6 longitudi- 
nal rows 
light green 
more or less white 



Winter buds 

*Shape of scales long tapering point short-pointed 
Immature (year-old) cones 

*Stalk long, 5 to 8 mm long short, 2 to 3 ™i^ 

long 

Length 8 to 10 ram 8 to l^J- mm 

Prickle on cone scale inconspicuous nearly l/2 mm long 



Mature cones 

Length 

Shape 
Seeds (nuts) 

Length 

Thickness of shell 
*Strength of shell 

Economic importance 
in United States 



20 to 25 mm 
nearly spherical 

8 to 13 mm 
2/3 to 1 mm 
cannot be cracked 
with the teeth 



25 to 50 ram 
longer than broad 

10 to 15 ram 
less than I/3 mm 
easily cracked 
with the teeth 

high 



"Other distinguishing characteristics have been proposed. 
There are a fev;^ microscopic differences in wood anatomy. Sud- 
worth (U.S. Dept. Agr. Buli.^6o) mentions the different numbers 
of cotyledons or seedleaves, 8 to I5 in Pinus cembroides and 7 to 
10 (the author finds 6 to 12) in Pinus edulis , but these numbers 
vary widely. The author is unable to detect differences in bark. 

"The two species may easily be distin-guished by testing the 
nuts with the teeth. The most practicable vegetative character- 
istic in the field is the presence or absence of the white longi- 
tudinal rows of stoms,ta on the dorsal or outer surface of the 
needles. This character is less conspicuous in herbarium speci- 
mens because the color fades, and, according to Shaw (The Pines 
of Mexico), does not hold in specimens from Mexico. In the field 
the slender, spreading needles of Pinus cembroides present a 
color contrast of dark green and white, while the stouter, light 
green needles of Pinus edulis, being mostly in tvos, do not ex- 
pose their whitish inner surfaces as much. . . . 



3U2 P H Y T L G I A Vol. 17, no. h 

"In a fe\/ placec in Arizona and Ue\r Mexico the ranges of Pinuc 
cerobroides and Pinus edulic meet, but they do not overlap much. 
Treec of the two species throwing side by side are not distin- 
guishable at a distance but may readily be identified when the 
characters previously mentioned are examined. No intergrades or 
hybrids have been observed. It would be interectia^ to kno-^ whe- 
ther these closely related species shed their pollen at the same 
time. 

"Although it is reported that they are gathered and eaten on a 
large scale in Mexico, nuts of Mexican pinons are of no commer- 
cial value in the United States at present because of their hard 
shells. Mr. Karl Pitschner, of the Albuquerque Food Producte 
Co., reports that in fairly well dried nuts the shells of Pinus 
cembroides make up 65 to 6? percent of total weight and shells of 
Pinus edulis only 48 percent. The abundance of Pinus cembroides 
is insignificant in compari; .1 with Pinus edulis . Nut crops of 
Mexican pinon trees in the United States apparently are light or 
frequently failures. Only a fe\T nuts are contained in one cone, 
and many nuts are empty. 

"Thus, Pinus cembroides and Pinus edulis in the United States 
are sufficiently distinct to be regarded as separate species, 
even under a conservative interpretation. Additional taxonomic 
study of Mexican trees and specimens, including examination of 
the type specimen of Pinus cembroides , is needed to test the con- 
stancy of the characteristics enumerated here." 

KEY TO ARIZONA PUFYONS 

The following key to the i^ tsixonomic groups of pinyons (nut 
pines) native in Arizona will serve as a summary: 

Needles 1 (rarely 2) in fascicle 

Needles stout (I.5-2 mm. wide), light gray green; seeds 
very thin-A-ralled, with mealy taste; northwestern 
Arizona Pinus raonoiDhvlla 

Needles slender (1-1.4 ram. wide), light green; seeds 
thin-walled, with oily taste; central and eastern 

Arizona Pinus edulis var. fallax 

Needles 2 or 3 in fascicle 

Needles mostly 2 in fascicle, stout (more than 1 ram. wide), 
green, with k-6 rows of dorsal stomata; seeds thin- 
walled; northern and eastern Arizona 

Pinus edulis var. edulis 

Needles mostly 3 in fascicle, slender (less than 1 mm. wide) , 
two-colored, dorsal surface dark green without dorsal 
stomata, ventral surfaces white; seeds thick--(-7alled; 
southeastern Arizona .... Pinus cembroides var. bicolor 

Forest Service, United States Department of Agriculture, 
Washington, D. C. 20250. 



THE ERUPTION IN HIIAKA CRATER, ISLAND OF HAWAII 
Otto fi: Isa Degener 



Hiiaka*- Crater, a minor pit crater in Hawaii Volcanoes National 
Park and named in honor of the sister of the Hawaiian Volcano God- 
dess Pele, showed activity at 6.U5 A^., August 22, 1968, the first 
time within the memor;,'' of man. Clouds of acrid fumes rose into the 
air to be carried by the trade Yd.nd in a southwesterly direction. 
Though the activity culminated in a 75-foot lava fountain, and a 
pool of lava 300 feet in circumference accumulated in the crater 
before draining away underground, the area became quiescent before 
noon of the same day. 

On August 2ii we drove along the Chain-of-Craters Road, turning 
into the side road leading to Ainahou Ranch. For a distance of 
about a mile along this road or up to one and a half miles leeward 
as the crow flies from Hiiaka Crater, we noted the following ccnimon 
plants affected by the fumes: Sadleria cyatheoides (endemic), 
Lycopodium cernutim var. crassifolium (native), Arundina bambusi- 
folia (naturalized weed), Santalum panic ulatum (endemic), Osteo- 
meles anthyllidifolia (native), Dodonaea viscosa s.l, (native), 
Styphelia tameiameiae (endemic), Vaccinium reticulatum (endemic), 
Buddleja asiatica (naturalized weed), Pluchea odorata (naturalized 
weed), and Railliardia ciliolata var. laxi flora (endemic). Their 
leaves showed signs of wilting andor yellovdng and often death. 
What impressed us was that the endemic Lletrosideros , often in- 
correctly considered conspecific with the New Zealand M, collina, 
showed no damage frcm the fumes at all, apparently having devel- 
oped an efficient immunity over the ages. 



*Incorrectly spelled "Heake" on the United States Geological 
Survey map, edition of 1933. 



3U3 



NOTES OK NEW AND JCTEWOkTHY PUNTS. LI 
Harold li, Itoldenke 



DURWJTA CAJAilARCENSiS Koldenke, sp. nov. 

Arbuscula valde armata; ramis rigidis tetragonis glabris niti- 
dis parce piinctato-lenticillatis, angulis acutis saepe suunargina- 
tis; ranulis abbreviatis spiniformibus 2 — 5 cm. longis ritridissi- 
mis apice argutis; laminis decussato-oppositis coriaceis ellipticis 
integris, apice acutis, ad basin longe attenuatis, juventute subtus 
pubesc aitibus, maturitate utrinque glabris; corollis albis . 

Very spiny shrub; branches nedium-slender, rigid, acutely tet- 
ragonal, glabrous, shiny, s^' . sely lenticillate with nore or less 
punctiform lenticels, the angles often submargined; Vivigs abbrev- 
iated, decussate-opposite, verj' rigid, 2 — S cm. long, composed of 
several nodes and intemodes, very sharply aculeate at the apex; 
leaves petiolate, decussate-opposite; petioles about 5 nun. long; 
leaf-blades coriaceous when mature, el3_Lptic, 3 — ? cm. long, 
1 — 1,7 cm. Td.de, acute at the apex, entire on the narQins, long- 
attenuate at the base, pubescent beneath when iijiiature, glabrous 
and shiny on both surfaces \fhen mature; flowers apparently a 
single pair on filiform peduncles about p mn. long at the apex 
of a very much abbreviated filiform twiglet in the axil of the 
leaf subtending a spine, this twiglet and peduncles puberulent; 
pedicels less than 1 mn. long, filiform, puberulent; calyx ob- 
conic, about h mm. long, 2 mm. wide at the top, puberulent; 
corolla white, its tube equalling i-he calyx, its limb spreading 
or reflexed. 

The type of this species was collected by A. Sag^stegiii A. 
( no. 638I|. ) on a shrubby slope, at an altitude of 2^00 netei^, at 
Guzmango, in the province of Contimaza, Cajamarca, Peru, on 
October $, 196?, and is deposited in my personal herbarium at 
Plainfield, New Jersey. 

VERBEMA OCCULTA f . AURAIITIACA Koldenke, f. nov. 

Haec forma a forma typica speciei corollis avirantiacis rece- 
dit. 

This form differs from the typical form of the species in 
having the corollas orange-red according to notes by the collector 
on the label of the type specimen. 

The type of the form was collected by ri. Angulo ( no. 1383) in 
a cultivated field at Humachuco, in the province of Humachuco, La 
Libertad, Peru, at an altitude of 2170 meters, on July 16, 1951, 
and is deposited in n;.^ personal herbarium at Plainfield, New Jer- 
sey. The tern actually used by the collector to describe the 
color of the corollas is "naranjadas". The tj'pical forn of the 
species is described as having purple corollas, and there is an 
albino form knovm with wiiite corollas , 

3kk 



BOOK REVIEWS 
Alma L. Moldenke 



•THE ELEMENTS OF CYTOGENETICS" by G. B. Wilson, 120 pp., illus., 
Reinhold Book Corporation, New York 10022, Amsterdaa & Lon- 
don. 1968. $2.25. 

This paper-back is the eighth in the series edited by Peter 
Gray on SELECTED TOPICS IN MODERN BIOLCGT, and like the others 
provides the beginning student vath a clearcut, interesting and 
intelligent survey of the field. It develops excellently basic 
principles, chrcmosome distributions in mitosis and meiosis or 
chromosome constitution, analysis, sex determination, and 
mechanics . 

There are included a useful glossary, a good bibliography, a 
needed index, and very helpful illustrations. 

This book will have its greatest use among college freshmen. 
It should also be used as enrichment for brighter high school 
biology students and for the interested layman. 

"FLORA NEOTROPICA" Monograph I "SWARTZIA" by Richard S. Cowan, 
227 pp., illus., Kafner Publishing Company, London & New 
York 10003. 1968. $13.00. 

The Organization for Flora Neotropica has just inaugurated 
this journal as a vehicle for monographic taxonomic accounts of 
plants growing spontaneously within the '»7estern Hemisphere 
tropics. The contributions will have geographii'", ecologic, 
cytologic, anatomic, chemical and economic dataj they will be 
organized in similar formats with bibliographies, citation of 
specimens and indexes. The executive director is Dr. Bassett 
Maguire who is at the New Yoric Botanical Garden. 

This first issue contains a very carefully developed mono- 
graphic study of the legume genus Swart zi a of this area, 
omitting Bocoa , which may be a distinct genus. It is the 
thorough work of a well experienced field botanist and system- 
atist. There are clear geographic distribution maps, excellent 
drawings, a list of collectors with their collections, a numer- 
ical list of 127 species and 67 varieties, an exclusion list, 
and a correction sheet, all in clear print, 

"TAXONOMY OF ALISRICAK SPECIES OF LII^DEN ( Tilia )" by George 

Neville Jones, 156 pp., illus., Illinois Biological Mono- 
graphs No. y^ ^ University of Illinois Press, Urbana, Illin- 
ois, paperbound. I968. 50 sh 6 d. or $5.95* 

Even though this study is the thorough one of an experienced 
teacher/field man/taxonooiist, the introduction and many descrip- 

3l;5 



3li6 PHYTOLOGIA Vol. 17, no, U 

tive items make interestinp and easily comprehended reading. 
Like the study above, it is well provided with a few geographic 
maps, specimen photographs, specimen citations, list of collec- 
tors, and index, 

"TJE GENIUS PlinJS" hy N. T. Hirov, viii Sc 602 pp., lllus., Ronald 
Press, New York 10016. I967. JlS^.OO. 

This book represents a thorough and somc-That monographic study 
through interesting facts and easy writing style of the economic- 
ally important and conspicuous pine genus. The topics covered are: 
history, paloobotanor, geography (a specialty of the author), 
genetics, morphology, reproduction, physiological ecology, bio- 
chemistry (omitting limber studies but including many of the 
author's studies), classical and modern taxonomy (omitting the 
citation of specimens and still using the sections H^lo:xylon 
and Di ploxylon ) , and an evaluating summary, 

Pinus dates back to the Mesozoic, has become a leirger group 
since glaciation even with man's onslaught considered, is well 
established throughout the North Temperate Zone, has spread ef- 
fectively into subtropical areas, and reproduces well and freely 
since all species have the same number and kind of chronosocies . 

This book is botanically needed because much has been learned 
(and published in many isolated places) since Shaw's monograph 
of 19lli and P^lger's treatment in the Engler i Prantl of 1926. 
Literature is cited at the end of chapters, diagrams and fine 
photographs are included, and excellent species distribution and 
chemical distribution maps are presented. The print is easilj' 
read. The index is full, 

"A MAIIUAL OF PENICILLIA" by Kenneth B. Raper and Charles Then, 
ix & 87^ pp., illus., copjTighted 19U2 and reprinted hy 
arrangement, Hafner Publishing Company, London Sc Nevr York 
10003. 1968. $27.50, 

The biochemical, medical and industrial development of peni- 
cillin from suitable members of the thousands of tested newer 
strains has meant that there are now thousands of pharmacists, 
pharmacologists, medical doctors, chemists, chemical engineers, 
technicians and other research assistants who need handj*- and 
accurate sources for the identification of their cultures. This 
offset printing of this earlier classic with its obvious de- 
scriptions, fine line drawings and photographs of many cultures 
in color and in black-and-white will fill much of that need very 
effectively. The system of nomenclatvire followed is more in 
manual than taxonomic monograph style stressing a workable sys- 
tem of descriptive diagnoses irtiich will enable the user to iden- 
tify the PenicillivBn mold in his culture with the genus deaned 
to include all penicillate green molds with or without asco- 
spores . 

It is interesting to note that the authors persist in using 



1968 Moldenke, Book reviews 3^7 

the plural form of the genus as a proper name, 

"PLANT NaiATOLOGT" by W. R. Jenkins and D. P. Taylor, xvii & 270 
pp., illus., Reinhold Publishing Compary, London, Amsterdam, 
& New York 10022 . 1967. $12 ..^0. 

Herewith a useful, fine book has been added to the "Reinhold 
Books in the Biological Sciences" series by eminently qualified 
authors . After a general introduction it covers anaton^y and 
morphology with many fine o-n.T'inal drawings and good photographs, 
nature and range of nematode damage to plants directly or indi- 
rectly in association vdth other pathogens, and descriptive 
accounts of the following nematodes — lancers, lesioners, bur- 
rowers, cyst-forraers, root-knotters, bulb and stem residents, 
stylet possessors, seed and leaf gallers, ring-formers, pins, 
sheathers , leaf dwellers , awlers , daggers , stubby rooters , etc . , 
as well as several typical non-parasitic soil forms. Some 
chapters are devoted to kinds of chemical, physical and cultural 
control. Biological control by natural predators, by trap crops 
that prevent maturation after entry, by growth of antagonistic 
plants nearby and by development of resistant varieties and make 
very interesting reading. 

With onl;'- an estimated 2 percent of nematodes scientifically 
described to date and with an estimated 10 — 2$ percent crop 
damage in the United States due to their depredations, this book 
should prove a fine introductory text and reference work for 
students in the field now or to be enticed into it. More materi- 
al of an ecological and physiological nature would have rounded 
out this survey better. 

"THE PRESERVATION OF NATURAL HISTORY SPECIMENS", Volume II by 
Reginald '.Vagstaffe and J, Havelock Fidler, xv & UOli pp., 
illus., Philosophical Library Press, New York 10016. I968. 
$17.50. 

These British authors have a marked advantage over comparably 
trained American workers because the country of the former has 
been effectively preserving a world-wide assortment of nature 
materials for centuries longer than we in herbaria, museums, 
universities, etc. 

This volxme explains clearly the most successful preservation 
methods for parts of, products of, or all of chordates ( including 
Walter's plastic method for the reproduction of reptiles and am- 
phibians), all types of plants, and geologic materials (rocks, 
minerals, fossils, relief maps;. 

The first volume appeared in 1955 and dealt with inverte- 
brates and their preservation. 

Several excellent appendices cover apparatus, preservatives, 
labelling, storage structures and problems, maintenance of col- 
lections, photographic records, and microscopy. 

This work makes an excellent, easily understood, yet thorough 
text for museum techniques and an excellent reference for the 



3li8 P U Y T L G I A Vol. 17, no. U 

professional and the aniateur's guidance. 

"THE PYRAMID OF LIVING THINaS" by Edith Raskin, 192 pp., illus., 
Mclraw-IIill Book Company, Toronto, London, Sydney, fi New 
York 10036. 1967. iiJU.^O. 

In this book the important ecological concept of the title io 
explained several times over in interesting, simple, quick- 
reading language. Young students as well as adults could profit 
from it and enjoy it. The interdependence of all life especially 
through the food chain is demonstrated among some of the major 
creatures of the following bioraes — the arctic and antarctic 
tundras, the tiagas, the deciduous forests, the middle latitude 
grasslsuids, the deserts, the tropical rain forests, and the 
savannas . 



ADDITIONAL NOTES ON THE SRIOGAULACEAE . HII 
Harold N. lioldenke 



ERIOCATJLACEAE Lindl. 

Additional synonymy: Eriocauloneae Desv., Ann. Sci. Nat. 13: 
U5. 1828. Eriocaulaceae Desv. ex Bullock, Taxon 7: 15. 19^8. 

Additional & emended bibliography: L., Sp. PI., ed. 1, 87 & 
129. 1753i Crantz, Inst. 1: 360. 1766j Hill, Herb. Brit. pi. 66. 
1769; Hope, Phil. Trans. Roy. Soc. 59: 2J41--2U5, pi. 12. 1770; 
Scop., Introd. Hist. Nat. 20U. 1777; Kuds., Fl. Angl., ed. 2, 2: 
iaU. 1778; Walt., Fl. Carol. 83. 1733; Lour., Fl. Cochinch. 1: 
60. 1790; Cothen., Disp. 16. 1790; Schreb., Gen. 2: 666. 1791; 
L. C, Rich., Act. Soc, Hist. Nat. Paris 1: 113. 1792; Vahl, Syaib. 
Bot. 3: 99* 1791; Roxb., Hort. Beng. 68. iSlIi; Veil., Fl. Flum. 
35. 1825; Lodd., Bot. Cab. lli: pi. 1310. 1828; Desv., Ann. Sci. 
Nat. 13: li5. 1828; Bong., M§m. Acad. Sci. St. P^tersb., ser. 6, 
1: I--7I1, pl. 1—19. 18 31; Wall., Numer. List 207. 1832; Hook, 
in Curtis, Bot. Mag. 59: pl. 3126. 1832; Bong., M^o. Acad. Sci. 
St. P^tersb., ser. 6, 2: 219—238, pl. 11—19 (1833) and ser. 6, 
Sci. Nat. 1: 5U5 — 560. 1835; Mart., Nov, Act. Acad. Leopold. - 
Carol. 17 (1): 1—72. 1835; Bong. M&n. Acad. Sci. St. P^tersb., 
ser. 3. Bot. 9—29, pl. 20—25. I8I1O; Kunth, Enum. Pl. 3: 1^92— 
530. 18la; Steud., Nom. Bot., ed. 2, 2: 65U. I8U; Griff., Itin. 
Notes [Posthum. Papers 2:] 65. 18U8; F. Muell., Trans. Philos. 
Soc. Victoria 1: 23— 2U. 1855; Steud., Syn. Pl. Gyp. 2: 261 & 268- 
233. 1855; Kym., Linnaea 27: 561—692. 1856; Steud., S^m. Pl. 
Glum. 2. 1856; Bsnth., Fl. Hongkong 382. I86i; K»rn. in Mart., 
Fl. Bras. 3 (1): 273—508, pl. 33—62. 1863; T. Camel, L:6m. Soc. 
Imp. Sci. Nat. Cherbourg lU [ser. 2, li] : 5 — 16. I868; K»rn. in 
Warning, Vidensk. Medd. Naturh. For. Kj/ibenh. 23: 309 — 3l6. I87I- 



1968 Moldenke, Notes on Erlocaulaceae 3U9 

l872j Benth., Fl. Austral, 7: 192. 1878; F. Muell., Syst. Census 
Austral. PI. 123. 1882; F. M. Bailey, Syn. Queensl. Fl. $78. 
1383; Benth. & Hook, f., Gen. PI. 3 (2): 1018— 102$. 1833; Hieron. 
in Engl. & Prantl, Nat. Pf lanzenf am . , ed. 1, 2 (U): 21—27. 1887; 
Poulsen, Vidensk. Medd. Naturh. For. Kj;ibenh. UO: 221—283, pi. 
6—12. 1888; A. W. Chapm., Fl. South. U. S., ed. 2, $02— ^Olt, 
658, & 696. 1889; F. Muell., Proc. Linn. Soc. New S, V^ales 5: 2^0. 
1890 ; F. Muell., Bot. Centralbl. UU: 302. I89O; Korong, Bull. 
Torrey Bot. Club 18: 351—362. I89I; Maxim., Diagn. PI. Nov. As. 
8: 25. 1892; Jacks, in Hook, f . & Jacks., Ind. Kew., pr. 1, 1: 
879. 1893; Moore & Betche, Handb . Fl. N. South Wales khO, 1893; 
J. G. Baker, Journ. Linn. Soc. Lond. Bot. 20: 227. 1893; Coult., 
Contrib. U. S. Nat. Herb. 2: I;59. I89U; Jacks, in Hook. f. & 
Jacks., Ind. Kew., pr. 1, 2: 681. 1895; Britton & Br., 111. Fl., 
ed. 1, 1: 371—373 & 602, fig. 899—901. I896; Ruhl. in Engl., 
Bot. Jahrb. 27: 65—85. 1899; Tate, Trans. Roy. Soc. S. Austral. 
23: 291. 1899; H. T. Holm, Bot. Gaz. 31: 17—37. 1901; N. E. 
Br. in Thiselt.-Dyer, Fl. Trop. Afr. 8: 255. 1901; F. M. Bailey, 
Queensl. Fl. 6: 1715. 1902; Prain, Bengal PI., pr. 1, 1: 121. 
1903; B. L. Robinson, Rhodora ^i 175—176. 1903; J. K. Snail, 
Fl. Southeast. U. S., ed. 1, 236. 1903; Post & Kuntze, Lexicon 
^hh, I90U; Ruhl. in Urb., Engl. Bot. Jahrb. 37: 519—520. I906; 
Ann. N. I. Acad. Sci. 17 (1): pl. 2U, fi^. 1. 1906; C. H. Wright, 
Kew Bull. Misc. Inf. 1907: 3—k* 1907; Robinson & Fern, in A. 
Gr^, New Man. Bot., ed. 7, 260—261 & 898. I908; M. A. Day, 
Check List 39. I908; Beauverd, Boiss., ser. 2, 8: 986 — 988. 1909; 
Nakai, Bot. Mag. Tokyo 2U: 5—6. 1910; Kawakami, List PI, Formos. 
130. 1910 ; G. T. Stevens, 111. Guide Flow. PI. 9, fig. 5. 1910; 
A. Chev., Sudania 1: 7. 1911; W, H, Br., Contrib, U, S, Nat. 
Herb. 13: 323. 1911; Nakai, Bot. Mag. Tokyo 26: [93— 9U] . 1912; 
Ann. Rep. N. J, State Mus. 1910: pl. 28, fig. 2. 1912; F. M. 
bailey, Compreh. Cat. Queensl. Pl. 58U. 1913; J. K. Steiall, Fl. 
Southeast. U. S., ed. 2, 236. 1913; Britton & Br., 111. Fl., ed. 
2, 1: U53— U55 & [678]. 1913; Domin, Bibl. Bot. 20: 506. 1915; 
Maiden & Betche, Census New S. Wales Pl. 38. 1916; Fern., Rho- 
dora 23: 92. I92I; Fyson, Joum. Indian Bot. 2: 133—150, 192— 
207, 259—266, & 307—320, pl. 1— )40 (1921) and 3: 12—18 & 91— 
115, pl. 11—32. 1922; Kolfino, Physis 6: 361—363. 1923; Ltltzel- 
burg, Estud. Bot. NordSste 3: lii7 & 150. 1923; Anon., Kew Bull. 
Misc. Inf. 1923: 303. 1923; Fyson, Indian Sp. Erioc. 1—88, pl, 
1—51. 1923; Herzog in Fedde, Repert. Spec. Nov. 20: 82—38. 
I92U; Ruhl. in Fedde Repert. Spec. Nov. 22: 29—35- 1925; Alv. 
Silv., Fl. Mont. 1— U26, pl. 1— 25U. 1928; A. S. Hitchc, Prop. 
Brit. Bot. 122. 1929; Uohof in Karst. L Schenck, Veg etationsb . 
21 (1-2): n.p. 1930; Ruhl., Notizbl. Bot. Gart. Berlin 10: 
lOUO— IOI4I;. 19 30; Herzog in Fedde, Repert. Spec. Nov. 29: 202— 
213. I93I; N. E. Br., Kew Bull. Misc. Inf. 1931: 61. 1931; Ewart, 
Fl. Vict. 263. 19 31; Solomon, Journ. Indian Bot. Soc. 10: 139 — 
lUi. 1931; Tu, Chinese Bot. Diet., abrdg. ed., 13U7. 1933; R. M. 
Adam, New Fl. & Silv. 6: 60— 63, pl, 2h ^ 2$. 1933; J. K. Small, 
Man. Southeast. Fl. 258. 1933; Tang, Contrib. Inst. Bot. Nat. 
Acad. Peiping 2: 133. 193U; H, B. Davis, Life Z: ,'.orks Pringle U3, 



3^0 P }! Y T L G I A Vol. 17, no. h 

55, 56, 9k, 105, 123, lla, 219, '/ 655. 1936; 'J&n Steenis, Trop. 
Natuur 25: 2. 1936; Moldenke, N. Am. Fl . 19: 17—50. 1937; Cory, 
Texas Agr. Exp. Sta. Bull. 550: 29. 1937; Merr. '/ Walker, Eibl. 
East. Asiat. Bot. 3l3. 1938; Satake, Joum. Jap. Eot. 15: HiO— 
lh5 & 627—632. 1939; Moldenko, PhytoloRia 1: 309—336. 1939; 
Moldenke, Camecie Inst. V/ash. Publ. 522: 137— lli8. 19h0; I'olden- 
kc, Phytolo£:ia 2: 6—7. 19U.; Wells. Pot. Rev. 3: 537. 19U2; I'ol- 
denke in Lundell, Fl. Texas 3 (1): h — 5. 19U2; Carolin. Florist 
Gov. J. Drayton S. C. Hi. 19U3; Black, Fl. S. Austral., ed. 2, 1: 
179. 19U3; Moldenke in V/oodson & Schery, Ann. Lo. Bot. Card. 31: 
65 — 71. 19Ui; Rouleau, Gontrtb. Inst. Bot. Univ. Montreal 5Ii: 
161 & 313. I9hhi furies & Robertson, U. S. Pub. l.'ealth Bull. 286: 
106. 19llli; W. A. Murrill, Guide Fla. PI. 2h. 19U5; Castellar.os in 
Descole, Gen. Sp. PI. Argent. Erioc. 87, pi. 17. 19U5i Abbiatti, 
Revist. Mus. La Plata Bot. 6 (26): 329—330, pi. 2 (1), fig. h (d) 
& 6. 19U6; Razi, Jovim. Itysore Univ. 7 (U): 77. 19U6; PJnodora U8: 
iv & 58. I9U6; Moldenke, Knovm Geogr. Distrib. Erioc. [1] — 62. 
I9U6; R. n. Tatnall, Fl. Del. 75. 19U6; Abbiatti, Soc. Argent. 
Bot. Bui. 1: 280—281. I9I16; W. Robyns, Bull. Jard. Eot. Erux. 
18: 135. I9U6; J. Hutchinson, Botanist in South. Afr. 591, 65ii, & 
660. I9U6; Moldenke, Phytologia 2: 3ii9— 352. 19^7; Moldenke, 
Wrightia 1: 220—221. I9I18; Castellanos, Lilloa 20: 2Uh. 19l49; 
Raizada, Sci. & Cult, ll: 387—388. 19U9; Faegri k Iversen, 
Text-book Mod. Pollen Analys . 193 & 221. 1950; Hare, Jo\irn. Linn. 
Soc. Lond. Eot. 53: 1^22— lj;8. 1950; Herter, Rev. Sudam. Bot. 8: 
16>-I61i. I95O; Reitz, Anais Bot. 2: 32 & 3U. 1950; Satake, Joum. 
Jap. Bot. 26: 221. I95IJ Maekarra, Joum. Jap. Eot. 26: II6. 
1951 ; Meikle, Am. Joum. Bot. 39: hh—^1. 1952; Penfound, Eot. 
Rev. 18: li31. 1952; Beard, Ecol. Monog. 23: 177. 1953; Zinderen- 
bakker, S. Afr. Pollen 1: 32, 36, & 79, pl. 7, fig. 33 «= Uii. 1953; 
Moldenke, Biol. Abstr. 27: 981|, 2026, & 3121. 1953; Moacyr Lisboa, 
Cent, Nascijn. Leon. Bot. Damazio [2]. 195U; Reitz, Sellowia 6: 
256. 195U; Koyama, Philip. Journ. Sci. Bot. 8U: 367—368 L 378, 
pl. 6. 1955; Razi, Journ. Ifysore Univ. E.li; (10): U60. 1955; Razi, 
Contrib. Bot. UO: 92. 1955; Razi, Proc. Nat. Inst. Sci. India 21B 
(2): 82. 1955; Anon., Assoc. Etud. Tax. Fl. Afr. Trop. Index 1955: 
29—30. 1956; Koyama, Joum. Jap. Bot. 31: 9—11, fig. 3. 1956; 
Masa Ikusi, Pollen Gr. Jap. 1956; Angely, Cat. Estat. 10: [2]. 
1956; Mutisia 25: 26, 1956; Hanbo, Sellovria 7: 263 £t 292. 1956; 
Soukup, Biota 1: 122 £: 208. 1956; Boivin, Bull. Bot. Soc. France 
103: 503. 1956; Reitz, Sellowia 7: 9U ^ 12U— 125. 1956; Moldenl:e, 
Bull. Jard. Bot. Brux. 27: 116 — Ihl* 1957; M. T. Davis, Taxon 6: 
179 & 181. 1957; J. A. Steyerm., Fieldiana 28: 1188. 1957; Anon., 
Biol. Abstr. 29: 32U8 £t 3626. 1957; Angely, Fl. Paran. 7: 7 
(1957) and 10: 1—16. 1957; Anon., Biol. Abstr. 29: 32U8 & 3626 
(1957) and 30: 3931 & U393. 1958; Anon., U. S. Dept. Agr. Bot. 
Subj. Index 5: U226 — U277. 1958; Kostermans, Proc. Sympos . Humid 
Trop. Veg. 159. 1958; Suvatabandhu, Proc. Sympos. Humid Trop. Veg. 
173. 1958; Alain, Revist. Soc. Cub. Bot. 15: U & U9. 1958; de 
Roon, Intemat. Direct. Spec. 79, 201, & 231. 1958; Angely, Eol. 
Inst. Par. Bot. 8: 3li. 1958; Soukup, Biota 2: 57. 1958; V^ Robvns 
Fl. Cong. Belg. Tabl. Anal. 17, 57, & 61^. 1958; Bullock, Taxon 7:' 



1968 Moldenke, Notes on Eriocaulaceae 35l 

15. 19^8; P. van Royen, Nov. Guin., new ser,, 10: 21 — hh, fig. 1- 
5. 1959} Moldenke, R^sum^ [1], 3—16, 19, 22, 23, 25, 27, 32, 35- 
38, la, U3— U8, 51— 5U, 57, 63, 66—80, 83, 8I1 87—89, 91, 92, 
9U— 109, 112—117, 119, 123, 126, 129, 133—138, lliO. Iii2— 151, 
153, 151, 156—163, 165—167, I69— 176, 178— 181, iSh, 186, 188, 
190—193, 201, 20ii— 209, 211, 218, 220, 237, 2liO, 2U9, 277, 279— 
281, 28U— 29U, 301, 302, 309, 320, 323—329, 33ii, 3U2, 3li5, 350— 
352, 355, 395— I4O2, UlU— 1:20, U2I;, U26, h28, & U79— h93. 1959i 
Soukup, Biota 5: 300—301. 1959; Moldenke, Biol, Abstr. 33: 3171. 
1959} T^-on 8: 77. 1959} Bullock, Taxon 8: 171. 1959} Anon., As- 
soc. Etude Tax. Fl. Afr. Trop, Index 1958: 31. 1959} J. Hutchin- 
son, Fam. FlOTT. PI. 2: 57U. 1959} Razi, Rec. Bot. Surv, India 18: 
19. 1959} Anon., Kew. Bull. Gen. Index 1929-1956, 111. 1959} 
Reitz, Sellovria U: 103 . 1959; Moldenke, R5sum5 Suppl. 1: [1]— 3, 
5—19, 21, 23, 25, & 26 (1959) and 2: [1], 2, h— 7, 9, & 16. 
I96O} P. van Royen, Blxoraea 10: 126 — 135, fig. 1. I960} Leenhouts 
in Lam & Leenhouts, Blumea 10 (2): xvi. I96O} Angely, Liv. Gen. 
Bot. Kras. 9, 19, ■^i hh. i960} Angely, Fl. Paran. 15: 6 & li^ 
(i960) and 16: 33. I960} Straka, 3rdkunde lit: 60, 87, 90, & 9$» 
I96O} Renn6, Levant. Herb. Inst. Agron. 68. I96O} Moldenke, Biol. 
Abstr. 35: 983, 1688, & 2177. I960} Santapau, Fl. Bombay & Sal- 
sette [3]. I96O} Nath, Bot. Surv. South. Shan States 9 <Sc 20. 
I96O} Panigrahi & Naik, Bull. Bot. Surv. India 3- 383. 1961} 
Runner, Rep. G. W. Groff Coll. 292. I96I} Fables, Bartonia 32: 
9. 1961 : Reitz, Sellowia I3: 37. 1961} Hamann, vaildenorria 2: 
6U0, 6kh, chart 1 6U5, 651— 65U, 656—658, 66O 663, 665—685, 
687, 69U, 711— 7lii, 7iiO, 7hh, IhS, 7U6, 752, 75I4, 761, & 763. 
I96I; Angely, Fl. Paran. 17: 10, 12, & 2ii. I96I; Tamayo, Bol. 
Soc. Venez. Cienc. Nat. 22: UO & 88. I96I} Moldenke, Biol. Abstr. 
36: 28U3. I96I} Clapham, Tutin, & Y/arburg, Fl. Brit. Isles, ed. 
2, 961—962. 1962} Hamann, !7illdenovri.a y. 179 & 191. 1962} Mol- 
denke, Biol. Abstr. UO: 250 k 1560. 1962} Van Steonis-Kruseman, 
Fl. Males. Bull, 3: xli, 687, 780, 78I, 861, & 862, 1962} J. H. 
Willis, Handb, PI, Vict, 281, 1962} K. Larsen, Nat, Hist. Bull. 
Siam Soc, 20: 113, 1962} Hatusima, Mem, South, Indust, Sci. Inst, 
Kagoshima Univ, 3 (1): 123 (1962) and 3 (2): 123 & 131. 1962} G. 
L, Shah, Biill. Bot, Surv. India h'. 237. 1962} L. B. Sm., Gontrib. 
U. S. Nat. Herb. 3^1 225. 1962} Angeily, Fl. Bacia Par. 22: 15, 

25, &31. 1962} Moldenke, P.Ssum^ Suppl. y. [1]— 5, 9, 12, 15— 21;, 

26, 28, 31, & 32 (1962), U: [1]— 7, 11, £c 21 (I962), and 5: [1], 
2, 5, L S, 1962} Hocking, Excerpt. Bot. A.l;: ^91—^93 (I962) and 
A.6: li5U. 1963} H. P. Riley, Fam. Flow. PI. S. Afr. I99. 1963} 

Prain, Bengal PI., pr. 2, 1: 121 (I963) and 2: 81;7— 81;8. 1963} 
Moldenke, Biol. Abstr. U2: 1517. 1963} Soukup, Biota hi 320. 
1963} E. H. Bryan, Pacific Botanists. 1963} Killick, Bot. Surv. 
S. Afr. Mem. 3l: 87, 108, IO9, & 119. 1963} Kegnauer, Cheraotax. 
Pfl. 2: 152— l51i & 517. 1963} Arker, Water PI., ed. 2, 286. 1963} 
J. Joseph, Bull. Bot. Surv. India 5: 283 & 297. 1963} Montgomery 
Sc Fairbrothers, Bull. Torrey Bot. Club 90: 92 & 96. 1963} Glea- 
son (£c Cronquist, Man. Vase. PI. I83 — l81i. 1963} Espirato Santo, 
Junta Invest, Ultramar Est. Ens. & Docxim. lOh: 51i t 88. 1963} 



3^2 P H Y T L n I A Vol. 17, no. U 

Moldonke R6oum6 Guppl. 6: [1], 2, 5, 6, 8, «r 9 (1963), 7: 3—6 
(1963), B: 2, 3, f. 5 (19610, 9: 6 (196U), 10: 1, 2, v. 1—7 (1961i), 
and 11: [1] &. h—6, I961i; Moldenke, Diol. Abstr. U^: 3521, i;019, 
l^.h3, B.iii;, D.117, t D.118. l96Ui Anpely, Dibl. Vge. ?aran. 90, 
98, 101, 15>'5, 1^0, 132, 19:^—197, &- 2:;3. 196h; Ancely, II. Paran. 
32: 60. I96U; Koldenke, I.i&n. LIus. iiist, liat, Paris, new ser. B, 
Bot. 1^: 6. 1961ij Koyana in Kitamura, Liurata, -^^ rloyana, Col. 
Illustr. Herb. PI. Japan 175 — 185, pi. );8. 196ii; Anon., biol. Ab- 
str. U5: B.U2 fv. B.116. I96U; Hocking, Excerpt. Bot. A. 8: 190. 
I96U; Soukup, Biota ^: I9U. I96U; P. Tomlinson, Jo\irn. Linn. boo. 
Lond. Bot. 59: 16 3-— 173. 196U; Anon., Assoc. Etud. Tax, Kl. Afr. 
Trop. Index I963: 9. 196U; Anon., Ann. Uo. Bot. Card. 51: iv. 
I96UJ Langman, Select. Guide Lit. Flow. PI. Mex. 911. 196ii; Anon,, 
Assoc. Etud. Tax. Fl. Afr. Trop. Bull. 15: 23. 1961^; llaexiire, 
Assoc. Trop. Biol. Bull. 3: 25. 1961;; Altman &t. Dittmer, oiol. 
Data Book 568. 1961;; Klein, Anais AV Congr. Soc. Bot. Bras. 271. 
1961;; Marie-Victorin, Fl. Laurent., ed. 2, 83, 90, 679, & 857. 
I96I;; Hao It Sastry, Bull. Bot. Surv. India 6: 281 & 281i. I96I;; 
Piint, Reg. Veg. 9. 1961; Batson, ".Vild Fls. S. G. 28. I96I;; Pani- 
gralii, Chowdhury, Raju, & Deka, Bull. Bot. Surv. India 6: 260 — 
261. I96I;; Bhattacharyya, Bull. Bot. Surv. India 6: 208. I96I;; 
C. M. & D. S. Patel, Vidya 7: [58]— 70. I96I;; J.'.oldenke, Biol. 
Abstr. 1;6: 1012 & 2131 (1965), U6 (3): B.i;6 & B.121 (1965), and 
1;6 (6): E.l;5. 1965; F. A. Larkley, List Ord. Fan. Anthoph. 15, 
113, & 16U. 1965; Naurois & Roux, Bull. Inst. Fr. Afr. Noire A. 
27: 851;. 1?65; S. A. J.:anning, Syst. Guide Flow. PI. 21; 2c 237. 
1965; Anon., Biol. Abstr. 1;6 (6): B,l;5. 1965; iiunter, Assoc. 
Trop. Biol. Bull. $1 33. 1965; Hedberg, Assoc. Etud. Tax. Fl. 
Afr. Trop. Bull. 16: h. 1965; R. C. Cook, Leaders Am. Sci., ed. 
6, Uli;. 1965; A. Robyns, Ann. Mo. Bot. Card. 52: 238, 239, 2l;2, 
& 2l;6. 1965; F. H. Llontgonery, Native ".iild PI., pr. 2, 9. 1965; 
Thanikaimoni, Pollen &: Spores 7: 131 — 191. 1965; J. S. Beard, 
Descrip, Cat. 'J. Austral. PI. 9. 1965; D, V.'alker, Govt. Sarawak 
Sympos. Ecol. Res. Humid Trop. Veg. lUl. 1965; Thanikaimoni, 
Mlia. L'us. Hat. Hist. Hat. Paris, new ser. B, Hi: 9 — 38. 1965; 
Hedberg, V/ebbia I9: 526. 1965; Humbert, Trav. Sect. Scient. Sc 
Techn. Inst. Fran^. Pond,, ser. 6, Hot. Carte Ladag. 66. 1965; 
Stocking, Hat. Conserv. Ecol. Stud. Leafl. 6: [15]. 1965; F. K. 
Fosberg, Govt. Sarawak Sympo-^ . licol. Res. Humid Trop. Veg. 286. 
1965; Moldenke, R^sum^ Suppl. 12: [1]— 5 & 7—12 (1965), 13: [1], 
3, 5, & 7 (1966), and lU: [1]— 3, 8, & 9. 1966; F. H. Montgomery, 
Plants f-^om Sea to Sea U & 9. I966} Thanikaimoni, Biol. .^bstr. 
Ii7: lj.69. 1966; 0. D, Evans, Contrib. New S, '.Vales Nat. Herb. Fl. 
Ser. 27/28: 9—12. I966; Krai, Sida 2: 285—332. I966; Erdtoan, 
Pollen Morph. I63, fig. 9h. 1966; Anon., Ind. Bibliog. Bot. Trop. 
3 (1): la. 1966; P. B. Tomlinson, Excerpt. Bot. 10 (1;): A. 310. 
1966; Airy Shaw in Willis, Diet, Flow. PI., ed. 7, vii, lii,l68, 
223, 221;, 3ii9, 396, 1;17, ia8, 1;83, 620, 61;7, 758, 950, 956, 1057, 
1091, & 1092. 1966; Anon., Gen. Costa Ric. Phan. 2. I966; R. C, 
Jacks., Reg. Veget. 1;3: 33. 1966; Anon., Biol. Abstr. 1;7 (10): S. 
50. 1966; J. A. Steyerm., Act. Bot. Venez. 1: 15, 19, 78, 135, & 
1U8. 1966; Shinners, Sida 2: li;l. I966. 



PHYTOLOGIA 

Designed to expedite botanical publication 
Vol. 17 November, 1968 No. 5 



CONTENTS 



CHUNG, In-Cho, Studies in Manctt/a I Ruhiaceae) Section 

P\nha>ithos Schuin 353 

LITTLE. E. L., [r.. I ransters to ii iia pira fro)ii iorruhia 

(\\ctagi)iaceae) 367 

DEGENER. O. & I.. Rei ieii of F. I:. W'iitnner. i.ampa}iulaceae- 

Lobelicjideae Supple n/ei/tum et C.a>npnnulaceae-(..\phioideae. 
Das Pflanzenreich. IV . 2^bc (108. Heft). l-\. S]b-1024; uith 
descri pt io>i of l rematolohe I ia n- iiniiieri De g. & Deg.. sp. nor. 369 

MOLDENKE. H. N., Additional notes on the liriocaulaceae . \l\ . .372 



KOWWl.A, T.. Iconograpbia C.yperacearuiii II 396 

.MOLDENKE. A. L., /^oo/C,t.//V/rs 422 



Published by Harold N. Moldenke and Alma L. Moldenke 

303 Parkside Road ^ 

Plainfield, New Jersey 07060 tX. CO \^ 

U.S.A. ^^ 22 P' 

<l "^ O 

CO >- 

Price at this number, SI; per volume, $6.75 in advar|Pt^ tH 



or $7 at close of volume 



•z. z 



"1 



STUDIES IN MANETTIA (RUBIACEAE) 
SECTION PYRRHANTHOS SCHUM. 

In-Cho Chung, Mansfield State College 

The first of this series of papers, section Heterochlora , 
appeared in Phytologia vol. 15 (no. 4): 272-288, 1967. 

Six species are recognized in section Pyrrhanthos in this 
paper. The morphological characteristics of the section are as 
follows. The corolla is red, clavate-tubular or infundibular, 
qradually V7idened from the base upwards, 2-6 cm. long, mostly 
glabrous outside, sometimes sparsely puberulent with 1-few-celled 
hairs, or rarely densely pubescent with slender multicellular 
hairs outside, glabrous within except for a band of hairs near 
the base of the tube. Anthers are subsessile to with more or 
less apparent filaments, the filaments 0.5-5 mm. long, inserted 
at the summit of the corolla-tube. Stigmas are ovate to oblong, 
obtuse at the apex. The disk is free from the calyx; calyx- 
lobes 4. 

Key to Species 

1. Leaves very small, 7-15 (24) mm. long, with no apparent 

lateral veins; corolla glabrous outside. 

2. Leaves coriaceous, elliptical, rarely ovate, 4-7 mm. 
wide, obtuse at apex, rounded to obtuse at base; 
calyx-lobes oblong, obtuse, about 2 mm. long; stip- 
ules coriaceous, truncate, glandular-toothed, hispid 
outside; branches subtetragonal to subterete, striate, 
scaberulous , with hispid, short hairs turned upward 
on the angles of young branches. 
West Indies M. domingensis 

2. Leaves membranous, narrowly lanceolate, 1 . 5-3 mm. 
wide, acute to acuminate at apex, attenuate at base; 
calyx-lobes linear, acute, 3-4 mm. long; stipules 
membranous, with deltoid, entire, short, free portion, 
glabrous; branches strongly tetragonal, winged, 
smooth, glabrous. Uruguay, S. Bra'zil. . M. tweedieana 

1. Leaves large, 2.5-14 cm. long with conspicuoui lateral veins, 

3. Flowers 4 or 5 in terminal cymes; calyx-lobes linear, 
attenuate-acuminate, 11-18 mm. long; free portion 

of stipules with 3 unequal aristae; corolla glabrous 

outside. Mexico M. zimapanica 

3. Flowers 1 or 2 in leaf-axils and 1 to several at 

the end of short branches; calyx-lobes broadly ovate 
to linear, if linear or linear-subulate, then pedi- 
cels, ovaries, calyx-lobes, and outer surface of 
corolla densely pubescent with slender multicellular 
hairs. Central S. America. 
353 



35U P H Y T L I A Vol. 17, no. 5 

4. Corolla and capsules densely pubescent with slender 

multicellular hairs outside. S. Brazil. . M. pubescens 
4. Corolla and capsules glabrous or puberulent with 

1-few-cellod hairs. 

5. Pedicels capillary, with 2 basally conate 

bracts at the base; capsules oblong to oblong- 
turbinate, 5-8 mm. long, 3-4 mm. wide; stigmas 
oblong. S. Brazil M. gracilis 

5. Pedicels not capillary, naked or with 2 distinct 
bracts near the middle; capsules mostly subcom- 
pressed ellipsoidal with slightly tapered apex 
or subglobose, mostly 8-12 mm. long, 6-7 mm. 
wide; stigmas ovate. Central S. America. 



M. cordifolia 



M. domingensis Sprague, Bull. Herb. Boiss. II. 5: 266. 
1905. Type: Eggers 2178 (K, isotypes BM,G). Fig. 1. 

Branches slender, subtetragonal to subterete, striate, sca- 
berulous, with hispid short hairs turned upward on the angles 
of young branches, internodes slightly shorter than the leaves; 
stipules truncate without free portion, coriaceous, glandular- 
toothed, hispid outside; petioles very short, 0.5-2 mm. long, 
hispid, leaf-blades coriaceous, elliptical, rarely ovate, obtuse 
at the apex, rounded to obtuse at the base, 7-15 (24) mm. long, 
4-8 mm. wide, with no apparent lateral veins, glabrous on the 
wrinkled upper surface, glabrous beneath except for the short 
hispid hairs on the midvein, ciliate on the reflexed margins; 
flowers solitary rarely two at the ends of short lateral and 
terminal branchlets; pedicels slender, 5-9 mm. long, glabrous; 
calyx-lobes oblong, mostly about 2 mm. long and about 1 mm. 
wide, obtuse, glabrous, 1-nerved, connate at the base for about 
0.2 mm.; corolla 20-30 mm. long (the lobes 4-6 mm. long)f about 1.- 
mm. wide at the base, 4-6 mm. wide at the apex of the tube, 
glabrous outside, glabrous within except for a band of hairs near 
the base; stamens reaching the apex of the corolla; filaments 
3-5 mm. long, inserted at the apex of the corolla tube; anthers 
2-2.8 mm. long; style exserted for about 3 mm., stigmas short, 
ovate, obtuse at the apex; disk free; capsules turbinate to 
oblong-turbinate, 5-7 mm. long, glabrous. West Indies: Sto. 
Domingo: Valle Nuevo, Eggers 2178 (BM,G,K) , Santiago Bueno 1772 
(F,GH); Prov. Azua, Ekman H6784 (S,US) , H11988 (S) . 
Prov. La Vega: Fuertes 1754 (G,GH, NY,P,US); Prov. San Juan: 
Sanbana Nueva, Howard 9 012 (BM, NY, P,S, US). 



1968 Chiui£, Studies in llanettia 35$ 

M. tweedieana Schum. Mart. Fl. Brass. 6 (6) : 1889. 
Type: Tweedie. Fig. 2. 

Branches slender, strongly tetragonal, winged on the angles, 
glabrous, smooth; stipules membranous, glabrous, 1-1.5 mm. long 
including the deltoid, entire free portion; leaf-blades narrowly 
lanceolate, acute to acuminate at the apex, attenuate to very 
short petioles, about 10 mm. long, 1.5-3 mm. wide, with no apparent 
lateral veins, glabrous on both sides except for the minute his- 
pidulous hairs near the reflexed margins; flowers solitary, term- 
inal; pedicels 8-18 mm. long, glabrous; calyx-lobes linear, 3-4 mm. 
long, about 0.3 mm. wide, acute, glabrous, connate at the base, 
alternate with glandular- toothed deltoid lobules (0.5 mm. long); 
corolla 30-37 mm. long (the lobes 2-3 mm. long) , 1.7-2 mm. wide 
at the base, 6 mm. wide at the apex of the tube, glabrous outside, 
glabrous within except for a band of hairs near the base; anthers 
about 2.5 mm. long, nearly sessile, the very short filaments 
about 0.5 mm. long, inserted at the apex of the tube; style ex- 
serted for about 4 mm; stigmas ovate, obtuse at the apex. 
Uruguay: Rio Grande, Tweedie (type, B, photo!), Brazil: Santa 
Catharina, Tweedie (GH, K) . 

M. zimapanica Hemsl. Diagn. Pi. Nov. Mex. 30. 1878. 
Type: Coulter 203 (K) . 

Flower-bearing branchlets subterete to somewhat compressed, 
moderately pubescent with short, spreading hairs; free portion 
of stipules trisetose, the middle arista 2-2.5 mm. long, the 
lateral aristae 1-1.5 mm. long, the sheath pubescent outside; 
petioles 4-5 mm. long, sulcate, pubescent; leaf-blades ovate- 
lanceolate, attenuate-acuminate at the apex, attenuate at the 
base, 6-9.5 cm. long, 2-4 cm. wide, sparsely pubescent except the 
upper surface of the midvein with dense short broad hairs and 
the lower surface of 3 or 4 pairs of lateral veins with moderate 
hairs; flowers 4 or 5 in terminal cymes; pedicels slender, 5-20 
mm. long, minutely puberulent; ovaries semispherical , 1.5-2 mm. 
long, glabrous; calyx-lobes linear, attenuate-acuminate, 11-18 mm. 
long, connate at the very base, 1-3-nerved, sparsely ciliate, 
glabrous on both sides; corolla 35-39 mm. long (the lobes 5-7 mm. 
long), glabrous outside, glabrous within except the pubescent 
basal 10 mm; anthers about 3 mm. long, subsessile, inserted 
near the apex of the tube in short-styled flowers; stigmas oblong, 
obtuse at the apex. 
Mexico: Zimapan, Coulter 203 (K) . 

The disk is slightly convex and free from the calyx, or 
nearly flat to slightly concave and adnate to the calyx-tube. 



356 PHYTOLOGIA Vol. 17, no. 5 

M. pubescena Cham. & Schl. Linnaea 4: 170. 1829. 

Type: Sellow . Fiqs. 4-9. 

M. villosa Cham. & Schl. I.e. 172. 1829, Type: Sellow . 

gellow 17^0 (B, photo) I 

M. confertif lora Benth. Linnaea 23: 443. 1850. 

Type Regnell I: 3681 

M. pubescens var. villosa Schum. Mart. Fl . Bras. 6(6): 173. 

T88T: 

Branches, petioles, lower surface of leaves, pedicels, ovaries, 
capsules, calyx-lobes, outer surface of corolla densely pubescent 
with slender multicellular hairs; free portion of stipules erect, 
mostly lanceolate, often tapering to an arista, frequently 3-lobed, 
with the lateral ones shorter and setaceous, the middle one up to 
11 mm. long; calyx-lobes linear-lanceolate to linear-subulate, 
5-13 mm. long, subulate short (1-5 mm. long) lobules usually 
present; corolla 33-50 mm. long, 4-8 mm. wide; filaments about 1 mm. 
long, anthers about 5 mm. long; capsules subcompressed ellipsoi- 
dal, 6-15 mm. long, 5-7 mm. wide; leaf-blades mostly ovate, rounded 
to obtuse at the base, acuminate at the apex, 25-80 mm. long, 
12-38 mm. wide. 

Brazil: Prov. Minas Geraes: Damazio 966 (G) , 40153 (NY), 
Glaziou 17630 (P) , Macedo 2895 (BM, G, MO, US) , Mosen 905 (S) , 
1864 (S) , Regnell I: 368 (K, P, S) , Saint-Hilaire 529Tn part (P) , 
Widqren 189 (S) , s.n. anno 1845 (GH, K, S) ; Rio deJaneiro: 
Widgren TFT; S. Paulo: Leite 3951 (F) , Loefgren 3445 (F) , 
P. Campos Porto 3273 (F) ; Prov. Parana: Dusen 16539 (MO, S) ; 
Prov. Sta. Catharina: Dusen 8416 (F, S) , Ule 1261 (P) ; Prov. 
Rio Grande doSul: Leite~T772 (F) , Rambo 47^6 (F) . 

M. gracilis Cham. & Schl. Linnaea 4: 169. 1829. 
Type: Sellow photo! 

Var. gracilis . Fig. 3. 

Young branches slender, subterete, striate, densely pube- 
scent; stipules deltoid, erect, toothed; petioles 12-18 mm. long, 
densely pubescent; leaf-blades ovate-lanceolate, rounded at the 
base, acuminate at the apex, 3-7 mm. long, 6-25 mm. wide, moderately 
pubescent beneath, with 4 or 5 lateral veins on each side of the 
midvein; flowers 1 or 2 in the leaf axils or several at the end 
of the branchlets ; pedicels capillary, 12-18 mm. long, glabrous 
or puberulent, with 2 small, basally connate bracts at the base; 
ovary glabrous to puberulent; calyx-lobes deltoid to lanceolate, 
more or less pubescent, 1-1.5 mm. long, connate at the base for 
about 0.5 mm; corolla 23-30 mm. long, slender, 3-5 mm. wide, gla- 
brous or sparsely pubescent outside with slender few-celled 
hairs, glabrous within except for a band (3-5 mm. long) of hairs 
near the base; stigmas oblong, obtuse; anthers 2.5-4 mm. long; 
filaments 1.5-3 mm. long, inserted at the apex of the corolla- 
tube; capsules oblong to oblong-turbinate, 5-8 mm. long, 3-4 mm. 
wide, glabrous or sparsely puberulent. 

Brazil: Prov. Minas Geraes: Regnell I : 366 in part (S) f Prov. 
San Paulo: Hcehne 1642 (F) , Weir s.n. (BM) ; Prov. Parana: Dusen 
9984 in part (NY, PH, S) ; Prov. Santa Catharina: Reitz & Klein 3401 
in part (NY, UC, US), 4115 (G, NY, UC, US), 6912 (UC, US). 



1968 Chung, Studies in Manettia 357 

M. gracilis var. glabra Benth. Linnaea 23: 444. 1850. 
Type : Regne ll I : 366 ! 

M, burchellii Wernh. Journ. Bot. 57 Suppl . 18. 1919. 
Typel Burchell 4855 (K) ! 

Branches glabrous; petioles ciliate; leaf-blades glabrous 
except for the puberulent veins on the lower surface; pedicels 
and calyx-lobes glabrous. 

Brazil: Prov. Minas Geraes: Hoehne 10-5-1927 (NY), 19536 (F) , 
Novaes 3634 (F) , Regnell I : 3 6 6 (S, US) , Widgren 191 (S) ; Rio 
deJaneiro: Brade 14642 (F) , 21224 (F) ; Prov. S. Paulo: Brade 6314 
(S) , Hoehne 17637 (f) , 42644 (F) , Eiten 2793 (NY, US) , Mosen 13?0' 
(S) , Usteri 128 (K) , Weir 226 (F, BM) , s.n. (K) ; Prov. Parana: 
Dusen 9765 (BM, F, G, S, USTT 9904 (NY) , 9984 in part (UC) , Jonsson 
283a (BM) ; Prov. Santa Catharina: Bowie & Cunningham s.n. (BM) , 
Reitz & Klein 4240 (NY, S, UC, US) , Schwacke 123 (FTT Burchell 
4855 (K) . 

Key to the Varieties and Forms of 
M. cordifolia 

1. Calyx-lobes connate at the base. 

2. Corolla glabrous outside or nearly so. 
3. Leaves more or less pubescent. 

4. Leaves at least puberulent on the veins 
beneath, but not tomentose. 
var. cordifolia 

4. Leaves tomentose beneath with gray or 
yellowish-brown hairs. 
var. cordifolia form, incana 

3. Leaves glabrous. 

5. Branches subterete, glabrous. . .var. glabra 
5. Branches tetragonal, with short, ref lexed, 

hispid hairs on the angles 

var. glabra form, boliviana 

2. Corolla puberulent outside; leaves and branches 

puberulent var. hassleriana 

1. Calyx-lobes free to the base, broadly ovate. 

6. Young branches subterete, densely pubescent; leaves 
more or less pubescent above; pedicels densely 
pubescent; ovary glabrous or nearly so; calyx-lobes 
glabrous or with a few hairs near the base of margins; 
corolla glabrous outside var. paranensis 

6. Young branches compressed, minutely puberulent; 
leaves glabrous or nearly so above; pedicels 
minutely puberulent; ovary minutely puberulent to 
glabrous; calyx-lobes sparsely minutely puberulent 
to glabrous except ciliolate margins; corolla more 
or less puberulent to glabrous. . . var. chrysoderma 



358 PHYTOLOOIA Vol. 17, no. 5 

M. cordifolia Mart. Denkshr. K. Acad. Muench. 9: 95, t.7 
1824. 

Var. cordifolia . Figs. 10-16. 

M. asperula Benth. Linnaea 23: 444. 1850. Type: Regnell I: 
367 (K) 1 

M. attenuata Nees & Mart. Nov. Act. Nat, Cur. 12(1): 14. 
1824. 

M. cordifolia var. attenuata (Nees d Mart.) Wernh. Journ. 
Bot. ^7 Suppl. 20. 1919. 

M. grandiflora Miq. Linnaea 22: 803, 1849. Type: Blanchet 
3600 1 

Guagnebina ignita Veil. Flor. Plum. 45, I. t. 115, 1825. 

M. ignita (Veil. ) Schum, var. cordifolia Schum. Mart. Flor. 
Bras, 6(Fn 111. 1889. 

M. leianthiflora Griseb. Abh. Wiss. Goett. 19: 159. 1874; 
Plant. Lorentz. 111. 1874, Type: Lorentz 365 1 

M. Stipulosa Wernh. Journ. BotT 57 Suppl. 21, 1919. Type: 
Gardner s.n. (K) ! 

Branches subtetragonal to subterete, striate, densely pub- 
escent to glabrous; stipular sheath about 1 mm. high, pubescent 
to glabrous; free portion of stipules erect, deltoid to subulate, 
mostly 1-2 mm. long rarely 3-4 mm. long, pubescent to glabrous, 
toothed; leaf-blades membranous to chartaceous, ovate to ovate- 
lanceolate, rounded to attenuate at the base, acuminate to long- 
attenuate at the apex, with 3-5 lateral veins on each side of 
the midvein, densely pubescent on both sides to nearly glabrous 
except on the veins beneath, 2.5-12 (14,5) cm. long, 1-6.5 (9) 
cm. wide; petioles 2-15 (rarely up to 40) mm. long, pubescent; 
upper leaves on flowering branchlets much smaller, nearly orbi- 
cular to lanceolate, cordate to obtuse at the base, acute to 
acuminate at the apex; flowers axillary and terminal on branch- 
lets, solitary to in cymelike inflorescences; pedicels 13-60 mm, 
long, pubescent to glabrous, naked or with small bracts near 
the middle or above the basal part; ovaries oblong to oblong- 
turbinate 3-5 mm. long, pubescent to glabrous; calyx-lobes 
connate at the base for 0,5-1.5 mm., free portions ovate-lan- 
ceolate to lanceolate, acute to acuminate, pubescent to glabrous, 
2-8 mm, (rarely 10-11.5 mm.) long and 0.5-2 (rarely 3-4) mm. wide, 
sometimes with small teeth or lobules in between; corolla 3,5-6 cm. 
long (lobes 3-5 mm. long), glabrous outside, densely pubescent 
within for 5-10 mm. near the base of the tube; anthers about 4 mm. 
long, half-exserted; filaments about 2 mm. long, inserted at the 
apex of the tube; stigmas exserted, ovate, obtuse; disk free from 
the calyx-tube; capsules mostly subcompressed ellipsoidal with 
slightly tapered apex, (6-7) 8-12 (15) mm. long, (4-5) 6-7 (8) 
mm. wide, mostly glabrous. 



1968 Chung, Studies in Manettia 309 

Brazil: Prov. Bahia: Blanchet 2092 (BM, G, NY), 3600 (BM, G, K, 
MO, P) ; Prov. Goyaz: Gardner 3769 (BM, F, G, K, NY, P) ; Prov. 
Mato Grosso: Lindman A3417 (S) , Moore 553 (BM) , Weddell 3323 (F) ; 
Prov. Amazonasl Rio Acre, Ule 986T fc, K) ; Prov. Minas Geraes: 
Brade 17862 (F) , Claussen 689 (NY, P) , s.n. (BM, G, GH , K, S) , 
Duarte"TT5~(F) , Gardner 47r7~ (BM) , 4718 (K) , Heringer 69 (F) , 
Irwin 2TIF (F, NY, UC, US) , Macedo 1685 (MO, S) , MagalKaes 3254 
(US) , Mexia 5697a (BM, F, G, GH , MO, NY, S, UC , US) , Mosen 1863 
(F, S) , 4474 (s) , Regnell I: 367 (K, S, US) , Saint-Hilaire ~2TT 
(P) , Vauthier 208 (G), Williams, L.O. & V. Assli~8218 (F, GHTT" 
Prov. Sao Paulo: Glaziou 12774 (P, US), Valio 16 (US) ; Prov. 
Parana: Dusen 3426 (F,S) , 7582 (S) , 23872 (F) , Reiss 54 (F, NY, S) ; 
Prov. Santa Catharina: Hoehne 24406 (F) , Reitz & Klein~3"401 in 
part (US), 4174 (Ny;; Ceara, Gardner 1699 in part (BM, G, GH, K, 
NY) ; Schott"~§3T (K) . 

Paraguay: vTTTarrica, Hassler 4132 in part (S) , 8841a (G) ; Villa 
Rica: Joergensen 7269 (F) ; Prov. Tobaty: Hassler 6246 (BM, G, NY, 
UC) ; Caballero, Morong 512 (NY) . 

Argentina: Prov. Misiones: Bertoni s.n.; Prov. Jujuy: Bartlett 
20396 (US) , Fries 242 (S) , Schreiter 11173 (F) , Venturi 87?7 
(BM, F, GH, K, MO, US), West 836S~ TGH, MO, UC , US) ; Prov. Salta: 
Borrea 37073 (GH) , Cabrera 3104 (F, NY) , Donell 3137, Ragonesi & 
Coras 37800 (GH) , Schreiter~g72"2 (F) , 10114 (US)7 37686 (GH) , 
Venturi 5317 (F, GH , US) , 8279 (F, US) , West 6131 (MO, UC , US); 
Prov. Tucuman: Dinelli s.rT! Tbm) , Gonzalez 21657 (G) , Lorentz 321 
(G) , Meyer 9827 (F), Schreiter 748 (US) , 2191 (F) , Venturi 1204 
(US) , 1574 TUCT US) , 4086 (GH, US) , 4244 (BM, F) , 7887 (GH, US) , 
8037 (F) , 10359 (BM, MO, S, UC) . 

Bolivia: Prov. Beni: Rurhenabaque , Fleischmann 215 (S) ; Huah- 
uanus-Reye, Cardenas 5390 (US) ; Prov. Cochabamba: Bang 1255 
(BM, F, GH, K, MO, NY, PH, US), Kuntze s.n. (US); Nord-Yungas: 
Milluguaya, Buchtien 252 in part (GH, MO) , 4723 (NY, US); Chaco: 
Cardenas 2605 (F) . 

Peru: Dept. San Martin: Ferreyra 5067 (US) , 7776 (US) , 7851 
(US) , 7911 (US) , Klug 3541 (F, G, GH , MO, S, US) , Spruce 39?9 
(K, S) , Williams LI. 7479 (F, G) , 7773 (F) , Woykowski 35259 (F) , 
35339 (F) ; Dept. Junin: Woytkowski 6333 (US) ; Pozuzo, Macbride 
4722 (F, S, US) , Pearce 215 (BM) . 



Gardner s.n. (K) , the holotype of M. stipulosa Wernh. (Journ. 
Bot. 57 Suppl. 21. 1919. Gardner s.n., K) is characterized by 
the large leafy calyx-lobes which are about 9 mm. long, 4 mm. 
wide, ovate-lanceolate, acuminate, and glabrous; glabrous petioles 
and leaf-blades except for the minutely ciliolate margins; erect 
stipular free postions lanceolate, 2-4 mm. long; and the moderately 
pubescent young branches. Venturi 7887 (GH) from Argentina and 
Dusen 9180 (BM) from Parana^ Brazil"^ aXso have similar, large, 
leafy calyx- lobes. In some of the stipules on Dusen 9180 (S) 
the free portion is as long as 4 mm. long. Although the leaf 
surfaces are glabrous, M. stipulosa Wernh is considered as a 
form of M. cordifolia var. cordifolia. 



360 PHYTOLOGIA Vol. 17, oo. 5 

M. cordifolla var. cordifolia form, incana (Schum.) Chunq , 
comb, nov. 

M. iqnlta (Veil.) var. incana Schum. Mart. Fl. P.ras, 6(6): 
171. TSBC Type: Balansa 2Tm 

M. sublanata Wernh. Journ. Bot . 57 Suppl, 21. 1919, 
Type: Hassler 8841 1 

Distinguished from var. cordifolia by the tomentum on the 
lower surface of the leaves, which is gray or yellov/ish-brown. 
Corolla 25-45 mm. long; calyx-lobes 2-4 mm. long; ovaries often 
tomentose; capsules 7-10 mm. long, pubescent; pedicels and 
branches pubescent. 

Brazil: Minas Geraes, Regnell I : 367 in part (S) . 
Paraguay: Villa Rica, Balansa" 2135 (G) ; Cordillera de Villa Rica, 
Hassler 8841 (BM, G, GH , NY, S,~ucy; Villarrica, Jorgensen 4132 
in part (F, GH, NY, PH, US). 
Argentina: Prov. Corrientes: Santo Tome, Ybarrola 1503 (S) . 

M. cordifolia var. glabra (Cham. & Schl.) Standi. Field 
Mus. Bot. 7(3) : ?61. 19 3T: 

M. glabra Cham. & Schl. Linnaea 4: 169. 1829. Type: Sellow 
Photo (BM 

M. ignita var. glabra (Cham. & Schl.) Schum. Mart. Fl. Bras. 
6(6): 171. 1889. 

M. micans Poepp & Endl . Nov. Gen. & Sp. 3: 24. 1845. 

M. ignita var. micans (Poep. & Endl.) Schum. Mart. Fl. Bras. 
6(6): 171. 1889. Poeppig 2415 (G) ! 

M. cordifolia var. f iliformis Wernh. Journ. Bot. 57 Suppl. 20 
1918. Typil Fiebrig 4636 ! 

Branches, petioles, leaf-blades, pedicels, and calyx-lobes 
glabrous. 

Brazil: Prov. Bahia: Blanchet 3281 (G) , Rose 20034 (US); 
Prov. Mato Grosso: Kuntze s.n. (NY, US) ; Prov. Sao Paulo: 
Loefgren 1252 (F) , Weir 192 in part (K) ; Prov. Parana: Dusen 9059 
(S) , 9180 in part (PHTT 11758 (S) , 16171 (F, GH , MO, S, US) , 
Hatschbach 3160 (US), 3763 (US) ; Prov. St. Catharina: Schwacke (F) , 
Tweed ie TghT! 

Uruguay: Dept. Artigas: Herter 1157 (F, G, MO, NY, UC, US) , 
Rosengurtt B-3663 (US); Dept. Salto: Osten 5451 B (US); Dept. 
Paysandi: Calot 94 (P) , 95^ (P) ; Concepcion, Lorentz anno 1895 
(GH) ; Fray Bentos, Fruchard (P) ; Islands of the Uruguay River, 
Tweed ie (K) . 

Paraguay: Between River Apa and River Aquidaban, Fiebrig 4636 
(BM, G, GH) ; Upper River Apa, Hassler 8327 (BM, F, G, GH , MO, NY, 
S, UC) ; Sierra de Amambahy, Hassler 11200 (BM, G, GH) ; Campo 
Duarte, Hassler 1237 (G, K) ; Yerbalium de Maracayu, Hassler 4435 
(BM, F, G, NY) ; Central Cordillera, Hassler 7026 (BM, G, NY); 
La)te Ypacaray, Hassler 11783 (BM, F, G, GH, MO, NY, S, UC, US); 
Paraguari, Balansa 2134 (G, K) . 



1968 Chung, Studies in Manettia 361 

Argentina: Prov. Misiones: Bertoni 1485 (UC, Ekman 1383 , (MO, 

NY, S) , Gruener 35 (F) , Meyer 5319 (F) , Rodriguez 337 (F) , Spega - 

zzini 20715 (F) ; Prov. Corrientes: Bonpland 599 (P) , Ibarrola 

1872 (S) , Meyer 8972 (S) , Wurth 74 (S) ; Prov. Entre Rios: Lorentz 

107 (PH) , s . n . ( PH ) , Tweedie s.n. (BM, K) . 

Bolivia: Dept. Cochabamba: Steinbach 9055 (GH) ; Dept. Santa 

Cruz: Maguire 44492 (NY), Steinbach 6240 in part (G, GH) ; S. Bolivia, 

Chignica, Fiebrig 2686 in part (BM) . 

Peru: Dept. Loreto: Rio Marano Valley, Killip, Smith & Dennis 

29200 (F, US); Yurimaguas , Killip & Smith~7990 (F, NY, US), Poeppig 

2415 (G) ; Balsapuerto, Klug~7?80 (BM, F, G, MO, S, US); Tarapoto, 

Mathews 1343 (BM, F, G, GH) ; Haallaya River, Spruce 4592 (BM, F, G, 

GH, K, NY) ; Dept. San Martin: Ferreyra 4521 (US) , Klug 2606 (BM, 

F, G, GH, MO, NY, S, US), 4075 (BM, F, GH , MO, NY, S, UC) , Woytkowsl<^i 

7242 (MO, US); Dept. Huannuco: Killip & Smith 26823 (F, NY, US) ; 

Dipt. Junin: Ferreyra 3603 (US) , 3678 (US) , Killip & Smith 23519 

(NY, US), 24748 (F, NY, US) , 25213 (US) , Macbride 5585 (F, US) , 

Schunke 480 (F, S) , 1430 (F) , 1519 (F) , Woytkowski 395 (US) ; 

74 51 (USFfPept. Cuzco: Sandeman 3646 (K) , Vargas 1843 (GH, MO) 

3791 (US) , 15430 (MO) . 

The following specimens are rather intermediate between 
var. cordifolia and var. glabra in that the leaves are glabrous 
but the young branches are pubescent. Brazil: Bahia: Blanchet 
3281 in part (BM, F, NY, P) ; Ceara, Loefgren 587 (S) ; Mato Grosso: 
Malme 174 (F, S) . Argentina: Corrientes: Ibarrola 1234 (S) . 

M. cordifolia var. glabra form, boliviana (Wernh.) Chung, 
comb . nov . 

M. boliviana Wernh. Journ. Bot. 57 Suppl . 20. 1919. 
Syntypes: Bang 1372 1 , Bridges 1 

Tetragnal branches with short reflexed hispid hairs or recur- 
ved teeth on the narrow wings. 

Brazil: Bahia: Lemos Froes 20165 (US); Espirato Santo: Freire 
68 (F) ; Mato Grosso: Kuntze 92 in part (NY) ; Ceara, Gardner 1699 
in part (P) . 

Bolivia: Dept. Santa Cruz: Brooke 5774 (BM, F, NY), Kuntze (NY), 
Steinbach 6240 in part (G) , 7092 (BM, F, G, GH , MO, NY, PH, S, UC) , 
Yapacani, Kuntze (NY); Dept. Cochabamba: Cardenas 3135 (F, US); 
Dept. La Paz: Mapire, Rusby 1126 (BM, F, GH , K, MO, NY, PH) , 
Buchtien 1449 (S) ; Guanai, Bang 1372 (BM, F, G, MO, NY, PH, US), 
Rusby If 27 (F, NY); Nordyungas, Buchtien 262 in part (BM, F, G, GH , 
NY) ; Chignica, Fiebrig 2686 in part (G, K, S, US); Gran Choca, 
Fries 1372 (S) . 

Argentina: Prov. Jujuy: Bartlett 20341 (US) , Hunziker 1974 
(GH) , Spegazzini 381 (F) ; Prov. Salta, Dept. Oran : Hunidoboro s.n. 
(S, UC) , Meyer 5072 (UC) , Pierotti s.n. (S) , Spegazzini 14280 (F) . 
Peru: Dept. Cuzco: Storke, Horton & Vargas 10465 (F, UC) . 



362 PHYTOLOGIA Vol. 17, no. 5 

M. cordifolia var. haasleriana (Chod.) Chung, comb, nov, 

M. haaalorlana Chod. Bull. Herb. Boiss. 7 App. 1: 82. 1899. 

Type: Haaaler 2509 (from near Rio Apa. , Paraguay, not see) 
M. angustifolia Wernh. Journ. Bot. 57 Suppl. 22. 1919. 

Type : Fiebrlg 5734 1 

Branches subterete, striate, puberulent; leaves lanceolate 
to ovate-lanceolate, acute to rounded at the base, attenuate- 
acuminate at the apex, 35-65 mm. long, 10-24 mm. wide, puberulent; 
calyx-lobes ovate-lanceolate, more or less narrower at the base, 
acuminate at the apex, 5-8 (12-13) mm. long, 2-5 mm. wide, puber- 
ulent; corolla 3-4 cm. long, moderately puberulent with short 
broad hairs; inside of corolla glabrous except for a band (6mm. 
long) of dense hairs in the narrow tube about 4 mm. above the 
base; filaments about 2 mm. long, inserted at the apex of the 
tube, anthers 3.5-4 mm. long; stigmas ovate, obtuse; capsules 
oblong to oblong- turbinate , 6-7 mm. long. 

Brazil: Parana: Duarte 1912 (F, NY); Sta. Catharina: Dusen 11894 
( GH , MO , S ) . 

Paraguay: River Alto Parana, Fiebrig 5734 (BM, G, GH, K, US). 
Argentina: Misiones: Iguazu Falls , Sandeman 4770 (K) , Smith 355 
(K) ; Dept. Iguazu: Rodrigo 3681 (F) . 

M. cordifolia var. paranensis (Standi.) Chung, comb. nov. 
M. paranensTs Standi, Field Mus. Bot. 8(5): 331. 1931. 
Type : Dusen 8964 1 

Distinguished from var. cordifolia mainly by the large 
leafy calyx- lobes which are free to the base; short, turbinate 
ovaries 1-2 mm. long; small subglobose capsules 5-6 mm. long. 
Branches subterete, densely pubescent; stipular sheath 0.5-1 mm. 
long; often ill defined, free portion of stipules 1.8-2.5 mm. 
long, triangular-lanceolate, often bilobed at the apex; petioles 
7-13 mm. long, pubescent; leaf-blades ovate, 2.8-5.5 cm. long, 
1-3 cm. wide, sparsely pubescent, with 2 or 3 lateral veins 
on each side of the midvein; calyx-lobes broadly ovate, acute to 
acuminate, 8-11 mm. long, 4-5 mm. wide, 3-nerved, glabrous or 
with a few hairs near the base of the margins; corolla 3.5-4.2 cm. 
long, glabrous outside. 

Brazil: Parana: Serra do Mar, Dusen 8964 (G, GH , NY, S, US), 
9007 (S) , Morretes, Hatschbach 1741 (S) ; Piraquara, Hatschbach 
2840 (US); Guaratuba, Hatschbach 6574 (US). 

M. cordifolia var. chrysoderma (Sprague) Chung, comb. nov. 

M. chrysoderma Sprague, Bull. Herb. Boiss. II. 5: 264. 
1905. Ty^el Mueller 123 (K) 1 

M. paulina Standi. Field Mus. Bot. 8(5): 328. 1931. 
Type: Saint-Hilaire 12291 



1968 Chung, Studies in Manettia 363 

Young branches compressed, puberulent; free portion of 
stipules deltoid to round and apiculate; leaf-blades ovate to 
ovate-lanceolate, rounded to acute at the base, acuminate at the 
apex, 4-6 (-11) cm. long, 1.8-3 (4) cm. wide, glabrous or nearly 
so above, puberulent beneath mainly on the veins, with 4 or 5 
lateral veins on each side of the midvein; pedicels more or 
less puberulent; calyx-lobes free to the base, ov?te to ovate- 
lanceolate; acute to acuminate, 4-5 mm. long, 2-3 mm. wide, more 
or less puberulent to glabrous except the ciliate margins; 
corolla 35-55 mm. long, 6-9 mm. wide, more or less puberulent 
with short broad hairs or glabrous outside. 

Brazil: S. Paulo: Edwall 1900 (F) , Krieger anno 1863 (NY) , 
Saint-Hilaire 1229 (F, P) ; Parana: Dusen s.n. (GH, MO) , Glaziou 
(P) , Hatschbach 4136 (US); Sta. Catherina: Mueller 123 (K) , Smith 
& Kleirr^450 (NY, US) , Reitz 4363 (US) , Reitz & KTe'in~T028 
(S, US) , 4700 (US), 5218 (UC,~USr. 



Acknowledgement: I wish to express my appreciation to the 
curators at the institutions indicated for making it possible 
for me to study the collections in their herbaria. The illus- 
trations have been prepared by Mr. Walter L. Boyer of Field 
Museium of Natural History after the author's drawings. 



36b 



PHITOLOOIA 



Vol. 17, r>o. 5 




Fig. 1. Manettia domingensis , flower, x4 ( Ekman H6284 , S) 

Fig. 2, Manettia tweedieana , flower, x4 ( Tweedie^ GH) 

Fig. 3. Manettia gracilis , flower, x4 ( Reitz & Klein 4115 , G) 



1968 



Chung, Studies in Manettia 



365 




Figs. 4-9. Manettia pubescens . 4: flower, x2 ( Macedo 2895 , PH) , 
5: ovary and calyx, x2 ( Damazio 966 , G) , 6: fruit, x2 (Damazio 
966 , G) ; 7-9: stipules, xlO (7; 
I: 368, S; 9: 



stipules, xlO (7 : Regnell Ij 
Macedo 2895, PH) . 



368, K; 8: Regnell 



366 



PHYTOLOGIA 



Vol. 17, no. 5 




Figs. 10-16 

xlO (10: Dusen 3426, S ;11 



Manettia cordifolia var. cordifolia. 10-12: stipules, 



Gardner, K, M. 



Dusen 9180 , S) 13-14: calyx-lobe, x4 (13 

Gardner, K, M. stipulosa) , 15: flowering branchlet, x 



stipulosa ; 12; 
Venturi 7887, GH; 14: 

(Mosen 1863, 



S) , 16: ovary and calyx-lobes, x4 ( Gardner 1699 ,K) 



TRANSFERS TO GUAPIRA FROM TORRUBIA (NYCTAGINACEAE) 

Elbert L. Little, Jr. 



Eight new combinations in Guapira (family Nyctaginaceae) are 
made here for forthcoming publications on common trees of the 
United States, Puerto Rico and the Virgin Islands, Province of 
Esmeraldas in Ecuador, and Venezuela. 

Guapira Aublet (Hist. PI. Guian. Franc. 1: 3O8; 3: t, II9 . 
1775) with a single species (G. guianensis ) Tra.s of uncertain posi- 
tion until identified as a synonym of Pisonia by Hallier (Leiden 
Meded. Herb. 35: 18-20. I918) . The genus Guapira was not 
accepted by a second author until I96I, when revived with 1 new 
species and 1 new combination by Woodson in Woodson and Schery 
(Fl. Panama k (k) : 403-^+06, fig. 12^; Mo. Bot. Gard . Ann. kQ: 6I- 
6k, fig. 12^. 1961). Lundell (Wrightia 3: 22. 1962) trans- 
ferred 2 species to Guapira , and Miranda (Soc. Bot. Mdx. Bol. 29: 
3h. 1965), 1. 

My proposal (No. IU9) to conserve the generic name Torrubia 
Vellozo was submitted in August I963 and published in the pro- 
posals for the Tenth International Botanical Congress at Edin- 
burgh (Regn. Veg. 3U: 58-59- 1964). Now the Committee for 
Spermatophj'ta (Taxon 17: 462-463. I968) by a vote of 1-9 "de- 
clines to recommend Torrub ia for conservation, chiefly because 
the identity of the type-species is uncertain and probably will 
remain so. No type- specimens of Vellozo are kno-\-m, and it is 
impossible to identify his plant from the protologue." However, 
many of Vellozo' s names have been typified and taken up. The 
type species, Torrubia opposita Veil., was illustrated by a draw- 
ing of foliage with male flowers and ^ras described from coastal 
forests at Rio de Janeiro, Brazil. My suggestion that it may be 
the same as T. olfersiana (Link, Kl. & Otto) Standi, var. nitida 
(Mart.) Reitz (Sellowia 12: I69. I96O) could be checked by a 
local specialist. About 50 names must be transferred to Guapira . 

Three species of small trees native in southern Florida 
(Little, Check List Native Naturalized Trees U.S. 422. 1953.)are: 

GUAPIRA BRACEI (Britton) Little, comb. nov. Brace blolly 

Torrubia brace i Britton, Torrey Bot. Club t^uI. 3I: 6l4. 1904. 

GUAPIRA GLOBOSA (Small) Little, comb. nov. roundleaf blolly 
Torrubia globosa Small, Man. Southeast. Fl. 49O, 1504. I933. 

GUAPIRA LONGIFOLIA (Heimerl) Little, comb. nov. longleaf blolly 
Pisonia discolor y longifolia Heimerl in Urban, Bot. Jahrb. 

21: 627. 1896. 
Torrubia longifolia (Heimerl) Britton, Torrey Bot. Club. Bui. 

367 



368 P H T T L G I A Vol. 17, no. 5 

31: 6lh. 190U. 
Plsonla longifolla Sarg., Man. Treec No. Amer. 3IU, fig , g^l . 

1905. 
A fourth speciec Torrubla florldana Britton (Torrey Bet. Club- 
Bui. 3I: 615. I90U) was described ac a low chrub. It was col- 
lected at Rock Key near Key West more than a century ago and has 
not been found since. 

The three specLeE listed below are trees native in Puerto Rico. 
Two \rere recorded by Britton and Wilson (Sci. Surv. Porto Rico 
Virgin Is. 5: 286-287. 192^1) and by Little and Wadsworth (Common 
Trees Puerto Rico Virgin Is. 92-93, fig . 196^^). Alain Liogier 
(Rhodora 67: 329. I965) has suggested the transfer to Guapira . 
The third species is knovm from southwestern Puerto Rico also. 

GUAPIRA DISCOLOR (Spreng.) Little, comb. nov. barrehomo 
Pisonia discolor Spreng., Syst. Veget. ed . I6, 2: I68. I825. 
Torrubia discolor (Spreng.) Britton, Torrey Bot. Club Bui. 31: 
613. I90U. 

GUAPIRA FRAGRANS (Dum.-Cours . ) Little, comb. nov. 

corcho, black marapoo 
Pisonia fragrans Dum.-Cours., Bot. Cult. ed. 2, 7: 11^- l8l4. 
Torrubia fragrans (Dum.-Cours.) Standley, U.S. Natl. Herb. 
Contrib. I8: 100. I916. 

GUAPIRA OBTUSATA (jacq.) Little, comb. nov. 

Pisonia obtusata Jacq., PI. Rar. Hort. Caes. Schoenbr. 3^ 35 > 

t. 31'^ 1798 . 
Torrubia obtusata (jacq.) Britton, Torrey Bot. Club Bui. 3I: 
612. 1904. 

The next species was found in I965 in the Province of Esmer- 
aldas, Ecuador. It was described from Peru, where it i^as reported 
to be the only species of Torrubia (Standley in Macbride, Fl. 
Peru pt. 2 (No. 2): 528. I937) • 

GUAPIRA MYRTIFLORA (Standi.) Little, comb. nov. 

Torrubia myrtiflora Standi., Field Mus. Pub. Bot. 8: 307. I93I. 

The last i-ra.s selected as an example of the genus in a book on 
common trees of Venezuela noA'? being edited for publication. Seven 
species of Torrubia were listed from Venezuela by Pittier et el . 
(Cat. Fl. Venez. 1: 29O-29I. I9U5) . 

GUAPIRA PACURERO (H.B.K.) Little, comb. nov. pacurero 

Pisonia pacurero H.B.K., Nov. Gen. Sp. 2: 2l8. I8I7. 
Torrubia pacurero (H.B.K.) Standley, U.S. Natl. Herb. Contrib. 
13: 101. 1916. 

Forest Service, United States Department of Agriculture, Wash- 
ington, D.C. 20250. 



REVIEW OF F. E. WIMIER, CAMPANULACEAE-LOBELIOIDEAE SUPPLEKENTUM 

ET CAMPANULACEAE-CIPraOIDEAE. DAS PFLANZENEEICH, IV. 276c (108. 

HEFT), I - X, 816 - 102U; WITH DESCRIPTION OF TREUATOLODELU 

WIMMERI DEG. & DBG., SP. NOV. 

Otto & Isa Degener 
Volcano, Hawaii 

Shortly before his death on May 2, 1961, Dr. Franz Elfried Wim- 
mer submitted his ccanpleted manuscript about Lobelioideae and 
Cyphioideae to Dr. K. H. Rechinger. The latter then sent the work 
to Drs. H. Stubbe and S. Danert. It was published on March 15, 
1968, in East Berlin. Of the Lobelioideae there are 29 plates and 
11 figures J of the Cyphioideae , $1 plates. The drawings, probably 
reproduced in the same size as executed by the illustrator instead 
of being reduced by half, are not as good as the photographs, 
those borrowed from the late Dr. J. F. Rock being outstanding. 

As we are not familiar with the Cyphioideae, we shall not re- 
view the almost 100 pages devoted to them. In fact, we shall limit 
ourselves to the Lobelioideae so far as represented in the Hawaiian 
Islands. Dr. trimmer lists the following genera endemic to these 
islands, with the number of species known up to his time, as: 

Brighamia 1 Delis sea __-_ 8 

Clermontia 32 Rollandia 12 

Cyanea Ih Trematolobelia 3 

For the presumably cosmopolitan genus Lobelia , he lists 388 species 
for the world. 

Being a bit less conservative perhaps than Dr. Wimmer, we prefer 
Lobeliaceae to Lobelioideae ; and so far as the genus Lobelia is 
concerned, do not recognize it as native to the Hawaiian Archipel- 
ago. Instead, we prefer to place most of the taxa reposing there 
into three small, endemic genera. 

As Hawaiian place names are confusing in their spelling and as 
plant labels, particularly ours, are often a bit illegible in script, 
we here wish to put on record some necessary orthographic changes: 

Page 817, for Kanehaha read Kanahahaj for Hononau, HonaTinau. 

Page 818, for Anny Greenwell read Amy Greenwell. 

Page 820, for McKandles read McCandless . 

Page 823, for thelephone read telephone. 

Page 825, for Pololo read Pololu; for Maunakui read Mauna Huij for 

Kapoho Puna read Kapoho, Puna. 
Page 826, for Papaiku read Papaikouj for Kala, Kikalaj for Pitso, 

Piko; for Jao, lao; for Kaulelewelewe, Kaulalewelewej for 

369 



370 PHTTOLOOIA Vol. 17, no. 5 

Pololo, Pololu; for Honokanenul, Honokane Nul; for Pololo, 

Pololu. 
Page 828, for Komakawei read Komakawai; for L'cCandlea, ycCandless . 
Page 829, for Olau read Olaa; delete Kohala before Kulanij for 

Pololo read Pololu; for Honokanenul, Honokane !^il; for Ann;' 

Greenwell, kmy Greenwell. 
Page 831, for Kawaihe read Kawaihae. 
Page 887, for Honokanenul read Honokane Nuij for Kaholuamano, 

Kaholuamanu; for Hamakue, Hamakua. 
Page 888, for Lehua makanoe read Lehuamakanoi . 
Page 892, for Waiahuattia read Waiahxiakua. 
Page 901, for Fam read Fern. 
Page 906, for liiray read Hirai. 
Page 909, for Hetheway read Hatheway. 

Regarding lobelias in the Hawaiian Islands, we are convinced 
many new taxa still exist; but most of these probably will be ex- 
terminated before they can be collected by the botanist due to 
the ravages of man's bxill dozing, his agricultural and timber in- 
dustries, his livestock raising, his building boom with apparent- 
ly a desert-like golf course next to every tourist hotel, and his 
introduction by accident and design of exotic plants and animals 
injurious to the endemic biota. One of these many taxa on the 
verge of extinction we here name, 

TREMATOLOBELIA WHflffiRI Deg. & Deg., sp. nov. Lobi calycini 7 ma . 

longij capsula 1$ mm. longa et 16 mm. lata. 
Trematolobelia macros tachys sensu Fagerlund & Mitchell, Checklist 

Plants Haw. Nat. Park Kilauea - Mauna Loa Sect. 58. 19Uh' 
Trematolobelia macrostachya (sic) sensu Fosberg; Doty ?c Mueller- 

Dombois in Haw. Bot. Sc. Paper 2: 231. 1966, 
Not Trematolobelia macrostachya Zahlbr.; Rock in Coll. Haw. Publ. 

2: U5. 1913. 
Trematolobelia kauaiensis sensu 7/immer in Pflanzenreich IV. 276c 
(108. Heft). 901. 1968. (As to Island of Hawaii only). 

Plant with single slender erect stem 2 meters tall. Leaves ob- 
lanceolate, about lU cm. long and 2 cm. wide, glabrous throughout, 
acuminate to sessile base, sharply cuspidate at apex, faintly un- 
dxilate with submarginal hydathode at each indentation. Flowers 
about 20 per horizontal 30 cm. long branch of inflorescence, with 
pair of bractlets at lower third of pedicel: in bud with hypanth- 
ium 3 mm. long and 2 mm. wide; with calyx lobes 5 nm. long amd 
almost 2 mm. wide, oblong, obtuse at apex but with faint cusp; in 
anthesis (flower in poor, decayed condition) with staminate column 
and style and stigma probably about 6 cm. long. Capsule on 
thickened 3 cm, long pedicel, 20 mm. wide, 13 mm. high without 
the persistent somewhat incurved 5 mm. long calyx lobes. 

Type locality: Hawaii, Kilauea, near Fern Forest, rich moist 
8\inny locality; only this one seen. Degener 7860, Februajry 18, 
1922. (Vienna). 



1968 Degener & Degener, Review 371 

The type, collected by Otto Degener in 1922, uras not available 
for study when the writers visited the Natural History Musevun, 
Vienna, in the summer of 196U. Even though the type specimen 
normally must have died after fruiting, the Degeners combed the 
type area in August 1968 with the hope of perhaps discovering an 
offspring of the 1922 plant. Though the area had escaped the 
usual ravages of "civilization" in the vicinity, no Trematolobel- 
ia plants were found. Three sheets (Fagerlund &. Mitchell 8ii7) in 
the Hawaii Volcanoes National Park herbarium, however, evidently 
belong to this taxon, are considered cotypes, and here have been 
used to augment the description. Fagerlimd & Mitchell collected 
the young flowering material September h, 19li3, and fruiting ma- 
terial from the same specimen February 22, 19kh» As the sheets 
cite the locality as being "In wet forest between Crater Rim 
road suxd Kilauea Iki," the writers visited the area in the hope 
of finding specimens. The search was of no avail - the area had 
been devastated by the 1959 Kilauea-Iki Eruption I In place of 
Trematolobelia , the unwelcome exotics Anemone japonic a , Buddleja 
asiatica and Rubus penetrans were taking over the area. We fear 
Trematolobelia wimmeri Deg. & Deg., a species with capsules 
reminiscent in size to those of T. kauaiensis (Rock) Skottsb., 
to be on the verge of extinction if not already extinct* 



ADDITIOHAL NOTES ON THE ERIOCAULA.CEAE . HV 
Harold N. Uoldenke 



ERIOCAUUCEA.E Undl. 

Additional & emended bibliography: Petiv., Gaz. pi. 6, fig. 2. 
1702; Pluk,, Aim. pi. U09, fig. 5. 1769; Lam., Encycl. 3: 276. 
1789; Willd. in L., Sp. PI., ed. h, 1: U86. 1797; L'-ichx., Fl. 
Bor.-am. 2: 165. 1803; Pursh, Fl. Am. Sept. 1: 91. l8lii; Roem. ^ 
Schult. in L., Syst, Veg., ed. 15 nova, 2: 86I4. 1817; Nutt., Gen. 
1: 90. 1818; Ell., Sketch Bot. 2: $6$. 182U; Wall., Plant. As. 
Rar. 3: 28. 1832; Wall., Numer. List 207—208 ["207"]. 1832; 
Beck, Bot. 370. 1833; Benth. in Hook., Niger Fl. 5U7. 18U9; Hook, 
f ., Fl. Brit. Ind. 6: 571—585. 1893; Jacks, in Hook. f. & Jacks., 
Ind. Kew., pr. 1, 1: 877—880. 1893; Britton & Br., 111. Fl., ed. 
1, 1: 371—373, 602— 60li, & 6U, fig. 899—903 (I896) and 3: 536, 
537, 51a, 5U5, & 577. 1896; Ruhl. in Engl., Pflanzenreich 13 (IV, 
30): 1—108. 1903; R. M. Harper, Ann. N. Y. Acad. Sci. 17: 267— 
268, pi. 2U, fig. 1. 1906; Alv. Silv., Archiv. Mus. Nac. Rio Jan. 
23: 162, pi. U. 1921; Fern., Rhodora U8: iv S: 58. 19U6: Jacks, in 
Hook. f. & Jacks., Ind. Kew., pr. 2, 1: 877—880 (19i;6) and pr. 
3, 1: 877—830. I96O; B. G. Eriggs, Contrib. N. S. Wales Nat. Herb. 
U: 2k & 26. 1966; Anon., Assoc. Etud. Tax. Fl. Afr. Trop. Bull. 
17: 19. 1966; G. L. Davis, Syst. Einbryol. Angiosp. 1966; K. Lar- 
sen, Dansk Bot. Arkiv 23: 375 — 399. 1966; C. G. Tovmsend, Excerpt. 
Bot. A. 10: 310. 1966; Anon., Assoc. Etud. Tax. Fl. Afr. Trop. In- 
dex 1965: 31. 1966; S. V. Ramaswami, Study Flow. PI. Bangalore 
[thesis] 219—221 & lli06— lii07. I966; Goodland, Bol. Soc. Venez. 
Cienc. Nat. 26: 3i;5. 1966; Klots, New Field Book Freshw. Life 91^. 
1966; Begum, Cvirr. Sci. [India] 35: 262—263. 1966; R. H. CoBq)ton, 
Journ. S. Afr, Bot. Suppl. 6: 19, 33, & 92. I966; Subramanyam k 
Henry, Bull. Bot. Surv. India 8: 211; . I966; Sebastine & Ramamur- 
thy. Bull. Bot. Surv. India 8: 182. I966; J. L. Ellis, Bull. Bot. 
Surv. India 8: 329 & 339. 1966; 0, D. Evans, Biol. Abstr. U8: 
1;562 Sc li563. 1967; Soukup, Biota 6: 359. 1967; Anon., Pollen & 
Spores 9: 61*2. 1967; Krai, Biol. Abstr. U8: 3190. 1967; Anon., 
Ind. Bibliog. Bot. Trop, h (1)» 53 & 88. 1967; T, H, Harrison, 
Biol, Abstr, I48: 8707. 1967; Anon., Biol. Abstr. U8: 3190 & li563 
(1967), Ue (10): S,60& S,117 (1967), and U8 (22): S,65. 1967; 
Moldenke, Biol. Abstr. h&: xxii & 10099 (1967) and U8 (20): S, 61, 
S,l6l, S,l65, & S.I83. 1967; Dombrowski & Kuniyoshi, Araucariana 
1: 15 & 18, 1967; J, de J. Jimenez, Archiv. Bot. & Biogeog. Ital. 
k3'' U. 1967; Beguai, Bioresearch Index 1967: 2255. 1967; Anon., 
Assoc. Etud. Tax. Fl. Afr. Trop. Bull. 18: Ii5. 1967; Koldenke, 
R4stm6 Suppl. 15: [1]— 5, 8, 10, 12, lU, 20, & 21. 1967; W. G. 
Burger, Fam. Flow. PI, Ethiop. 132. 1967j Sculthorpe, Biol. Aquat. 
Vase. PI. 23, 389—391, 393, & 39U. 1967; L. V. Barton, Bibl. 
Seeds 782. 1967; Satake, Nat. Sci. ^ Mus. 3ii: I6I & 162. 1967; 
Fulling, Ind. Bot. Record. Bot. Review I78. 1967; R. M. Harper, 
Castanea 32: 17. 1967; Rickett, kVild Fls. U, S. 2 (1): 135, pl. 

372 



1968 Moldenke, Notes on Eriocaulaceae 373 

27 (1967) and 2 (2): 659 & 666. 1967; Friedrich-Holzhanmer in 
Merxmtaier, Prodr. Fl. Sttdw. Afr. 1^9: 1—2. 1967; Berhaut, Fl. 
Sln^gal, ed. 2, 311. 1967; J. ^ A. R^rnal, Adansonia ?: 329. 1967; 
L. S. Thomas. Pine Barrens 23. 1967; D, A. Livingstone, Ecol, 
Uonog. 37 (1): U3. 1967; L. 0. Williams, Fieldiana Bot. 31: 2U9— 
269. 1967; Anon,, Assoc. Etud, Tax. Fl. Afr. Trop. Index I966: 9 
(1967) and 1967: 31. 1968; Cronquist, Evol. & Class. Flow. PI. 
335, 336, & 390. 1968; F, A. Barkley, Outline Classif . Organisms, 
ed. 2, 10. 1968; Moldenke, Biol. Abstr. U9: U88 (I968) and k9 
(9): S.58. 1968; Anon., Biol. Abstr. ii9: 1975 (1968) and k9 (7): 
S.58, S.I33, i: S.180. 1968; Moldenke, R6sum5 Suppl. 16: [1], 2, 
5—9, 12, 19, 21, 23, 25—27, & 30. 1968; R. M. White, Irish 
Naturl. Joum. 16: UO. I968; Meikle, Kevr Bull. 22: Ha—lhh. 
1968; Justice & Bell, Wild Fls. N. C. 13 & 209. 1968; Moldenke, 
Phytologia 17: 3U8— 352. 1968; Fassett, Index Rep. Fl. Wise, [1]. 
n.d. 

Burger (1967) inforrus us that in this fanily "a pistillode [is] 
often present in starainate flowers". Airy Shaw (I966) states 
that the genus Reilia Steud. may belong in either the Eriocaula- 
ceae or the Juncaceae; he also tells us that the RrLocaulaeeae 
was included by Bentham &: Hooker in a "Series" called Glumaceae . 
Tomlinson (I96U) compares the Eriocaulaceae with the genus 
Aphyllanthes in the Liliaceae . Runner (I96I) places the genera 
Streptolirion and Juncoides in the Eriocaulaceae by the apparent 
error of omitting the name of family "31" between families "30" 
and "32". Tanayo (I96I) places Leucothog venezuelensis A, C, 
Sm. in the Eriocaulaceae instead of in the Ericaceae . Larsen 
(1966) reports the chromosome numbers for seven species in this 
family from Thailand. 

BUSTOCAULON Ruhl. 

Synonymy: Blastocaular Angely, Fl. Bacia Paran. 22: 31, sphalm, 
1962. 

Additional bibliography: Ruhl. in Engl,, Pflanzenreich 13 (IV, 
30): 223. 1903; Moldenke, Known Geogr. Distrib. Erioc. 7, 28, 31, 
39, Ui, 52, 53, 55, fie 59. I9U6; Moldenke, Phytologia h: 338. 
1953; Angely, Cat. Estat. 10: [2]. 1956; Angely, Fl. Paran. 10: 6, 
7, 9, 4 10. 1957; Moldenke, R6sum« 87, 237, 279, 281, 285, 292, 
323, 327, 328, 33U, 352, U02, & 1;79. 1959; Angely, Liv. Gen. Bot. 
Bras. 19 & 39. i960; Angely, Fl. Bacia Paran. 22: 3I. 1962; Heg- 
nauer. Chemotax. Pfl. 2: 153. 1963; Moldenke, R6sum5 Suppl. 7: 7 
(1963) and 12: 11. 1965; F. A. Barkley, List Ord. Fan. Anthoph. 
113 £c lli5. 1965; Airy Shaw in Willis, Diet. Flow. Pi., ed. 7, 
138. 1966. 

The genejric name is taken from the Greek words, fiXccOTC^S^aind 
'tw'vKci , meaning "young branchlet stems" because the stems pro- 
duce small branchlets, 

BLAST OCAULON ALBIDUM (Gardn.) Ruhl. 

Additional bibliography: Moldenke, Known Geogr. Distrib. Erioc. 
7, 23, hh, & ^$* 19li6; Moldenke, Phytologia I4: 338. 1953; Moldenke, 



37U PHYTOLOGIA Vol, 17, no. 5 

RfiBumS 87, 279, 285, 323, 33U, & h79. 1959. 

Additional citations i BRAZIL: Minaa Geraist G. Gardner 5273 
(B — isotypo, N— isotype). MOUITTED ILLUSTRA.TIONS: drawingsTliotes 
by KWmicke (B). 

BLASTOCAULON PROSTRATUM (KOrn.) Ruhl. 

Additional bibliography: Moldanke, Knoirn Geogr, Distrib. Eri- 
ocaul. 7, 28, 31, 52, & 55. 19U6} Moldenke, Rfiaun^ 37, 281, 327, 
33U, & U79. 1959. 

Pereira reports that this plant grows in pure stands with no 
other plants in the formation. The species has been collected in 
anthesis in May. 

Additional citations: BRAZIL: Minas Gerais: Martius s.n, [Cabo 
Agosto; Macbride photos 18733] (B — isotype, Mu— 292— isotype, N — 
photo of isotype, W — photo of isotype) j E. Pereira 2802 [Pabst 
3638] (Bd— 38U7, Z); J. E. Pohl s.a. (ku--293) . 14GUin:ED ILLUSTRA- 
TIONS: drawings & notes 'Ey Ktfmicke (B). 

BUSTOCAULON RUPESTRE (Gardn.) Ruhl, 

Additional synonymy: Blastocaulon rupestris (Gardn.) Ruhl, ex 
Uoldenke, Phytologia k'- 338, in syn. 1953. 

Additional bibliography: Moldenke, fjiown Geogr, Distrib, Erioc, 
7, 28, 39, 53, 55. & 59, 19l;6; Uoldenke, Phytologia k'- 338, 1953; 
Uoldenke, R6sum6 87, 237, 292, 328, 33U, 352, & U79. 1959; Uol- 
denke, R^sumfi Suppl, 12: 11, 1965. 

Additional citations: BRAZIL: Minas Gerais: G. Gardner 5272 
(B— isotype, N— isotype, W— 1067056 — isotype); Mexia 5779 (B, Ca- 
509U43, Mi, Ut— 50252a, Vi, W— 157I9OU), 5780 (Gg, Go, Mi, Ut— 
50251a, W— 1571905); E. Pereira 2805 [Pabst 36UI] (Bd— 33U6, Z); 
Schwacke 81i85 (B). MOUNTED ILLUSTRATIONS: drawings fie notes by 
KOmicke (B) . 

BLASTOCAULON SPELEICOLA Alv. Silv. 

Additional bibliography: Alv. Silv., Fl, Mont, 27U, pl. 182, 
1928} Moldenke. Known Geogr. Distrib. Eri^c. 7 & 28. 191*6 j Mol- 
denke, RSsum^ 87 & U79. 1959. 

CARPTOTEPALA Moldenke 

Bibliography: Moldenke, Fieldiana Bot, 28: llli. 1951; Angely, 
Cat, Estat, 10: [2], 1956; Uoldenke, Bull, Jard, Bot. Brux. 27: 
118. 1957; J, A, Steyem., Fieldiana Bot. 28: 1157, 1957; Angely, 
Fl, Paran, 10: 7, 9, & 10, 1957; Uoldenke, R^suafe 70, 7U, 2U9, 
326, UOl, & U79. 1959; F. A. Barkley, List Ord, Faa, Anthoph, 113 
& lii9. 1965; Aiiy Shaw in Willis, Diet, Flow, PI,, ed. 7, 202. 
1966. 

Type: C^. insolita Moldenke ["C, jenmani (Gleason) Uoldenke], 
The geneiric name is derived from the Latin, carptin, and the 
latinized French, tepala, meaning separate divisions of the peri- 
anth, because of the completely free sepals and petals in the 
flowers of both sexes. 



1968 Moldenke, Notes on Erlocaulaceae 375 

CARPTOTEPAIA JENMANI (Gleason) Meldenke 

Synonyaj: Paepalanthus jenmanl Gleason, Bull, Tonrey Bot. Club 
56: lU. 1929. Carptotepala insollta Moldenke, Fieldiana Bot. 28: 
llli — 116. 1951. Paepalanthus chimantensls Moldenke, Bull, Jard, 
Bot, Bnix. 27: 118, in syn. 1957. 

Additional bibliography: Gleason, Bull. Torrey Bot, Club 56: 
Hi. 1929; Moldenke, Fieldiana Bot, 28: llli— 116. 1951; Moldenke, 
Phytologia h'- 338. 1953; Moldenke, Mem. N. Y. Bot. Gard. 9: 278. 
1957; Moldenke, Bull. Jard. Bot, Brux. 27: 118—119. 1957; Mol- 
denke, R^sum6 70, 7U, 2U9, 326, & U79. 1959. 

Collectors describe the roots of this plant as thickish and 
orchid-like, the leaves borne in dense clusters, rigid, erect, 
firmly menibranous or rigid-coriaceous, varying from rich- or 
pale-green to grass-green on both surfaces, the involucre buff, 
the heads white or gray-white with blackish on the outer parts at 
the base, the flowers white or whitish, and the bracts gray-bronn. 
Gleason' s original description was "Leaves densely cespitose, 
soft and lax, 1 — 2 mm. wide, 8 — 12 cm. long, glabrous, subulate- 
tipped; pedxxncles 20 — 25 cm. long, costate, somewhat twisted, 
glabrous; sheaths strongly twisted, U — 5 cm. long, sparsely hir- 
sute; heads hemisphere, 3—5 mm. in diameter; bracts broadly 
ovate to orate-oblong, appressed, imbricate in several series, 
glabrous." 

The type of the species is Jeman 1032 , collected on the 
Kaieteur savanna in British Guiana, and deposited in the herbar- 
ium of the Royal Botanic Gardens at Kew. The conspecific Cj, in- 
solita , on the other hand, was based on £. A, Steyermark 60703, 
collected by a waterfall in a swampy savanna between Rio Karuai 
and Sal to de Itaba-naima along the Rio Kaniai, at the southwes- 
tern base of Ptari-tepui, at 1220 meters altitude, Bolivar, Ven- 
ezuela, on November 28, I9UU, and is deposited in the herbarium 
of the New Tork Botanical Garden. Paepalanthus chimantensis was 
based on Steyermark & V/urdack 365, also from Bolivar, Venezuela. 

The plant has been collected at altitudes of 65 to 2600 meters, 
in anthesis from January to March, and in July, August, October, 
and November, and in fruit in July. Steyermark records the ver- 
nacular name "leut" . He also states that the species forms dense 
mats on wet rocks at the base of waterfalls, that it is common 
along swift water and rapids, locally abundant in large colonies 
in rapid water among rocks, and found in the spray zone on top 
of waterfalls. Maguire found it on moist rocks, while Sandwith 
describes it as tufted in sand among boulders by falls. Maguire 
& Fanshawe found it by waterfalls, on sandstone savannas, and 
locally common by riversides. Steyermark & Wurdack describe it 
as locally frequent on moist mossy ground, in scrub forests, in 
dense cushions in thickets, in dry sand, and in large colonies 
in rapid water among rocks. On the label of their no. 365 they 
note that its "leaves nan*ower and caudex more elongate than 
36U but probably only an ecological variant". I agree with this 
conclusion, Whitton found the species growing on wet rocks, in 
noistish open ?^ite sand, and as recently exposed or still below 



376 PHTTOLOOIA Vol. 17, no. 5 

river irater. He notes "buda farther advanced the further up shore 
one coes". 

Additional citations: VEIffiZUELA: Bolivar: B. llaguire 33$l6a 
(N); J. A. Steyemark 6070 (U), 7U662 (Z), 76016 (Z). 76057 (Z); 
Steyermark & Wurdack 72 (N), 361 (N), 365 (N), U76 (N) . BRITISH 
GUIANA: S. G. Harrison 1391 (K, 3); Jenman 1032 [!.'. Y. Bot. Gard. 
Type Photo neg. 5007] (K — type, II — photo of type, IJ — photo of 
type), 7198 (K), 7U86 (Ut— 9107a) ; Maguire ?,. Fanshavre 32312 (IJ), 
326U3 (Uu, N); SandTfith 1258 (K, Ut— Mi22lia); Schomburgk s.n. 
(K); WMtton 36 (K), 77 (K) , 367 (K) . 

COUANTHERA L, B. Sm. 

Bibliography: L. B. Sa., Contrib. Gray Herb., ser. 2, 117: 33- 
39, pi. 2. 1937; Moldenke, Phytologia U: 338. 1953; Angely, Cat. 
Estat. 10: [2]. 1956; Angely, Fl. Paran. 10: 5, 7, 9, & H. 
1957; Anon., U. S. Dept. Agr. Bot. Sub j . Index 5: 1226. 1958; 
Moldenke, R^suml 70, 75, 88, 351, UOO, & U79. 1959; Moldenke, 
RSaum^ Suppl. 1: 5, 6, 16, 21, 23, & 25. 1959; Angely, Liv. Gen. 
Bot. Bras. 19 & U2. I960; F. A. Barkley, List Ord. Earn. Anthoph. 
113 & 15U. 1965; Airy Shaw in Willis, Diet. Flow. PI., ed. 7, 
268. 1966; Moldenke, Phytologia 13: 218. I966; Moldenke, Biol. 
Abstr. \xl'. 6792. 1966. 

The generic name is derived from the Greek, K<^ p^-i and ocrdo's^ 
meaning "hairy flower", since the anthers are long-haii-y. 

Mrs. A. GOrts-van Rijn, in a letter to me dated March 21, 
1966, casts some doubt on the validity of this genus. She says: 
"We have been looking very thoroughly to some Comanthera kegeli - 
ana specimens, partly annotated by you, and also used the publi- 
cation of L. B. Smith in Gontr. Gray Herb. 117: 38. 1937. He 
gives the description and some illustrations of this new genus 
and of C. linderl. Vfe cannot agree with him on the characteris- 
tics of the flowers. He describes the male flowers as having a 
very reduced perianth and only one stamen. The sterile flowers, 
according to him, have reduced stamens. We have been looking to 
the flowers of Syngonanthus or Gomajithera kegeliana , but could 
not find similar male flowers. We did, however, find overripe 
female flowers, where the fruits had come out and the perianth- 
segments had partly fallen off; these had the appearance of the 
described male flowers of Gcmanthera L. B. Smith. About the 
sterile flowers we are not quite sure, but they are supposed to 
be the immature male ones. They do have staaens, but it is 
difficult to say whether they are reduced or only verj'- young." 
In a letter to me dated August 3, 1967, Dr. Smith replies as 
follows: "I have just gotten around to studying my Comanthera 
that I borrowed from Harvard at your suggestion. It has stam- 
inate flowers as I described them. Yo\ir Syngonanthus akurimen - 
sis is the sane thing as regards the type but the Irwin col- 
lection shows no such stamens . Maybe the species is polymor- 
phic and some heads lack functional stamens." 

Type: Comanthera linderi L. B. Sm. [•'C. kegeliana (KOm.) 



1968 Uoldenke, Notes on Eriocaulaceae 377 

Moldenke . 

COMA.OTEERA KEGELIANA (KOm.) Moldenke 

Synonymy: Paepalanthus kegelianus KOrn. in Uart,, Fl. Eras. 3 
(1): U38. 1863. Dupatya kegeliana (KOrn.) Kiintze, Rev, Gen. PI. 
2: 7li5« 1891. Syngonanthus kegelianus (KWrn.) Ruhl, in Engl,, 
Pflanzenreich 13 (IV, 30): 273. 1903. Ccmanthera linderi L. B. 
Sm., Contrib. Gray Kerb., ser. 2, 117: 38—39, pi. 2. 1937. 
Syngonanthus sJcurimensis Moldenke, Phytologia 2: 371 — 372. 19U7. 
Syngonanthus akurimensis var. amazonicus Moldenke, Phytologia 3? 
U2. 19U8. 

Additional bibliography: KOm. in Mart., Fl. Bras. 3 (1): U38. 
1863; Kuntze, Rev, Gen, PI, 2: lh$» 1891} Ruhl. in Engl,, Pflan- 
zenreich 13 (IV, 30): 273. 1903; Moldenke, Bull. Jard. Bot, Brux. 
27: 119—120. 1957; Angely, Fl. Paran. 10: 5. 19^7; Anon., U, S. 
Dept. Agr. Bot, Sub j . Index $: 1^226, 19^8; Moldenke, R^sumS 70, 
75, 77, 88, 280, 326, 351, & U79. 1959; Moldenke, Rfesuml Suppl. 
1: 5, 6, 16, 21, 23, Sr 25. 1959; Moldenke, Phytologia 13: 218. 
1966; Moldenke, Biol. Abstr. hi: 6792. 1966, 

The species has been encountered by Lindeman on a large sand 
savanna. It has been collected in anthesis from llarch to July 
and in fruit in May. It is described by Tamayo as growing 5 — 8 
cm. tall. An isotype, Kegel llt73 , ""f^s photographed by Mac bride 
in the herbarium of the Conservatoire et Jardin Botaniques at 
Geneva and is his type photograph number 25170. Material has 
been misidentif ied and distributed in herbaria as " Compositae " . 

Additional & emended citations: VENEZUELA: Bolivar: Las ser 
1705 (K, N, N, Ve, W— 1901897); Tamayo 323U (F— photo, N, N— 
photo, Ve, W, Z — photo). Federal District: Lockhart s.n. [Ga- 
racasj (K; . BRITISH GUIANA: Cox & Hubbard 121 (N); Irwin BG, 
20 (W— 21lalilli, Z); Linder UO [N, Y, Bot, Gard. Type Photo neg. 
5006] (G, N— photo, N— photo); Martyn lli6 (K) . SURINAM: Kegel 
lli73 [Macbride photos 25170] (N— photo of isotj-pe, V^— photo of 
isotype); Lanjouw & Lindeman 298U (K, Ut — 178768); Lindeman 
UOI8 (Ac). BRAZIL: Amazonas: Fr6es 22U33 (Ca— 28252, N) . Par^: 
Ducke 5,n. [Herb, Mus, Goeldi 12088] (Bs), 

ERIOCAULON Gron. 

Additional & emended synonymy: Erioucaulon L., Mant. 580, 
sphalm, 1767. Cespa Hill, Herb. Brit. 1: pi. 66 [some copies]. 
1769. Nasmythla Huds., Fl. Angl., ed. 2, 2: UHi. 1778. 
E^icaulon Lour., Fl. Cochinch. 1: 60, sphalm. 1790. Eriocaulon 
L. ex Steud., Nom. Bot., ed. 1, 312. 1821. Randalia P. Beauv. 
ex Desv., Ann. Sci. Nat. Paris 13: U7. 1828. Sphaerochloa P. 
Beauv. ex Desv., Ann. Sci. Nat. Paris, ser. 1, 13: 1;7. 1828. 
Sphoerochloa P. Beauv. ex Desv., Ann. Sci. Nat. Paris, ser. 1, 
13: pi. 5, fig. 1. 1828. Symphachne P. Beauv. ex Desv., Ann. 
Sci. Nat. Paris, ser. 1, 13: U7. 1328. Leucocephala RoA., Fl. 
Ind. 3: 612. 1832, Busseuillia Lesson in Bougainville, Journ. 



378 P H y T L Q I A Vol. 17, no. 5 

Navig. Aut. Freg. Thetis & Corv, Esp^r. 2: 3U8. 1837. Syapachne 
P. Beauv. ex Steud., Noo. Bot., ed, 2, 2: 65Ii. 18U1. Chaetodlacua 
Steud., Syn. Pi, Cyp. 2: 261. 18$$. Electroaperma F. liuell,, 
Trana. Philos. Soc. Victoria 1: 23. l8$$. Dichrolepia Welw., 
Apont. Phyt.-geogr. $U2. 18 $9. Lasiolepis Boock. (in part), flo- 
ra $6: 90. 1873 [not Laaiolepie Bennett, I838]. Syaphyachna Post 
& Kuntze, Lexicon $Uh» 190U. Eric anion lierr. ?c VTalker, Bibl. 
East. A3iat. Bot. 3U3, sphalm. 1938. Laaiolepsis BtJck. apud 
Milne-Redhead, Kew Bull. Misc. Inf. 19U3: 172, sphaLn. I9U8 . 
Randalia Petit apud Moldenke in Humbert, Fl. Madag. 36: 2, in 
sjm. sphalm. 19$$. Randalia Tetiv. ex Desv." ex Angely, Cat, 
Estat. 10: [2], in syn. sphalm, 19$6. Sphaerochloa "P. Eeauv. 
ex Desv." apud Angely, Cat, Estat. 10: [2], in syn. 19$6. 
Symphachne "P. Beauv. ex Desv," apud Angely, Cat. Estat. 10: [2], 
in syn. 19$6. EIriaucolon L. ex Moldenke, Rifisum^ 28$, in syn. 
19$9. Randalia Beauv. & Desv. ex Moldenke, R6sum5 3U2, in syn, 
19$9. Randalia Petiv. ex Moldenke, R^sum^ 3ii2, in syn. 19$9. 
Sphaerochloa Beauv. & Desv. ex Moldenke, R^sumi 3ii$, in syn. 
19$9" Eriocaullon With, ex Moldenke, Rigsvm6 Suppl. 3: 31, in 
syn. 1962. Eriocaulum KOm. apud Angely, Eibl. Veg. Paran. 1$$, 
sphalm. I96U. Randalia "Beauv. ex Desv," apud Airy Shaw in 
Willis, Diet. Flow. PI., ed. 7, 9$0, in syn. I966. Sympachne 
Steud. apud Airy Shaw in Willis, Diet. Flow, PI., ed. 7, IO9I, 
in syn. I966. 

Additional &; emended bibliography: Petiv., Gaz. pi. 6, fig. 2. 
1702i L., Sp, Pi,, ©d. 1, 87 & 129. 17$3i Crantz, Inst. 1: 36O, 
1766; Pluk., Aim, pi. U09, fig. $. 1769; Hope, Phil. Trans. Roy. 
Soc. $9: 2ia-— 2li$, pi. 12. 1770} Scop., Introd. Hist. I^at, 20U. 
1777; Kuds., Fl. Angl., ed. 2, 2: UHi. 1778; Walt., Fl, Carol, 
83. 1788; Lam., Encycl. 3: 276, 1789; Lour., Fl. Cochinch. 1: 60. 
I79O; L. C. Rich., Act. Soc. Hist. Nat. Paris 1: 113. 1792; 
Willd. in L., Sp. PI., ed. k, 1: li86. 1797; Michx., Fl. Bor.-am. 
2: 16$. 1803; Pursh, Fl. Am. Sept. 1: 91. 1811;; Roxb,, Hort. 
Beng. 68. 1811;; Roem. & Schult. in L,, Syst, Veg,, ed, 1$ nova, 
2: 86U. 1817; Nutt., Gen. 1: 90. I8I8; Ell., Sketch Bot. 2: $6$. 
I82it; Lodd., Bot. Cab. Hi', pi. 1310. 1828; Bong., M6n. Acad. Sci. 
St. P^tersb., ser. 6, Sci. Math. Phys. & Nat. 1: 601— 6$6, pi. 
1—10. 1831; Wall., Numer. List 207—208 ["207"]. 1832; Hook, in 
Curtis, Bot. Mag. $9: pi. 3126. 1832; Wall., Plant. As. Rar. 3: 
28. 1832; Beck, Bot. 370. 1833; Bong., Mfin. Acad. Sci. St. Pfet- 
ersb. ser. 6, Sci. Math. Phys. & Nat. 2: 219—238, pi. 11—19 
(1833;, ser. 6, Sci. Nat. 1: $U$— $60 (183$), and ser. 3, Bot, 
9—29, pl. 20—2$, I81i0; Steud., Nbm. Bot., ed. 2, 2: 6$U. 1811; 
Griff., Itin. Notes [Posthum. Papers 2:] 6$. 18U8; Benth. in 
Hook., Niger Fl. $U7. 18U9; Steud., Syn. PI. Cyp. 2: 261 & 268— 
283. 18$$; F. Muell., Trans. Philos. Soc. Victoria 1: 21;. 18$$; 
Benth., Fl. Hongkong 382. I86I; Benth., Fl. Austral. 7: 192. 
1878; F. Uuell., Syst. Census Austral. PI. 123. l882j F. M, Bai- 
ley, Syn, Queensl, Fl. $78. 1883} A. W, Ghapm., Fl. South. U. S., 



1968 Moldenke, Notes on Eriocaulaceae 379 

ed. 2, 502— 50U, 658, & 696. 1889} F. Muell., Proc. Unn. Soc. N. 
S. Wales 5: 250. I89OJ F. Muell., Bot. Centralbl. Uh: 302. I89O} 
Morong, Bull. Torrey Bot. Club 18; 35U. I89I; Maxia., Diagn. PI. 
Nov. As, 8: 25. 1892; Jacks, in Hook, f , & Jacks., Ind, Kew., pr. 
1, 1: 877—880. 1893} Moore & Betche, Handb. Fl. N. S. Wales UiO. 
1893} J. G. Baker, Journ. Linn. Soc. Lond. Bot. 20: 227. 1893} 
Hook, f., Fl. Brit. Ind. 6: 571—585. 1893} Coult., Contrib. U. 
S. Nat. Herb. 2: h^9* I89U} Jacks, in Hook. f. & Jacks., Ind. 
Keir., pr. 1, 2: 681. 1895} Britton & Br., HI. Fl., ed. 1, 1: 
371«_373^ 602, & 611. fig. 899—901. I896} Ruhl. in Engl., Bot. 
Jahrb. 27: 65—85. 1899} Tate, Trans. Roy. Soc. S. Austral. 23: 
291. 1899} H. T. Holm, Bot. Gaz. 31: 17—37. 1901} N. E. Br. in 
This elt .-Dyer, Fl. Trop. Afr. 8: 255. 1901} F. M. Bailey, Queen- 
si. Fl. 6: 1715. 1902} B. L. Robinson, Rhodora 5: 175—176. 1903} 
J. K. Small, Fl. Southeast. U. S., ed. 1, 236. 1903} Ruhl. in 
Engl. Pflanzenreich 13 (IV, 30): 1—108. 1903} Post & Kuntze, Lex- 
icon $Uh. I90U} Ruhl. in Urb., Engl. Bot. Jahrb. 37: 519—520. 
1906} R. M. Harper, Ann. N. I. Acad. Sci. 17: 267, pl. 2U, fig. 
1. 1906} C. H. Wright, Keir Bull. Misc. Inf. 1907: 3— U. 1907} 
Robins. & Fern, in A. Gray, New Man. Bot., ed. 7, 261 & 898. 
1908} M. A. Day, Check List 39. 1908 j Nakai, Bot. Mag. Tokyo 2U: 
5—6. 1910} Kawakami, List PI. Fomos. 130. 1910} G, T. Stevens, 
111. Guide Flow. PI. pl. 9, fig. 5. 1910} R. W. Sm., Bot. Gaz. 
U9: 281—289, pl. 19 & 20. 1910} A. Chev., Sudania 1: 7. 1911} 
W. H. Br., Contrib. U. S. Nat. Herb. 13: 323. 1911} Nakai, Bot. 
Mag. Tokyo 26: [93— 9li] . 1912} Ann. Rep. N. J. State Mus. I9IO: 
pl. 28, fig. 2. 1912} F, M. Bailey, Compreh. Cat. Queensl. Pl. 
58U. 1913} J. K. Small, Fl. Southeast. U. S.. ed. 2, 236. 1913} 
Britton & Br., Illustr. Fl., ed. 2, 1: U53— U55 & [678]. 1913} 
Domin, Bibl. Bot. 20: 506. 1915} Maiden & Betche, Census N. S. 
Wales Pl. 38. 1916} Fern., Rhodora 23: 92. 1921} Alv. Silv., 
Archiv. Mus. Nac. Rio Jan. 23: l62, pl. U. 1921} Fyson, Journ. 
Indian Bot. 2: 133—150, 192—207, 259—266, & 307—320, pl. 1— 
IiO (1921) and 3: 12—18 & 91—115, pl. 11—32. 1922} Anon., Kew 
Bull. Misc. Inf. 1923: 303. 1923} Ltftzelburg, Estud. Bot. Nord- 
6ste 3: U47 & 150. 1923} Fyson, Indian Sp. Erioc. 1—88, pl. 1— 
51. 1923} Alv. Silv., Fl. Mont. 17—19, pl. 5 & 5a. 1928} Uphof 
in Karst. & Schenck, Vegetationsbild. 21 (1-2): n.p. 1930} Ruhl., 
Notizbl. Bot. Gart. Berlin 10: lOUO— lOUi. 1930} N. E. Br., Kew 
Bun. Misc. Inf. 1931: 61. 1931} Ewart, Fl. Vict. 263. 1931} Sol- 
OBon, Joum. Indian Bot. Soc. 10: 139 — liiii. 1931} R. M. Adam, New 
Fl. & Silv. 6: 60 — 63, pl. 2li & 25. 1933} Tu, Chinese Bot. Diet., 
abrdg. ed., 1317. 1933} J. K. Small, Man. Southeast. Fl. 258. 
1933} Tang, Contrib. Inst. Bot. Nat. Acad. Peiping 2: 133. 193hi 
H. B. Davis, Life & Works Pringle U3, $^, 56, 9li, 105, 123, lia, 
219, & 655. 1936} Van Steenis, Trop. Natuur 25: 2. 1936} Moldenke, 
N. Am. Fl. 19: 17—37, UO, li3, hh, li6, & 50. 1937} Cory, Texas 
Agr. Exp. Sta. Bull. 550: 29. 1937} Merr. & Walker, Bibl. East. 
Asiat. Bot. 3U3. 1938} Satake, Joum. Jap. Bot. 15: ll;0— lli5 & 
627—632. 1939} Wells, Bot. Rev. 8: 537. 19U2} Moldenke in Lun- 
dell, Fl. Texas 3 (1): U— 5. 19^2} Carolin. Florist Gov. J. Dray- 
ton S. C. Ha. 19ii3} Black, Fl. S. Austral., ed. 2, 1: 179. 19U3} 



380 P H T T L I A Vol. 17, no. $ 

Rouleau, Contrib. Inst, Bot. Univ. L'ontreal SU: 161 h 313 . I9U1; 
Eyries h Robertson, U. S. Pub. Health Bull. 286: 1&6. 19Ui; W, A. 
Uurrlll, Guide Fla. PI. 3U. 19U5; Castellanos in Descole, Gen. Sp. 
PI. Argent. Eriocaulac. 87, pi. 17. l^U^i Abbiatti, Revist . Lus. 
La Plata Bot. 6 (26): 329—330 pi. 2 (1), fig. U (d) b^ 6. 19li6; 
Razi, Joum, L^sore Univ. 7 (h;: 77. 19U6; Fern., Rhodora UC: iv 
& 58. 19U6j L^oldenke, ijiov/n Geogr. Distrib. Erioc. [1] — 8, 19 — 28, 
30, 32— 142, UI4, U7, 53, 56, t 60—62. I9U6; it. R. Tatnall, Fl. 
Del. 75. I9U6; Jacks, in Hook. f. h Jacks., Ind. Kew. pr. 2, 1: 
877—880. I9U6; Abbiatti, 3oc. Argent. Bot. Bui. 1: 230—281. 
I9I46; Castellanos, Lilloa 20: 2iiii. 1919; Raizada, Sci, >j. Cult. 
lU: 387—388. 19U9; Faegri & Iversen, Text-book Mod. Pollen An- 
aTys. 193 &i 221. 1950} Hare, Linn. Soc . Lonri. Joum. Bot. 53: 
U22— UliS. 1950; Herter, Rev. Sudam. Bot. 8: 163— I6U. 1950; Mae- 
koira, Journ. Jap. Bot, 26: 116. 195l» Penfound, Bot, Rev, I8: 
li31. 1952; Zinderenbakker, S. Agr, Pollen 1: 32, 36, & 79, pl. 7, 
fig, 33 «: Ui. 1953; Moldenke, Biol, Abstr. 27: 98U, 2026 £c 3121, 
1953; Moacyr Lisboa, Cent, Nasclm. Leon. Bot. Damazio [2], 195ii; 
Koyama, Philip, Journ. Sci, Bot. 8U: 367—368 & 378, pl. 6, 1955; 
Razi, Joum, Itysore Univ, B.IU (10): U60. 1955; Razi, Contrib, 
Bot, UO: 92, 1955; Razi, Proc, Nat, Inst. Sci, India 21B (2): 82, 
1955; Anon,, Assoc, Etud. Tax. Fl. Afr. Trop. Index 1955: 29 — 30. 
1956; Koyama, Joum. Jap. Bot. 31: 9 — 11, fig. 3. 1956; Maaa Iku- 
si, Pollen Or. Jap. 1956; Angely, Cat, Estat. 10: [2]. 1956; K. 
Hess, Bericht. Schweitz. "'ot, Gesell. 67: 33. 1957; E. H, Yfalker, 
Proc. 8th Pacif, Sci, Cong. U: U06. 1957; Angely, Fl. Paran. 10: 
\^~3 & 11. 1957; Anon., Biol, Abstr. 29: 32U8 Sc 3626 (1957) and 
30: 3931 & U393. 1958; Anon., U, S, Dept, Agr. Bot, SubJ. Index 
5: U226 — U277. 1958; Kostemans, Proc, Sympos. Humid Trop. Veg. 
159. 1958; Suvatabandhu, Proc. Sympos. Hxuiid Trop. Veg. 173. 
1958; Alain, Revist. Soc. Cub. Bot. 15: 1^9. 1958; DeRoon, Inter- 
nat. Direct, Spec. 201, 1958; P, van Royen, Nov. Guin., new ser., 
10: 21— lili, fig, 1—5. 1959; Soukup, Biota 5: 300—301. 1959; 
Anon., Assoc. Stud, Tax. Fl. Afr. Trop, Index 1958: 31. 1959; 
Razi, Rec, Bot, Surv, India 18: 19, 1959; Anon., Kew Bull. Gen, 
Index 1929-1956, 111, 1959; Reitz, Sellowia 11: 103 . 1959: liol- 
denke, R6sura4 U— 12, H, 22, 23, 25, 27, 32, 35, 36, U, U3, U6, 
U8, 51—53, 63, 66, 70, 71, 75, 77—79, 83, 88, 89, 112, II3, 
116, 119, 123, 132—138, lUo, lUi— 151, 153, 156—163, 165—167, 
169—176, 178—181, 131;, 186, 188, 190—193, 196, 201, 20U, 205, 
207—209, 211, 213, 226, 2U0, 277, 278, 231, 28U— 291, 309, 320, 
323, 32li, 326, 328, 329, 342, 3ii5, 350, 351, 395—399, U-li, hlS, 
i;17— UI9, U2U, U26, U28, U79— U8U, k h9k. 1959; Moldenke, Rgsuafi 
Suppl. 1: [l]--3, 5—19, 21, 23, Sc 25. 1959; P. van Royen, Blumea 
10: 126—135, fig. 1. i960; Straka, Erdkxmde lU: 60 £c 87. i960; 
Angely, Fl. Paran. 15: lU. I96O; Renn6, Levant, 'lerb, Inst. Agron. 
68. I96O; Jacks, in Hook, f, & Jacks., Ind, Kew,, pr, 3, 1: 877 — 
830, I960; Angely, Liv. Gen, Bot, Bras. 19 & UU. I960; Moldenke, 
Biol. Abstr. 35: 1688 & 2177. I960; Santapau, Fl. Bombay & Sal- 
sette [3]. I96O; Moldenke, RfisumS Suppl. 2: [1], 2, U— 7, Sc 9. 
i960; Panigrahi Sc Naik, Bull. Bot, Surv, India 3: 383. 1961; Run- 
ner, Rep. G. W. Groff Coll. 292. I96I; Fables, Bartonia 32: 9. 



1968 Moldenke, Notes on Sriocaulaceae 381 

1961 i Angely, Fl. Paran. 1?: 2U. 1961; Van bteenis-Iiruseman, Fl. 
Males. Btai. 3: xli, 76I, 861, & 862. 1962; J, H. V/illis, Handb. 
PI. Vict. 281. 1962; Hocking, Excerpt. 3ot. A.Ii: $92 & 593. 1962; 
K. Larsen, Nat. Hist. Bull. Siam Soc. 20: 113. 1962; Moldenke, 
R6sum5 Suppl. 3: [1]— 5, 7, 9, 12, 15— 2U, 26, 28, 31, & 32 (1962), 
h: [l]—7 & 11 (1962), and 5: [1], 2, 5, & 6. 1962; Angely, Fl. 
Bacia Paran. 22: 31. 1962; Hatusima, iuea. South. Indust, Sci. 
Inst. Kagoshima Univ. 3 (1): 123 (1962) and 3 (2): 123 & 131. 
1962; G. L. Shah, Bull. Bot. Surv. India k'- 237. 1962; Prain, Ben- 
gal PI., ed. 2, 2: 8U7--3U8. I963; Arker, Water PI., ed. 2, 286. 
1963; J. Joseph, Bull. Bot. Surv. India $: 283 & 297. 1963; Heg- 
nauer, Chemotax Pfl. 2: 152. 1963; Montgomery & Fairbrothers, 
Bull. Torrey Bot. Club 90: 92 & 96. 1963; Gleason & Cronquist, 
Man. Vase. PI. 183 — 13U. 1963; Espirito Santo, Junt. Invest. Ul- 
tramar Est. Ens. Sc Docum. lOli: 5U & 88. 1963; H. P. Riley, Fam. 
Flow, PI. S. Afr. 199. 1963; Moldenke, RlsunS Suppl. 6: [1], 2, 
5, 6, 8, & 9 (1963), 7: 3 & 6 (I963), 8: 2 & 3 (196U), 10: U & 5 
(I96U), and 11: [1] & U-— 6. I96U; Rao & Sastry, Bvill. Bot. Surv. 
India 6: 281 & 28U. 196U; P^mt, Reg. Veg. 9. 196U; Langman, Sel- 
ect. Gviide Lit. Flow. PI. Mex. 911. I96U; Panigrahi, Chowdhury, 
Raju, & Deka, Bull. Bot. Surv. India 6: 260 — 261. 196ii; Bhatta- 
charyya. Bull. Bot. Surv. India 6: 208. I96U; C. U. & D. S. Patel, 
Vidya 7: [58]~70. 1961;; Moldenke, Biol. Abstr. US'- 5019. 196U; 
Batson, Wild Fls. S. C. 28. I96U; Koyama in Kitamura. Murata, & 
Koyama, Col. Illustr. Herb. PI. Japan 175—185, pl. 48. I96U; 
Angely, Bibl. Veg. Paran. 155 ^ 253. 196U: D. Walker, Govt. Sara- 
Trak Sympos. Ecol. Res. Humid Trop. Veg. lUl. 1965; F. A. Darkley, 
List Ord. Fan. Anthoph. 113 & 16U. 1965; Thanikaimoni, Pollen £c 
Spores 7: I8I— 189. 1965; Thanikaimoni, M^m, Mus. Nat. Hist. Nat. 
Paris, new ser. B, Hi: 9—38. 1965; J. S. Beard, Descrip. Gat. W. 
Austral. PI. 9. 1965; Hedberg, Webbia 19: 526. 1965; Humbert, 
Trav. Sect. Scient. &: Techn. Inst. Frang. Pond., ser. 6, Not, 
Carte Madag. 66. 1965; Stocking, Nat, Conserv. Ecolog. Stud. 
Leafl. 6: [15]. 1965; F. R. Fosberg, Govt. Sarawak Sympos. Ecol, 
Res. Humid Trop. Veg. 286. 1965; Moldenke, R6sum§ Suppl. 12: [1]- 
5 & 7—10 (1965), 13: [1], 3, 5, & 7 (1966), and lii: [1]~3 & 8. 
1966; Thanikaimoni, Biol, Abstr, kli UI69 * I966; S, V, Ramaswami, 
Study Flow, PI. Bangalore [thesis] 219—221 & lii06— 1U07. 1966; 
J. A. Steyerm,, Act. Bot. Venez. 1: 15 & 19. I966; Goodland, Bol, 
Soc. Venez. Cienc. Nat. 26: 3I45. 1966; Klots, New Field Book 
Freshw. Life 9U. 1966; B. G. Briggs, Contrib. N. S. Wales Nat. 
Herb. U: 2U & 26. I966; Shinners, Sida 2: Uil. 1966; R. C, Jacks., 
Reg, Veg, U3: 33. 1966; Anon., Gen. Costa Ric. Phan. 2. 1966; R. 
H. Coopton, Joum. S. Afr. Bot. Suppl. 6: 19, 33, & 92, I966; 
Krai, Sida 2: 290—312 & 330. I966; 0. D. Evans, Contrib. N. S. 
Wales Nat. Herb. Fl. Ser. 27/28: 9—12. I966; Aiiy Shaw in Willis, 
Diet. Flow. PI., ed. 7, I68, 223, 22U, 3U9, 396, Ul7, Ul8, 620, 
6U7, 758, 950, 1057, 1091, & 1092. 1966; Subramanyaa & Henry, 
Bull. Bot. Surv. India 8: 211;. I966; Sebastine & Ramanurthy, Bull. 
Bot. Surv. India 8: 182. I966; J. L. Ellis, Bull. Bot. Surv. India 
8: 329 & 339. 1966; Sculthorpe, Biol. Aquat. Vase. PI. 23, 389— 
391, 393, & 39U. 1967; L. V. Barton, Bibl. Seeds 782. 1967; Satake, 



382 P H Y T L I i Vol. 17, no. 5 

Nat. Sci. & Mu8. 3U: 161 & 162. 1967} Fulling, Ind. Lot. Record. 
Dot. Revierw 178. 1967; L. 0. Williama, Fieldiana hot. 31» 2U9— 
269. 1967; R. M. Harper, Castanea 32: 17. 1967; 0. D, Evana, idol, 
Abstr. U8: U562 . 1967; Rickett, Wild Fls. U. S. 2 (1): 135 (1967) 
and 2 (2): 659. 1967; Friedrich-Iiolzhaamer in lierxMtCLler, Prodr. 
Fl. Stfdw. Afr. 159 1 1—2. 1967; berhaut, Fl. S6n6gal, ed. 2, 311. 
1967; Anon., Biol, Abstr. U8 (10): S.60. 1967; J. «r. A. Raynal, 
Adansonia 7: 329. 1967; L. S. Thoaas, Pine Barrens 23. 1967; D. A. 
Livingstone, Ecolog. Monog. 37 (1): U3. 1967; Uoldenke, RSsumfi 
Suppl. 15: [1], 8, 10, 12, lU, & 20 (1967) and 16: [1], 2, 5, 7— 
9, 12, 19, 21, & 25—27. 1968; Justice & Bell, Wild Fls. N. C. 13 
& 209. 1968; Keikle, Kew Bull. 22: lUl— Hiii. 1968; R. M. White, 
Irish Naturl. Joum. 16: UO. 1968. 

The scientific name of this genus is taken frcm the Greek, 
£- p t o v^ and Kccv/Koi, meaning "hairy stem", since many sjsecies 
have pubescent scapes or peduncles. Eerhaut (1967) describes 
this genus, as knoim to him, as "Bract^es triangulares, sonnet 
beaucoup plus large, base cunSifonae, 6a bractSes de base seule- 
ment, dont 3 exterieures d^ojrdant la base du capitule.. ..capi- 
tules blanc-neigeux" . Riley (1963) reports the sporophytic 
chromosome number as 32 and 36. Thanikaimoni (1965) studied the 
pollen of ii6 species of the genus. Livingstone (1967) tells ua 
that the genus is among the minor taxa in the ericaceous belt of 
the Ruwenzori Motintains in equatorial Africa. Rickett (1967) re- 
cords the common names "hatpins" and "pipeworts" for the genus 
as a whole, and Espirito Santo (I963) records "or8" . 

The Laslolepis of Bennett, referred to in the synonymy above, 
is a synonym of Harrisonia R. Br, in the Rutaceae . The type 
species of Eriocaulon is E^ decangtilare L. [as established by 
Britton & Brown (1913)]} that of Chaetodlscus is £_. gllberti 
Steud., based on Gilbert 153 from Australia [Ruhland reduces 
this genus to synonjnay under Eriocaulon , but fails to dispose of 
the type binomial anywhere in his work] . The type of Electro - 
sperma is E, australasicua F, Muell, [" Eriocaulon australasicum 
(F, Muell,) KOm.]. Lasiolepis has no type indicated; three 
species were proposed in the original publication: L. aquatica 
Boeck,, L^ brevifolla Boeck., and L_. pilosa Boeck. — of these 
the first two are members of the genus Eriocaulon , while the 
last-mentioned belongs in the genus Paepalanthus . The type spe- 
cies of Nasmythia is N, articulata Huds. [" Eriocaulon septangu - 
lare With,], that of Randalia is R. d ecangulare (L,) P, Beauv. 
[" Eriocaulon decangulare L.], and that of Symphachne is S_. xyr- 
oides P. Beauv, [ " Eriocaulon decangulaz^ L.]. 

The Poilane 138U9, distributed to herbaria as a species of 
Eriocaulon , is actually Fimbristylis tetragona R, Br. in the Gy- 
peraceae . 

ERIOCADLON ABYSSDUCUM Hochst. 

Additional bibliography: Moldenke, Bull, Jard, Bot, Brux, 27: 



1968 koldenke. Notes on Eriocaulaceae 383 

122. 1957; Moldenke, RSsub* 135. 138, lii7, 153, & U79. 1959; Kil- 
lick, Bot. Surv. S. Afr. Men. 3U: 119. 1963; R. H. Compton, 
Journ. S, Afr. Bot. Suppl. 6: 33. 1966; Moldenke, R5sTam6 Suppl. 
16: 8. 1968. 

Compton (1966) records this species from Swaziland. The H. 
Wild 1162 [Govt. Herb. l5lOO] , distributed as £. abyssinictun , is 
actually E. ambo^nse Schinz. 

Additional citations: ETHIOPIA: Schlmper 61i8 (S), 19hh (B— 
isotype, Z — isotype) . 

ERIOCAULON ACHITON KOrn. 

Synonymy: Eriocaulon heteropeplon KWm. ex Moldenke, R6sum6 
Suppl. 1: 17, in syn. 1959 [not E. heteropeplon Alv. Silv., 1928] . 
Eriocaulon schlagintffeitii Rvihl. ex Moldenke, R^sum^ Suppl. 1: 18, 
in syn. 19^9^ Eriocaulon thomsoni Kffm. ex Moldenke, R4sum5 
Suppl. 1: 18, in syn. 1959. 

Additional bibliography: Moldenke, Bull. Jard. Bot. Brux. 27: 
122. 1957; Moldenke, R6sum6 l59, 161, 175, 178, & U79. 1959; Mol- 
denke, R63um6 Suppl. 1: 11, 17, & 18 (1959), 3: 16 (1962), and 
15: 8. 1967. 

This species has been collected on wet cliffs in open areas, 
at altitudes of 50 — 2000 meters, flowering from Januai^- to March 
and in October, and fruiting in February. Hansen & Smitinand de- 
scribe it as "conmon in wet locsilities" in Thailand, and tell us 
that the flowers are •'whitish'* or "dirty-white". Smitinand says 
of it "ccannon in sandy soil along edge of water hole". 

Th© riame, E. heteropeplon Ktfm., appears to be based on 
Schlagintweit 2653 , fi'om East Punjab, deposited in the herbarium 
of the Botanisches Museum at Berlin, and II3II , from Sind, d epos- 
ited in the herbarium of the Naturhistoriska Riksmiseum at Stock- 
hola, while E, schlagintweltii Ruhl. is based on Schlagintweit 
188, from Khasia, deposited at Berlin, and E. thomsoni KOrn. is 
based on £, ^ Hooker 3, from Sikkim, also deposited at Berlin. 
Ruhland also annotated"*the Hooker collection at Berlin as " Erio- 
caulon n. sp." 

Hansen & Smitinand 12388a is a mixture with E£ sexangulare L., 
while Ritchie~12U2 is a mixture with E, stellulatum K»m. and E. 
thwaitesil KWm. 

Material has been misidentified and distributed in herbaria as 
E. sexangulare L. The Smitinand 1982a , distributed as E. achiton , 
is actually E. alpestre Hook, f , 5c Thorns. 

Additional citations: PAKISTAN: East Bengal: Griffith 5576 (B, 
C, S). Sind: Schwagintweit II3II (S). INDU: Assam: Chand 2978 
(Mi); Koelz 31319a (Mi) . East Punjab: Sctilagintweit 13? (B), 
2653 (B) . Kerala: Stocks , Law , &c . s.n. [Malabar, Concan, &c] 
(B). Khasi States: Griffith U7 (B— type); Schlagintweit 188 (B) . 
Madras: Perrottet 1170 (Y, V— 96838, V— 270556). Mysore: S. N;. 
Raaaswamy 20 (Ac), 21 (Rf), 29 (Ac). Sikkim: J. D. Hooker 3 (B) . 



38U P H Y T L G I A Vol. 17, no, 5 

Stato undetermined: Ritchie I2I42 , in part (T). TIIAILAJIL: liar.sen 
& Smitinand 11097 (Cp, Rf ) , l?38Ba (Cp), 12389 (Cp, Rf ) j Snitl - 
nand ^60?~ (Gcy« 

ERI0CAU1/)N ADAin^SII Kclkle 

Additional bibliography: lloldenke, Phytologia 3'- l&l. 19U9; 
lloldenke, R6sum6 I36 L U79. 1959', Uoldenke, R6siim6 Suppl. 1: 8 «c 
9 {19S9) and h: 6. 1962. 

This species has been collected in floner in December, growing 
in poor sandy soil at the uppermost ends of tidal creeks, and also 
"common in wet ditches, often submerged", tleikle (I9US) cocments 
that E. adamesii "is a very distinct Kriocaulon , having closer af- 
finities with the West Indian E^ echinospermum C. Wright, and ita 
allies, than with any African representatives of the genus. E. 
mutatum is the only African species with which it could possibly 
be confused, but this has blackish capitula, and the sepals of the 
? flowers have broad wing-like keels," 

Additional citations: SEl^AL: £. G^ Adam 18299 (Z), 13377 (Z) . 
REPUBLIC OF GUINEA: Boismare Ul7 [Herb. Chillou 3937] (An); Chillou 
17li6 (An), LIBERIA: Dinklage 3009 (B) . 

ERIOGAULON AEQUI^DCTIALE Ruhl. 

Additional bibliography: Moldenke, Known Goegr. Distrib, Erioc . 
5 & 32. 19l;6; Moldenke, Phytologia 3' I8I. 19^9 J Moldenke, RlsuaiS 
70 & li79. 1959. 

ERIOGAULON AFRICANUM Hochst. 

Additional bibliography: J. Hutchinson, Botanist in South. Afr. 
678, 19U6j Moldenke Bull. Jard, Bot. Brux, 27: 122—123. 1957; 
Moldenke, R6sum6 lliQ, l5l, £c 1^3. 1959 j Moldenke, R6suin6 Suppl, 
2: 9 (i960) and 3: 16. 1962. 

This species has been collected at 6OOO feet altitude. liutch- 
inson (19li6) cites liis no. h32h» The Zeyher 1730 , distributed 
as E, africanum , is actually Syngonanthus "wahlbergii (Jikstr.) 
Ruhl. 

Additional citations: SOUTH AFRICA: Transvaal: F\. A. Rogers s. 
n, [Moss k Rogers 1921] (S) . 

ERIOGAULON AFZELIANUM Wikstr. 

Synonymy: Eriocaulon kourous sense Lecomte ex Moldenke, Pw^sumS 
289, in syn. 1959. Eriocaulon afzelil Wikstr. ax Moldenke, R6su- 
m4 Suppl, 1: 16, in syn, 1959. 

Additional & emended bibliography: Moldenke, Known Geogr, Dis- 
trib. Erioc. 20, 21, 32, & 36. 19U6; Moldenke, Bull. Jard. Bot. 
Brux. 27: 123. 1957; Moldenke, R^sumS 13U--138, 289, & U79. 1959} 
Moldenke, R6sum6 Suppl. 1: 16. 1959; Hepper, Bull. Inst. Fond, 
Afr. Noire 27: U20. 1965; Berhaut, Fl. S^n^gal, ed. 2, 311. I967. 

The nane, E, kouroussense Lecomte, appears to be based on Ray - 
nal & Raynal 6795 in the herbarium of the California Academy of 
Sciences at San Francisco. Hepper (1965) found the species grow- 



a 



A: 



>. 



Id 



^), 



^3; 



at. 

k. 

1: 



Fl. 

6. 

ct,, 



Figure 1, Distribution of the Eriocaulaceae in the United States 



Herbariuni curators who hare material of this family from additional 
counties are asked to send it to the author for verification and 
record, so that future editions of this map may be more complete 



Mapping by counties done by Andrew R, Moldenke 




1968 Koldenke, Notes on Eriocaulaceae 385 

ing in seasonally inimdated ricefielda in Northern Nigeria. Ber- 
haut (1967) cites his numbers I636 , 6h32, 6633 , & 66^1 from S6n6- 
gal. Material has been misidentified and distributed in herbaria 
as E. bongense Engl. & Ruhl. and under the name Utricularia spira- 
lis Sm. 

Additional citations: SEN&iAL: J, G. Adam 15887 (Z), 15922 (Z), 
159U7 (Z), I6968 (Z), 18U77 (Z), 18527 (Z); Raynal & Raynal 5231 
(Z, Z— dravring) , 6795 (Gg); Roberty 16336 (An). REPUBLIC OF GUI- 
NEA: Boismare ldi2 [Herb. Ghillou 3962] (An)j Chillou 7 (An), lU 
(An), 75U (Z), 789 (An, An), 935 (An), n55 (Z), 3ii35 (An), 3555 
(An), UOIO (An)i Pi tot s.n. [13.X.1950] (An). SIERRA lEONE: Af- 
zelius Hi (B — type, S — isotype, S — isotype, Z — isotype) . NIGERIA: 
Northern: C. Barter 1019 (B, S, Ut — 325). CHAD: Schvreinfurth £. 
n^ [Djur, 1869] (B). 

ERIOGAULON ALATUM H. Lecomte 

Additional bibliography: Moldenke, Known Geogr. Distrib, Eri- 
oc. 26 & 61. I9I16} Uoldenke, Phytologia 3: 181 (19l;9) and ki 339. 
1953; Moldenke, R6sum6 175, 181;, 201, L U79. 1959; Moldenke, R6- 
sumS Suppl. 1: 13 (1959) and 3: 20. 1962; Thanikaimoni, Pollen & 
Spores 7: 18 3. 1965. 

Collectors have found this species growing in savannas, de- 
scribe it as an herb Ydth yellowish heads, at 200 meters altitude, 
flowering in April, October, and December, fruiting in April, and 
called "chuk nok yung" in Thailand. 

Additional citations: THAIIAND: Bunnal 571b [Roy. Forest Dept. 
I826I4] (Bk); Larsen 8U25 (Z); S^rensen, Larsen , & Hansen 78U (Cp), 
8070 (S). WESTERN PACIFIC ISLANDS: PHILIPPINE ISLANDS: Luzon: 
Reillo 19270 (N). INDONESIA: GREATER SUNDA ISLANDS: Celebes: 
Eyaa 3383 (Ut— 11518b), 3996 (Ut— ll5lUb) . Sumatra: H. H. Bart- 
lett 7li56 (Mi) . 

ERIOGAULON ALLEIZETTEI Moldenke 

Additional bibliography: Moldenke, Biol. Abstr. 27: 2026. 1953; 
Moldenke, Bull. Jard. Bot. Brux. 27: 123. 1957; Moldenke, R6sum5 
156 & U79. 1959. 

ERIOGAULON ALPESTRE Hook, f . & Thorns. 

Synonyny: Ericaulon alpestre Merr. & Walker, Bibl, East. Asiat. 
Bot. 3U3, sphalm . 1938. Eriocaulon alpestre var. alpestre (Hook, 
f. & Themis.) Koyama, Philip. Joum. Sci. Bot. 81;: 368. 1955. 
Eriocaulon femineo-spathaceum Ruhl. ex Moldenke, R^sum^ Suppl. 1: 
17, in syn. 1959. 

Additional bibliography: Maxim., Diagn. PI. Nov. As, 8: 25. 
1892; Hook, f., Fl. Brit. Ind. 6: 578. I89U; Rxihl. in Engl., 
Pflanzenreich 13 (IV, 30): 9$. 1903; Kawakami, List PI. Formos. 
130. I9IO; Nakai, Bot. Mag. Tokyo 26: [93~9itJ . 1912; Lecomte, Fl. 
G6n. Indochine 7: 10, pi. 6D. 1922; Mak., Jap. Bot. Journ. 3: 26. 
1926; S. Sasaki, List Fl. Formos. 99. 1928; Tu, Chinese Bot. Diet., 



386 P H Y T L G I A Vol. 17, no. 5 

abrdg. ed., 1317. 1933} Uerr. & Walker, Blbl. Laat. Asiat. bot. 
3U3. 1938} Moldenke, Known Geogr. Distrlb. Erioc . 23, 2$. 26, 32, 
& 61. I9I46} Koyama, Philip. Joum. Sci. Bot. 8I4: 367—368. 1955} 
Moldenko, Rfisumfi I6I, I69, 171, 172, 175, 178, 13U, 285, & li79. 
1959} Moldenke, Rfisum^ Suppl. 1: 11 .& 17 (1959), 2: 6 (I960), and 
3: 19. 1962; Thanikaimoni, Pollen & Spores 7: I83. 1965} Moldenke, 
R6sum6 16: 21. I968. 

Koyama (1955) cites a Hayata a.n. froa Tonkin and remarks: 
"Having expected the occuirence of the present species in Indo- 
china, Lecorte included this in his Flora gSnSral de 1' Indo-Chine, 
without any citation of extant specimen from Indo-China, This 
Hayata's record may be the first one based upon a real sp)ecinen. 
E, alpestre in Ruhland's sense is composed of two taxa in the 
present days, namely E, alpestre in his meaning includes E. ro- 
bustius, a Japanese allied one. Examining E, alpestre , I, how- 
ever, found that there was not very inportant difference between 
the above two entities, and I was inclined to place E, robustius 
in a varietal rank as Maximowicz did in his first publication of 
this taxon." Koyama, therefore, recognizes E^ alpestre var. ro- 
bustius Maxim, and E^ alpestre var. niginaa (Satake) Koyama, -sAiich 
I maintain as E, robustius (Maxim.) L'ak. and E. robustixis var. 
nigrum Satake, respectively. 

Sriocaulon alpestre has been collected in bogs, at pond margins, 
and in rice paddies near carabao pastures and Ghara pools, at al- 
titudes of 50CO to 12,000 feet, floweidng in August, Coiamon nanea 
recorded for it are "hiroha-no-inunchige" , "hiroha-no-inunohige", 
"kok-cheng", and •♦kuro-inunohiga". Material has been mlsidenti- 
fied and distributed in herbaria under the names E. achiton KOna., 
E« atmm Nakai, E. j aponicum KWm., E. luzulaefolium Mart., and 
E. wallichianum Uart, The cheironymous bincmial, E^ femineo- 
spathaceuB Ruhl., was based by Ruhland on Warburg s«n. from lulu- 
po, Japan, deposited in the herbarium of the Botanisches UuseuM 
at Berlin. 

Additional citations: INDIA: Assaii: Jenkins s.n« [Assam} h.r. 
B. 3I0] (S) . Khasi States: C. B. Clarke l885la (B)} Hooker & 
Thomson 19 (B), s.n. [Mont. Khasia, 5-6000 ped.] (S, S, Ut— 3oU) . 
Sikkim: J. D. Hooker 18 (B), s.n. [Sikkia, 8-12,000 ped.] (S, 
Ut— 305). THAILAND: Smitinand 1982a (Gg) . INDOCHINA: Annaa: 
Clemens & Clemens U212 (Ca— 339315) . KOREA: Komarov 3U9 (N) . 
WESTERN PACIFIC ISLANDS: JAPAN: Honshu: Furuse s.n. [16 Sept. 
195U] (S), s^ [17 Sept. 195U] (S, S), S£U [Ih Sept. 19551 (S)} 
Saida s.n. [Matsushiro, Prov. Shinano, Aug. 1885] (B). Kiushu: 
Hayakawa s.n, (S) . Island undetermined: Warburg s,n, [Yulupo] 

TbT 

ERIOCAULON ALTO-GIBBOSUM Ruhl. 

Additional bibliography: Moldenke. Known Geogr, Distrib, Eri- 
oc. 7 & 32. I9U6} Moldenke, R6sum6 88 & U79. 1959. 



1968 Moldenke, Notes on Eilocaulaceae 387 

Additional citations: BRAZIL: Mattogrosso: Pilger 7$7 (B — type, 
Z — Isotype) . 

ERIOCAULON AJiANOANUU Koyama 

Bibliography: Koyama, Joiim, Jap. Bot. 31: 9 — 11, fig. 3. 
1956; Moldenke, R^sumfi 181 & U79. 1959; Hatusima, Mem. South. In- 
dust, Sci. Inst. Kagoshima Univ. 3 (1): 123. 1962j Moldenke, B&- 
suml Suppl. 12: 8. 196$, 

Th9 ■type of this species was collected by T. Amano ( no. k) — 
in trtiose honor it is named — at Ogimi-nura, Okinawa, in 1937, 
and is deposited in the herbarium of the National Science LIuseura. 
Koyama (1958) states that this species is related to E. latifoli\im 
J. Sa,, of Africa, but differs in its pilose receptacle, the se- 
pals of the staminate florets being glaucous-nigrescent, the anth- 
ers nigrescent, and the petals of the pistillate florets being 
smaller. He says that it resembles E. sexangulare L., which dif- 
fers in being dimerous. 

Additional citations: WESTERN PACIFIC ISLANDS: JAPAN: Kiushu: 
Hatusima & Sako 25289 (Z) . 

ERIOCAULON AUBOfiNSE Schinz 

Additional bibliography: Moldenke, Bull. Jard. Bot. Bmrx. 27: 
123. 1957; Moldenke, R6sum6 lU7, l5l, & 1^79. 1959; Moldenke, R5- 
siaaS Suppl. 3: 16 (1962) and U: 6 & 7. 1962; Friedrich-Holzhammer 
in Merzaataier, Prodr. Fl. Stldw. Afr. 159: 1 & 2. 1967. 

This species has been collected at 5UU0 feet altitude in South- 
era Rhodesia. Material has been misidentified and distributed in 
herbaria as E^. abyssinicum Hochst,, E. inyangense Arwidsson, and 
E. sexangulare L. 

Additional citations: S^N^L: £. G. Adaua 15709 (Z), 133622 
(Z). REPUBLIC OF GUINEA: Schuell 236g" (An) . RHODESIA: £. K. 
Brain liii70 (N) , 9010 (N) ; Horaky hTIWS [Govt. Herb. 13la7]"TN— 
photo); H. Wild 1162 [Govt. Herb. 15100] (N) . SOUrffwlST AFRICA: 
Baum 111 (S, Z); Pinter 7220 (S) . SOUTH AFRICA: Cape of Good 
Hope: F. A. Rogers s.n. [Moss L Rogers 1593] (S). 

SlIOCAULON AMPHIBIUU Rendle 

Additional bibliography: Moldenke, Phjrbologia 3s 181. 19U9; 
Moldenke, R6sum6 1U8 & U79. 1959. 

ERIOCAULON ANDONGENSE Welw. 

Additional bibliography: Moldenke, Known Geogr. Distrib, Eri- 
oc. 21 & 32. l9U6i Moldenke, R6sum6 1U7 & U79. 1959. 

Additional citations: ANGOLA: Loanda: Welwitsch 2]4ii3 (B — co- 
type, Z — cotype) . 

ERIOCAULON ANGUSTIFOLIUM KOm. 

Additional bibliography: Moldenke, Known Geogr. Distrib. Eri- 
oc. 7 & 32. 19U6; Moldenke, Phytologia ii: 3l;0. 1953; Moldenke, 
R^sumS 88 & U79. 1959; Moldenke, Rfesimi^ Suppl. lU: 2. 1966. 

This plant has been collected in flower and fruit in October. 



388 PHYTOLOGIA Vol. 17, no. $ 

Additional citations: BRAZIL: Brasilia: Sucre 339 [Luiza 6?] 
(Z). Golds: G. Gardner U332 [Llacbride photos lO:^^ (B— t:'-'e, JJ- 
isotype, W— photo of ty\ie) . MOUNTED ILLUSTRATIOIIS: drawings Sc 
notes by KOmicke (D). 

ERIOCAULON ANGUSTISEPALUL' }I. Hess 

Additional bibliography: Anon., Assoc. Btud. Tax. Fl. Afr. 
Trop. Index 1955: 29—30. 1956; Uoldenke, Bull. Jard. Bot. Brux. 
27: 12U. 1957} H. Hess, Bericht. Schweitz. Bot. Gesell. 67: 83. 
1957; Moldenke, R^suraS Ihl & U79. 1959; Moldenke, R6suin6 Suppl. 
1: 10. 1959. 

This plant has been collected at 1850 meters altitude in An- 
gola. 

Additional citations: ANGOLA: Huila: Antunea 168b (B); H. Hess 
52/175U (B, Z). MOUNTED ILLUSTRATIONS: fig. IklTTF) . 

ERIOCAULON ANNAMENSE H. Leconte 

Additional bibliography: Moldenke, Known Geogr. Distrib. Eri- 
oc. 26 & 61. 1916} Moldenke, R6sum6 175 & U79. 1959. 

ERIOCAULON ANNUUM Milne-Redhead 

Additional bibliography: Moldenke, Known Geogr. Distrib. Eri- 
oc. 21. 19U6; Moldenke, Phytologia 3: 131—182. 19U9; Moldenke, 
RSsum^ lUi, lliS, & U79. 1959; Moldenke, RSsumI Suppl. 1: 9 
(1959) and U: 6. 1962. 

Additional citations: REPUBLIC OF GUINEA: Boisaare li22 [Herb. 
Chillou 391*2] (An); Chillou 727 (An); Pitot s.n. [U.X.l9^0] (An), 
s.n. [13.X.1950] (An). MAFIA ISLAND: Schlieben 257U (B, N, S) . 

ERIOCAULON ANTUNESII Engl. & Ruhl. 

Synonymy: Eriocaulon antunesii Engl, ex Moldenke, R^sunS 
Suppl. 1: 16, in syn. 19^9^1 

Additional bibliography: Moldenke, Known Geogr. Distrib, Eri- 
oc. 21 & 32. 19l;6; Moldenke, R6sum6 lli7 ■Sc h79 . 1959; Moldenke, 
R63um6 Suppl. 1: 16 (1959) and U: 6 fie 7. 1962. 

Additional citations: VOLTAIC REPUBLIC: Vfinkony 3 (Z) . S^- 
GAL: Winkony 23 (Z) . IVORY COAST: Winkony 1 (Z). AlWOLA: Huila: 
Antunes 139 (B — type, B — isotype, Z — isotype) . 

ERIOCAULON APICULATUM H. Lecomte & Moldenke 

Additional bibliography: Moldenke, Bull. Jard. Bot. Brux. 27: 
125. 1957; Moldenke, Rlsum^ 156 & U79. 1959. 

ERIOCAULON AQUATILE KOm. 

Synonymy: Paepalanthus aquatilis Mart, ex Moldenke, R^sum^ 
323, in syn. 1959'. 

Additional bibliography: Moldenke, Known Geogr. Distrib. Eri- 
oc. 7, 32, & Uh. 19U6; Moldenke, Phytologia 3: 321. 1950; Molden- 
ke, R^suml 88, 323, & 179. 1959; Renn6, Levant. Herb. Inst. Ag- 
ron. 68. I960; Moldenke, Rfesum^ Suppl. Hi'- 2. 1966, 

The name, Paepalanthus aquatilis, is apparently a cheironym 



1968 Moldenke, Notes on Eriocaulaceae 389 

placed on the type collection of this taxon by Martius himself. 
The type specimen, Martius s.n. , deposited in the Munich herbari- 
um, was photographed there by Macbride as his type photograph 
nunber I868U. The species has been collected in anthesis in 
June. 

Additional citations: BRAZIL: Brasilia: Irwin & Soderstrom 
$822 (N). Minas Gerais: Martius s.n. [Macbride photos I868U] (N- 
photo of type, N — photo of type, W — ^photo of type); Sena s.n. 
[Herb. Schwacke 1U561] (B) . MOUNTED ILLUSTRATIONS: drawings & 
notes by Kt5micke (B); drawings of type collection by Martius (B). 

ERIOCAULON ARECHAVALETAE Herter 

Additional bibliography: Castellanos, Lilloa 20: 214;. 19h9i 
Moldenke, Bull. Jard. Bot, Brux. 27: 125. 1957} Moldenke, R^sum^ 
119, 285, 289, & U79. 1959. 

The Pedersen 812, distributed as E^ arechavaletae , is actually 

E. magnum Abbiatti. 

Additional citations: MOUNTED ILLUSTRATIONS: Descole, Gen. 3p. 
PI. Argent, pi. li; (N), pi. 1$ (N). 

ERIOCAULON ARENICOLA Britton & Small 

Additional bibliography: Moldenke, Bull. Jard. Bot. Brux. 27: 
125. 1957; Moldenke, R^sum^ 53 & l79. 1959. 

Additional citations: ISLA DS PINOS: Killip Ii27l5 (S); Marie- 
Victorin & Alain I66 (Vi) . 

ERIOCAULON ARISTATUH H. Kess 

Additional bibliography: Anon,, Assoc. Etud, Tax. Fl. Afr. 
Trop. Index 1955: 29—30. 1956; H. Hess, Bericht. Schweitz. Bot. 
Gesell. 67: 8>-8U. 1957; Moldenke, Bull. Jard. Bot. Brux. 27: 

125. 1957; Moldenke, R^sum^ lii7, 151, & U79. 1959; Moldenke, R6- 
sum$ Suppl. 1: 10. 1959; Friedr ic h-Holzhammer in Merxmttller, 
Prodr. Fl. Stidw. Afr. 159: 2. I967. 

Hess (1957) records this species from Southern Rhodesia. 
Friedrich-Holzhananer (1967) reduces E. welwitschii var. pygmaeum 
Rendle to synonymy under E. aristattun . 

ERIOCAULON ATABAPENSE Moldenke 

Additional bibliography: Moldenke, Bull. Jard. Bot. Brux, 27: 

126. 1957; Moldenke, R^sumS 66, 71, & li79. 1959. 

The Cruyent hi collection, cited below, is a mixture with some 
cyperaceous material . 

Additional citations: VENEZUELA: Amazonas: Cruyent h7, in part 
(Ve); Vareschi & Maegedfrau 66O8 (Ve~li2903); Ll. Williams 13858 
(Z — photo of type). 

ERIOCAULON ATRATUM KtJrn. 

Additional bibliography: Hook, f., Fl. Brit. Ind. 6: 57li. I89U; 
Ruhl. in Engl., Pflanzenreich 13 (IV, 30): 69. 1903; Fyson, Joum. 
Indian Bot. 2: 310. 1921; Moldenke, Known Geogr. Distrib, Erioc, 
2U <Sc 32. I9U6; Moldenke, Phytologia U: 3iiO. 1953; Moldenke, R6sum5 



390 P H Y T L G I A Vol. 17, no. 5 

167 & U79. 1959. 

Ruhland (1903), In his nonograph of this group, cites the type 
collection of this species aa " Gardner 972 ", but the actual type 
seems definitely to be number 932. The Collector undesi grated s. 
n. [18/10/13], distributed as E. at ra turn , is actually E. atrum 
Nakai. ~ 

Additional citations: CETLON: G_, Gardner 932 (B — type, Z — iso- 
type). 

ERIOCAULON ATRATUli var. MAJOR Thwaites 

Additional bibliography: Moldenke, Known Geogr. Distrib, Erioc . 
2U, 32, 33, 5: 38. I9U6; Moldenke, Phytologia h: 3liO. 1953; Kol- 
denke, R6sun6 167, 286, 291, & U79. 1959. 

The Herb. Koltermann s.n. specijuen, cited beloir, has steos to 
12 inches long and leafy throughout! 

Additional citations: CETLOH: Herb. Holtemaim s.n. (B); 
Thwaites C.V.131 (B — isotype, B — isotypiyi 

ERIOCAULON ATROIDES Satake 

Additional bibliography: Moldenke, Bull. Jard. Eot. Brux. 27: 
126. 1957; Moldenke, K^sumS 172 & 1;79. 1959. 

This species has been found growing in nuddy swamps. A coraaon 
name recorded for it is "kuro-inunohiga". Material has been 
misidentif ied and distributed in herbaria as E^ atrua l.aikai . 

Additional citations: WESTERN PACIFIC ISLAJtliS: JAPAN: Honshu: 
Furuse s.n. [6 Oct. 1955] (S), 3.n. [2 July 1956] (S) . 

ERIOCAULON ATROIDES f . NANUM Satake 

Additional bibliography: Moldenke, Bull, Jard. Bot. Brux. 27: 
126—127. 1957} Moldenke, Rlsum6 172 & li79. 1959. 

ERIOCAULON ATRUM Nakai 

Synonymy: Eriocaulon atratun Nakai, in herb, [not E. atratua 
K»m., 1856]. 

Additional bibliography: Moldenke, Known Geogr. Distrib. Erioc, 
25 & 61. 1916; Moldenke, Bull. Jard. Bot. Brux. 27: 127. 1957; 
Moldenke, R6sum6 171, 172, & U79. 1959; Moldenke, R^sumfi Suppl. 
3: 18 & 21. 1962; Koyama in Kitamura. Murata, k Koyana, Col. 111- 
ustr. Herb. Fl. Japan I8U— 185, pl. U8, fig. 310, text fig. 126 
(2). I96U. 

This species has been found growing in boggy pondsides. The 
Koyama plate, cited above, is in full color. The Funiae s.n. ' [6 
Oct. 1955], distributed as E. atrum , is actually E. atroides Sa- 
take, Trtiile Furuse s.n. [17 Sept. 195ii] is E, alpestre Hook, f . 
& Thorns, 

Additional citations: VVESTERN PACIFIC ISLANDS: JAPAN: Honshu: 
Collector undesignated s.n. [18/10/13] (S); Furuse s.n, [Hi Sept. 
1955] (Ca— 599I6), s.n, [2 July 1956] (S), s,n, [21 Sept. 1957] 
(S), 



1968 Uoldenke, Notes on Eriocaulaceae 391 

ERIOCAULON ATRUM var. INTERMEDIUM Nakai 

Additional bibliography: Moldenke, Bull. Jard, Bot. Brux. 27*. 
127—128. 1957; Moldenke, R^sumfi 172 & U79. 1959; Koyama in Kita- 
imra, Murata, & Koyama, Col. Illustr, Herb. PI, Japan 185. 196U. 

ERIOCAULON ATRUM var. PLATIPETALUM Satake 

Additional bibliography: Moldenke, Bull. Jard. Bot, Brux, 27: 
128. 1957; Moldenke, R6sum6 172 & Ii79. 1959. 

ERIOCAULON AUSTRALASICUM (F. Muell.) KOrn. 

Synonymy: Electrosperma australasicum F, Muell,, Trans, Philos. 
Soc. Victoria 1: 2k » 1855. Eriocaulon electrospermum F. Muell,, 
Syst. Census Austral. PI. 123. 1882. 

Bibliography: F. Muell., Trans. Philos. Soc, Victoria 1: 2U. 
1855; KOrn., Linnaea 27: 616. 1856; F, Muell., Syst. Census Aus- 
tral. PI, 123. 1882; Moore & Betche, Handb. Fl. New S. Wales khO. 
1893; Ruhl. in Engl., Pflanzenreich 13 (IV, 30): llU. 1903; Mai- 
den & Betche, Census New S. Wales Pl« 38. 1916; Moldenke, Known 
Geogr. Distrib, Erioc, 27, 32, & 3U. 19U6} Ewart, Fl. Vict, 263. 
1931; Moldenke, R6sum6 208. 28U, 286, 287, & U79. 1959; J. H. 
Willis, Handb. PI. Vict, 281, 1962; 0. D, Evans, Contrib. New S. 
Wales Nat, Herb, Fl, Ser, 27/28: 10, 1966, 

Evans (1966) describes this plant as follows: '♦anall annual 
scapigerous herb. Leaves 2~5 cm, long, ca. 1,5 am. broad, lin- 
ear-subulate, pellucid, fenestrate, 3- to 5-nerved. Scapes about 
as long as the leaves, erect, U- to 5-ribbed. Flower-heads ovate 
to subglobose, 3 — h nm, diam,; outer bracts almost lamceolate, 
obtuse to acute, glabrous; inner bracts narrow, acxnninate, glab- 
rous; receptacle conical. Male flowers central, pedicellate; 3 
outer tepals cohering at the base; 3 inner tepals fused into a 
tube, with 3 lobes at the apex each bearing a gland; stamens 6. 
Female flowers on short pedicels; perianth absent; style short 
with 3 filiform stigmatic branches. Capsule smooth, 3-locularj 
seeds solitary, smooth." He saj's that the type was collected 
by Ferdinand Jacob Heinrich von l!ueller, in December, 1853, in 
wet groiind along the Murray River towards the junction with the 
Murrumbidgee [ Nat. Herb, New South Wale s 58361 , part of the 
holotype] . He cooments "Known only from the type locality; if 
not extinct, it would be expected to occur on both the New South 
Wales (South-western Plains) and Victorian sides of the Murray 
River, Search for it is desirable." It should also be noted 
that if the above description is correot and there are really 
no sepals (as well as no petals) in the pistillate florets, 
then this species does not fit into the generic description of 
Eriocaulon and Mueller's genus Electrosperaa may well be revived 
for it, 

ailOCAULON AUSTRALE R. Br, 

Additional bibliography: Benth,, Fl, Austral, 7: 192, 1878; 
F. Muell,, Syst, Census Austral. PI. 123. 1882; F. M, Bailey, 
Syn, Queensl, Fl, 578, 1883; Moore & Betche, Handb, Fl, New S, 



392 P H Y T L G I A Vol. 17, no. 5 

Wales UjO. 1893; F. LI. Bailey, Queensl. Fl. 6: 171i>'. 1902; huhl. 
in P^gl., Pllanzenreich 13 (IV, 30): 66. 1903; V. U. Bailey, Com- 
preh. Cat, Queensl. PI. 58U. 1913; Domin, Blbl. Lot. 20: ^Od. 
1915; Maiden ?i Betche, Census lien S. .Vales PI. 3'3. 1916; Koyama, 
Philip. Journ. Sci. Bot. 8U: 368 ^t 378, pi. 6. 19t>t>i l^oldenke. 
Bull. Jard. Bot. Brux. 27: 128. 1957; Moldenke, R6anm6 169, 175, 
208, 211, '/ U79. 1959; Moldenke, K^sumS Suppl. 3: 17. 1962; 0. D. 
Evana, Contrib. New S. Wales Mat, Herb., Fl. Ser., 27/28: 10 — 11. 
1966; Moldenke, R^sum^ Suppl. 16: 12. I968. 

The specific epithet of this species is sometimes uppercaaed 
for no valid reason. Koysuiia (1955) cites Hayata 99 from Annam. 
Bailey (1913) records the common name "hat-pin plant". Bvana 
(1966) describes the plant as follows: "Annual scapigerous herb 
sprinkled with loose hairs at least on the lower parts of the 
leaves and scapes. Leaves basal, tufted, linear, up to 60 cm. 
long and 0.8 cm. wide. Scapes about half again as long as the 
leaves, ribbed when dry with 6 — 7 distinct ribs. Flower-heada 
hoary, semi-globose, changing to depressed-globose at maturity, 
up to 8 mm. wide; involucral bracts closely imbricate, broad, 
glabrous or nearly so, the margins entire or lacerate; fertile 
bracts closely imbricate, 3 mm. long, up to 3 nrni. wide, obconical, 
narrowed at the base into a short stalk, broad and rounded at 
the apex which is covered externally with a very short and dense, 
white, persistent tomentum. Flowers veiy nianerous, the male and 
female mixed together or sometimes one sex or the other predomin- 
ating, the tepals scarious or hyaline. Male flowers: outer tep- 
als 3, irregular, the 2 laterals ca. 2.5 nun. long, 0.5 nn. wide, 
the middle one linear, much narrower; inner tepals 3, equal, less 
than 1 mm. long, inserted on the receptacle close beneath the 
stamens , each fringed with a few white hairs . Stamens 3 — 6 on 
very short filaments. Faaale flowers: parts seen better in 
fruiting stage as follows: outer tepals 3, irregular, the 2 la- 
terals ca. 3 nnn. long, up to 3 nm. wide, complicate, the keel 
very broadly vringed, lacerate on the upper aiargin, the middle 
one lanceolate, concave, shorter than the laterals; inner tepals 
3, regular, ca. 2.5 ram. long, linear but with a broader base. 
Ovary 3-lobed, 3-locular} style branches 3, filiform. Capsule 
similar to the ovai7', slightly enlarged, opening by longitudinal 
slits. Seeds ellipsoid, ca. 0.8 nnn. long, brown, shining." He 
comments that the species flowers in summer "and possibly most 
of the year," growing in wet places in sandy heathland and on 
margins of swamps . From New South Wales he cites Collector un- 
designated s.n. [Nat, Herb. 58391], Constable s.n. [Jan. 1953; 
Nat. Herb. 22205] . Ingram 3.n. [Aug. I9UI} Nat. Herb. 633iiO] and 
s.n, [Jan, I96I; Nat, Herb. 633Ui4.], and Kaiden & Doorman s.n. 
[Nov. 1903; Nat. Herb. 58392]. He reports it also from Queens- 
land and Northern Territory. 

Additional citations: CHINA: Fukien: En 2lUl (Ca~288l23). 
Kwangtung: Tsang 330 [Herb. Lingnan Univ. I96II] (N), 3^ [Herb. 
Lingnan Univ. 19612] (N); Tso 21077 (N, N) . AUSTRALIAN REGION: 
AUSTRALIA: Queensland: Dallachy s.n. [Rockingham Bay] (V — 71557). 



1968 Moldenke, Notes on Erlocaulaceae 393 

State imdetennlned: Collector undesignated s.n. [Nov, Koll,] (V). 
MOUNTED ILLUSTRATIONS: Baur Icon 2U9 (V), 2^0 (V). 

ERIOCAULON BARBA-CAPRAE Fyson 

Additional bibliography: Fyson, Joum. Indian Bot. 2: 1921} 
Moldenke, Known Geogr, Distrib, Erioc. 23 f- 61. 19^6; Moldenke, 
Phytologia U: 3la. 1953; Moldenke, Rlsum^ 161 & li79. 1959; Thani- 
kaimoni. Pollen & Spores 7: 18U. 1965} Moldenke, R^sumfi Suppl. 
15: 20. 1967. 

ERIOCAULON BARBEYANUM RTihl. 

Additional bibliography: Moldenke, Known Geogr. Distrib. Erioc, 
23 & 32. 19ii6} Moldenke, R6sum6 161 & U79, 1959. 

Additional citations: INDIA: Mysore: Ritchie 12U7 (B—isotype, 
Z — isotype). 

ERIOCAULON BASSACENSE Moldenke 

Bibliography: Moldenke, Phytologia 3: 308—309 & 321, 1950; 
Moldenke, R^sun6 175 & U79, 1959. 

ERIOCAULON BAURII N, E. Br. 

Synonymy: Sriocaulon bauri N, E, Br. ex Zinderenbakker, S. Afr. 

Pollen 1: 32, 36, «c 79, pl. 7, Tig, 33 & Ui. 1953. 

Additional bibliography: Zinderenbakker, S, Afr, Pollen 1: 32, 
36, k 79, pl, 7, fig. 33 & hh* 1953; Moldenke, Bull. Jard. Bot. 
Brux. 27: 128, 1957; Moldenke, R^sumS 153 & U79. 1959; Moldenke, 
RSsum^ Suppl, 2: 9 (I960) and 3: 16, 1962} Thanikaimoni, Pollen & 
Spores 7: 182. 1965} R, H. Gompton, Journ, S. Afr, Bot, Suppl, 6: 
33, 1966} Moldenke, RfisumS Suppl, 16: 3, I968, 

This species has been collected at altitudes of 5600--6000 
feet, flowering in November. Killick states that it is "locally 
very abundant" in Natal. Gompton (I966) records it from Swazi- 
larKi. 

Additional citations: SOUTH AFRICA: Cape of Good Hope: Baur 
1166 (B—cotype, Z— cotype). Natal: Killick II6U (S) , Trans- 
vaal: F. A. Rogers 19580 (S) , 

ERIOCAULON BEAUVERDI Moldenke 

Additional bibliography: Moldenke, Known Geogr, Distrib, Erioc, 
35, 61, &62, I9I16} Moldenke, Phytologia 3: 133 (19U9), 3: 321 
(1950), and U: 3ia, 1953; Moldenke, R6sum6 88, 288, & U79. 1959. 

ERIOCAULON BENTHAMI Kunth 

Additional bibliography: H. B. Davis, Life >!^; Tiforks Pringle $6 
& 655, 1936; Moldenke, Bull, Jard, Bot, Brux, 27: 129. 1957; Mol- 
denke, RSsumS 35, 286, & U79, 1959; Moldenke, R6sum6 Suppl. k'- h 
(1962) and 12: [1] & 2, 1965; Thanikaimoni, Pollen & Spores 7: 
181. 1965, 

Recent collectors have found this plant growing in water, in 
scattered colonies in moist sandy soil of moist open wooded ra- 
vines, and in moist to wet places in low wet meadows along with 



39U PHYTOLOGIA Vol. 17, no. $ 

GypeiTus , Eleocharis , Juncua , Mimulua , P.anunculu3 , Spiranthaa , 
TrlTollum , crassea, etc., at altitudes of l^OO — 22$0 meters, 
flofwering in July. litis and hia asaoclates describe the flowers 
aa "chalk-wJiite". 

Material has been mi a identified and distributed in herbaria aa 
E. humboldtii Kunth. On the other hand, the Hitchcock h Stanford 
7201 and Pr ingle 2665 , distributed as E. benthami, appear to be 
E. ehrenber^ianum Klotzsch. The E. benthani of Sc'nlechtendal is 
a synonym of E. ehrenbergianum Klotzach. 

Additional citations: LOUISIANA: Beauregard Par.: R. L. Crock - 
ett 8280 (Ld) . IffiXICO: Chihuahua: E. W. Helaon 6028 (W— 35?7U5) • 
Federal District: Collector undesignated s .n. [Chapultepec, Au- 
guat 31, 1872] (W— U5278). Jalisco: Hartweg 258 (B—type); R. L. 
McGregor 16617 (Lw) ; R. McVaugh 20U73 (Mi). Mexico: Gilly , 
Alexander , & Hernandez Xolocotzi 83 (Mi); Hlnton 627 (S), 3638 
(Rf , S) , U5U9 (Rf . Rf , Ur) i Matuda 30855 (Z) ; Pringle 11371 (Mi) . 
Michoacin: litis , Koeppen, 4 litis U09 (S). Nayarit: J_, N_, Rose 
s.n. [Aug. 8, 1897] (W--3U2909). Tlaxcala: Arsene 1725 (W— 
IO327UI). Veracruz: Pringle 11871 (Gg--U23U00, W— IIS1266, W— 
1586U88) . 

ERIOCAULON BIFISTULOSUM Van Heurck i Muell.-Arg. 

Additional 5t emended synonymy: Eriocaulon flultans Griff., 
Itin. Notes [Posth. Papers 2:] 65. I8U8. Eriocaulon flultana J. 
G. Baker, Joum. Linn. Soc. Lond. 3ot. 20: 227. 1893. Eriocaulon 
limosum Engl. £: Ruhl. in Engl., Bet. Jahrb. 27: 7U. 1399. Erio- 
caulon schwa infurthii Engl. & Ruhl. in Engl., Bot. Jahrb. 27: 7U. 

1899. 

Additional fie emended bibliography; Griff., Itin. Notes [Posth. 
Papers 2:] 65. 13U8; A. Chev., Sudania 1: 7. 1911; Moldenke, 
Known Geogr. Distrib. Erioc. 20—23, 27, 32, 3>h, & 39. 19U6; Mol- 
denke, Bull. Jard. Bot. Brux. 27: 129. 1957; Anon., Biol. Abstr. 
29: 32U8. 1957; H. Hess, Bericht. Schweitz. Bot. Gesell. 66: 87— 
88. 1957; Moldenke, RSsum6 133, 13U, 136— 138, liiO, 11^6— IL18, 156, 
161, 208, 288, 289, 292, & U79--U80. 1959; Razi, Rec. Bot. Surv. 
India 18: 19. 1959; Moldenke, R^siuil Suppl. 3: 15 & 16. 1962. 

The binomial, E. limosua Engl, t Ruhl., is a cheironymous des- 
ignation placed by Riihland on the specimen of Barter 1021 from 
Nupe, Nigeria, in the Berlin herbarixaa, while the same workers 
apparently regarded Schweinfurth 22it , from '♦am Biri, Dar-Fertit, 
Gaaalquellen-gebiet", in the same herbarium, as the type of 
their E. schweinf urthii . 

Hesa (1957) claims that E. bifistulosum shoiild be induced to 
synonymy under the earlier E. melanocephalum Kunth of the New 
World, affirming that the differences which I list as differenti- 
ating the two taxa do not hold up. He records the taxon from 
Angola. Griffith (I31i8) records it from Assam. Recent collectors 
have found it at altitudes of 1000 — 1830 meters. Milne-Redhead 



1968 Moldenke, Notes on Eriocaulaceae 395 

& Taylor describe it as a "plant vaiying in height and size de- 
pending on age and depth of water; leaves very delicate, entirely 
sutmergedj heads blackish", growing in water 3 — $0 cm. deep in 
the open, rooting in crevices of laterite, flowering in April. 

Chevalier (1911) cites his no, 302. Material has been mis- 
identified and distributed in herbaria as E. heudelotii N. E. Br. 
€ind E^ melanocephalum Kunth. 

Additional citations: MALI: Collector vuidetermined s.n. [26. 
11.16] (An). CHAD: Schwelnfurth 2U76 (B, S) . SSNaSAL: J. G. 
Adam nlih2 (Z) ; Couey 1 (Z) ; Rsynal & Raynal 68 U6 (Z) . REPUBLIC 
OF GUINEA: Boismare IxlQ [Herb, Chillou 3938] (An); Chillou 1737 
(An) J Schuell 21la (An). NIGERIA: Northern: C. Barter 1021 (B— 
isotype, S-— isotype, Ut — 321; — isotype). TANGANYIKA: Milne- 
Redhead & Taylor 9929 (B, S) . AiraOU: Huila: H. Hess 52A678 
(B). PORTUGUESE EAST APRIGA: Gazaland: Schwelnfxirth 22U (B) . 
MADAGASCAR: Loher s.n. [Tananarive, 11.1911] (Mu--395) • 

ERIXAULON BUOBATUM Morong 

Additional & emended bibliography: H. B. Davis, Life & Works 
Pringle 9U & liil, 1936; Moldenke, Known Geogr, Distrib, Erioc, 
U, 32, & 36. 19U6; Moldenke. Bull. Jard. Bot. Drux. 27: 130. 
1957; Moldenke, R^sum6 3^, Ul, 289, & U30. 1959; Moldenke, R6su- 
m6 Suppl. 7: 3. 1963} Langman, Select, Guide Lit. Flow, PI. Max. 
911. 196U. 

Recent collectors have found this plant growing on slopes by 
streams in meadows in pinewoods, at altitudes of 2300 — 2li00 
meters, floweidng in September. Ruhland, on a label of the type 
collection in the Berlin herbariimi, states that he feels that 
this is the "E. sexangulare Auct." and that the taxon is probab- 
ly conspecific with Ej. cinereua R. Br. (which he cadled E, sie - 
boldtianup Sieb, & Zucc). The two taxa are certainly very sim- 
ilar. Pringle 6llt6 is a mixture with E. schiedeanum KOm. 

Additional citations: MEXICO: Durango: Moore & Bunting 8690 
(Z). Jalisco; Barnes S: Land 159 (S); R. McVaugh 17578 (Mi); 
Pringle 3855 (B — isotype, B — isotype, Ca — 2Ul5 — isotype , Ms — 
l5U65~isotype, S— isotype), 6llj6, in part (Ca— 115172), 6299 
(B, Ca— 115173, Ms— 151;6U, S, S) . GUATEMAU: Jutiapa: J. A. 
Steyermark 30U05 (W— 2022037) . 

ERIOCAULON BLDMEI Kt5m, 

Additional bibliography: Moldenke, Known Geogr. Distrib, Erioc. 
27 & 32. 19U6; Moldenke, Rlsum^ 190 & ii80. 1959} Moldenke, R^su- 
m6 Suppl. 1: 13. 195?. 

Backer found this plant growing at 1725 meters altitude, flow- 
ering in October. Koorders is of the opinion that E. blumei is 
conspecific with and should be regarded as a synonym of E. 
brownianum Mart, and he so identified Pulle 3079 . 

Additional citations: INDONESIA: GREATER SUNDA ISLANDS: Cele- 
bes: Eyma li009 (Ut— Il5l6b), li009 bis (Ut— Il5l5b) . 



ICONOGRAPIIIA CYPF.RACEARUM IT 

Tetsuo Koyama 
The New York Botanical Garden 



As the second part of this series of illustrations the pre- 
sent paper presents ten species, which are: Carex doenitzii , C. 
curvicollis , C. mitrata, C. Breviscapa, C. jackiana (ssp.). C. 
brownii , C. olivacea (ssp.), C. idzuroei, C. michauxiana (ssp.)j 
and Scirpus juncoides (ssp.)- The selection of these species 
was made from the species that are endemic to the Japanese flo- 
ristic region, or from those of which the main area of distri- 
bution lies in that particular floristic region. 

It has been known among the cyperologists that the genus 
Carex shows its highest endemism in the Japanese floristic re- 
gion. In 1962 I made a revision of Japanese Carices. In my 
revision of Japanese Carices (1962)** an emphasis was made on 
the taxonomic relationships of Japanese species with those of 
other floristic regions to elucidate the actual endemism of 
Japanese Carices. The following table reflecting the high en- 
demism of Carex in the Far East was based on 422 species that I 
recognized to be valid in my above-mentioned study. The Japa- 
nese floristic region as regarded here includes the Japanese 
Archipelago from Kuriles to northern Ryukyus, Korea and the 
montane region of Formosa. 

I would like to express my appreciation to Miss Josephine 
H. Ueno, who so ably typed my sometimes difficult manuscript in 
the form that suits the off-set printing. 



* Part I. Phytologia 15 (3): 201-221, pl£. 1-10. 1967. 

'* Koyama, T. Classification of the Family Cyperaceae (2). 
Journ. Fac. Sci. Univ. Tokyo, III, 8(4): 149-278. 1962, 

396 



1968 Koyama, Iconographia Cyperacearum 397 

Table 1. Geographical distribution of Carex occurring 
in the Japanese floristic region. 



1. Species endemic to the Japanese floristic region 59.3% 

2. Far Eastern species: 

2a. Species extending to eastern 

Siberia and Manchuria 16.3% 

2b. Species extending to Central China 4.9% 

3. Discontinuous species: 

3a. Species also occurring in Indian 
Himalayas, Southern China and 
occasionally as well as in Malaysia 5.0% 

3b. Species also occurring in North America 3.7% 

3c. Species also occurring in Australia 

and adjacent Malaysian Archipelago 0.8% 

4. Wide species: 

4a. Circum-polar and circum-boreal species 6.5% 

4b. Eurasian species 2.9-s 

4c. Cosmopolitan species 0.6% 

Plate 11. CAREX DOENITZIl BOckeler 

Carex doenitzii BOckeler, Flora 65: 61. 1882. 

Synonymy. Carex plocamostyla Maximowicz, Mel. Biol. 
12:565. 1887. 

Carex dicuspis Franchet , Bull. Soc. Phi lorn. Paris 8®ser. 
TT 42. 1895. 

Carex chrysolepis Franchet ^ Savatier var. modesta L^vei- 
ITe § Vaniot, Bull. Acad. Intern. Geogr. Bot . 10: 
279. 1901 

Carex dicraea C. B. Clarke, Kew Bull. Add. Ser. 8: 71. 
1907^ 



398 



PHYTOLOGIA 



Vol. 17, no. 5 







Plate 11. Carex doenitzii Bockeler 



1968 Koyama, Iconographia Cyperaceartun 399 

Carex nagatadakensis Masamune, Fl. Geobot. Stud. Yaku- 
shima, 526. 1934. 

Carex doenitzii Bockeler var. mollis Akiyama, Journ. 
Jap. Bot. 13: 648. 1937. 

Tufted in clumps; rhizome short, erect or erect -ascending, 
divided; roots densely yellow-hairy. Leaves radical, narrowly 
linear, shorter than to slightly longer than culms, linear, 3-5 
mm wide, flat, rather soft, lightly green above, densely papillo- 
se and white-powdery beneath, gradually acute at apex; sheaths 
dorsally reddish- or purplish-brown, ventrally yellow-brown, the 
basal sheaths short-bladed or cataphylloid; ligule auriculate, 
membranous. Culms slender, 10-60 cm tall, ca. 1 mm thick, 3- 
sided, scaberulous, erect or slightly curved above. Spikes 2 to 
4; upper 1 or 2 staminate, clavate, 0.7-2 cm long, 3-6 mm thick, 
deeply purple-brown; other spikes pistillate or rarely androgy- 
nous, obovate to elliptic, 1-3 cm long, 5-7 mm thick, densely 
many-flowered, cemuous or filiform peduncle, the lowest one of- 
ten spaced and long-peduncled, the upper ones somewhat approxi- 
mate and short-peduncled. Lower 1 or 2 bracts leaf-like equal- 
ling or slightly exceeding the inflorescence, not sheathing at 
base. Pistillate glumes lanceolate or ovate-lanctolate, 6-10 mm 
long, deeply purplish-fuscous, deeply red-purple or rarely gre- 
enish, gradually tapering above to an acute aristate apex; the 
awn scabird 0.1-0.2 mm long, the costa green, 3-veined. Perigy- 
nia slightly shorter than glume, erect, lanceoblong or ovate-ob- 
long, 4-6 (-10) mm long, unequally biconvex, membranous, lightly 
ferrugineous-green to yellowish-green, densely dotted with red- 
dish purple, sparsely hispidulous on both sides, minutely serru- 
late on both margins, contracted at short-stipitate base, gradu- 
ally tapering above to a long flattish beak, the orifice deeply 
bifurcate, the teeth shortly awn-shaped, serrulate. Achenes 
rather tightly enveloped, obovate-elliptic, biconvex, 2-2.2 mm 
long, contracted at both ends; style elongated, long-exserted 
beyond the orifice of perigynia; stigmas 2, filiform, up to 6 mm 
long, persistent. 

Voucher specimen: Japan, Mt . Fuji, U. Faurie 15570 (KYO) . 

Wet open grassland or on rocks in subalpine regions. Dis- 
tribution Endemic to Japan; from Hokkaido southwards to central 
Japan and southwestwards to Toyama Prefecture of Japan Sea side 
of the Mainland. 

Plate 11. A. Total plant; B. Staminate spike; C. Staminate 
glume; D. Pistillate spike; E. Perigynium at anthisis; F. Stami- 
nate glume; G. Perigynium; H. Basal part of perigynium; I. Por- 
tion of the margin of perigynium; J. Tooth of the orifice of 
perigynium; K. Achene with style-base. 



Uoo 



PHYTOLOGIA 



Vol. 17, no. 5 



'^ II / 



/ <>i 



^ 




Plate 12. Carex curvicollis Franchet 5 Savatier 



1968 Koyama, Iconographla Cyperacearum UOl 

Plate 12. CAREX CURVICOLLIS Franchet § Savatier 

Carex curvicollis Franchet S Savatier, Enum. PI. Japon. 2: 579. 
1879. 

Synonymy. Carex viridula Franchet § Savatier, Enum. PI. 
Japon. 2: 2: 151 § 579. 1879. Not of Michaux, 1803. 

Carex Savatieri Franchet, Nouv. Archiv. Museum 3^ ser. 
rO: 71. 1898. 

Densely tufted in large clumps; rhizome short, branching, 
stoloniferous; stolons slender, covered with reddish-brown sca- 
les, the inter-nodes 0.5-0.8 cm long; roots densely yellow-pube- 
scent. Leaves many, narrowly linear, 2-4 mm wide, equalling or 
somewhat shorter than culms, flattish, soft, lightly green; she- 
aths pale-brown to reddish-fuscrous, the basal ones short-bladed 
to cataphylloid, eventually split into soft brown fibers. Culms 
slender, acutely 3-angled, 10-35 cm long, soft, smoothish, in- 
clined to nodding. Spikes 3-6 (-8], all approximate to subfasti- 
giate; terminal spike staminate, clavate, 0.8-2.5 cm long, 1.5-3 
mm thick, purplish-fuscous, erect on a short peduncle or nearly 
sessile; lateral spikes pistillate, oblong to cylindrical, 1.5- 
4 cm long, 5-8 mm thick, densely many-flowered, the upper 3 or 4 
subsessile or short-peduncled, the lower ones on a capillary 
elongated peduncle, the lowest spike sometimes with 1 or 2 addi- 
tional spikes at the base of the body of the spike through bran- 
ching. The lower 1 or 2 bracts leaf-like, equalling to slightly 
longer than inflorescence, the upper bracts glumaceous or seta- 
ceous, none seathing at base. Pistillate glumes ovate, 1.5-2.25 
mm long, deeply purplish-fuscous to pale-ferrugineous, contract- 
ed to obtusish or mucronulate apex, the costa green, obscurely 
3-nerved. Perigynia much exceeding glumes, erect to erect- 
patent, lanceolate, 4-5 mm long, compressed trigonous, thinly 
membranous, pale-green, weakly and densely many-veined, smooth, 
glabrous, suddenly contracted at obtuse short-stipitate base, 
gradually tapering above to a long terete often slightly recur- 
ved beak, the orifice truncate or somewhat emarginate. Achenes 
loosely enveloped, elliptic-obovate, compressed-trigonous, 1.5- 
1.7 mm long, contracted at both ends; style elongated, slender, 
subpersistent, not thickened at base; stigmas 3, 2.5 mm long, 
recurved. 

Voucher specimen: Japan, Mainland, base of Mt. Bukosan in 
Saitama Prefecture, ca. 850 m alt., T. Koyama 6,703 (NY). 

Plate 12. A. Total plant; B. Prophyll at the base of pedun- 
cle; C. Staminate glume and its triandrous flower; D. Apex of 
anther showing the connective; E. Perigynium at anthesis; F. G. 
Pistillate glumes; H. Dorsal view of perigynium; I. Orifice of 
perigynium; J. Fruiting pistil showing mature achene. 



U02 



PHYTOLOOI/l 



Vol. 17, no. 5 




y -^ 










.A 




Plate 13. Carex mitrata Franchet 



1968 Koyama, Iconographia Cyperacearvnn U03 

Plate 13. CAREX MITRATA Franchet 



Carex mitrata Franchet, Bull. Soc. Philom. Paris 8^ ser. 7: 
887 1895. 

Tufted in large clumps with divided slender ascending rhi- 
zome. Leaves rather many, slenderly linear, stiffish, 1.5-2 mm 
wide, grass-green, folded, longer than culms, gradually attenu- 
ate to acute tip; sheaths yellow-brown or brown, eventually 
weakly split into brown parallel fibers. Culms many, slender 
but stiffish, 6-30 cm tall, obtusely trigonous, nearly smooth, 
glabrous. Spikes 3-5, the upper 3 or 4 approximate to contigu- 
ous at culm apex, the lowest one often basal; terminal spike 
staminate, linear, 5-15 mm long, 0.8-1 mm wide, whitish-brown- 
ish, short-peduncled to nearly sessile, slightly exceeding the 
next pistillate spike; lateral spikes pistillate, densely many- 
flowered, narrowly cylindrical to oblong, 0.5-1.5 mm long, 2- 
2.2 mm thick, green, erect, nearly sessile or on a short pedun- 
cle enclosed in bract sheath. Bracts short, setaceous or spa- 
thaceous, much shorter than the subtending spike, the base 
hardly or short- (±0.5 mm) -sheathing. Staminate glumes oblong, 
whitish-brownish, the margins broadly white-hyaline. Pistill- 
ate glumes oblong to oblong-obovate, 1.25-1.5 mm long, ca. 2/3 
mm wide, membranous, pale, truncate at hyaline apex, the keel 
green, 1-nerved, projecting beyond the glume apex into a short 
upright mucro. Perigynia nearly erect, slightly longer than 
glume, fusiform-obovate to fusiform, 2-3 mm long, ca. 1-75 mm 
wide, 3-sided, membranous, lightly green, weakly many-veined, 
sparsely pubescent, the base gradually attenuate, the apex con- 
tracted to a short conical occasionally slightly incurved beck, 
the orifice minute, nearly entire. Achenes tightly enveloped, 
elliptic, triquetrous, 1.5 mm long, the sides shallowly concave 
below, the beck discoid-annulate, 0.25 mm wide; style thickened 
at base; stigmas 3, short, slender. 

Voucher specimen: Japan, Mainland, Urawa in Saitama Prefec- 
ture , T . Koyama 6881 (NY) . 

Grassy hillsides somewhat sheltered by loose woods. Dis- 
tribution. Endemic to Japan, central and western Mainland, Shi- 
koku, Kyushu and southern Korea. 

Plate 13. A. Total plant; B. Staminate spike; C. Stami- 
nate glume; D. Pistillate spike; E. Pistillate glume; F. Peri- 
gynia; G. Achene. 



hOli 



PIIYTOLOGIA 



Vol. 17, no. $ 




llii-S w 



Plate 14. Carex breviscapa C. B. Clarke 



1968 Koyama, Iconographia Cyperacearum Uo5 

Plate 14. CAREX BREVISCAPA C. B. Clarke 

Carex breviscapa C. B. Clarke, Fl. Brit. Ind. 6: 736. 1894. 

Synonymy. Carex jackiana Boott var. breviculmis Thwai- 
tes 5 Hook, f., Enum. PI. Zeyl . 356. 1884. 

Carex curtisii Ridley, Mater. Fl. Malay Renins. 3: 117. 
T90T. 

[Carex obtuso-bracteata Hayata, Icon. PI. Formos. 6: 131. 
1916"! Nomen nudum . J 

Carex lutchuensis Ohwi, Mem. Coll. Sci. Kyoto Univ. B, 
5: 270. 1940. 

Densely tufted from short erect rhizome clothed with dark 
brown parallel fibers; roots rather stout. Leaves many, crowd- 
ed, all radical, linear, elongated, 4-7 mm wide, 25-60 (-90) cm 
long, herbaceous, somewhat roughned above, 3-costate, flat-pli- 
cate, gradually tapering above to long acute apex, the base 
short-sheathing, dark purplish-brown on veins, eventually dis- 
integrating into dusky-brown fibers. Culms much shorter than 
the leaves and almost hidden in leaf tussocks, 10-20 (-30) cm 
tall, slender, obtusely 3-angled, ca. 1 mm thick, 3- to 6-nod- 
ed, bearing spikes from above the base. Spikes usually panicu- 
late through branching; terminal spike staminate, slenderly li- 
near, 1-2 cm long, 1 mm thick, pale-brownish, erect on a short 
peduncle, equalled or surpassed by the next lateral spike; late- 
ral spikes pistillate or with short staminate part at apex, nar- 
rowly cylindrical, erect to erect-patent, 1-3 cm long, 3-4 mm 
thick, loosely many-flowered, the upper spikes short-peduncled 
or nearly sessile, the lower ones on exserted peduncles. The 
lower 2 or 3 leaf-like bracts elongated, much exceeding the in- 
florescence, reaching 40 cm in length; upper bracts much re- 
duced, short-bladed or spathaceous, the sheathing base 1-2.5 cm 
long. Pistillate glumes elliptic-oblong or ovate-oblong, 2-3 
mm long, 1-1.75 mm wide, membranous, pale- or whitish -brown, 
white on hyaline margins, obtuse or rounded at cilio-late apex, 
faintly several-nerved on both sides, the midvein greenish end- 
ing below the glume apex or projecting beyond the glume apex 
into a short straight mucro 1/2 to 1 mm long. Perigynia about 
twice as long as glumes, rhombic- lageniform, 3-sided, 3.75-5 mm 
long, broadest at about the middle, 1.25-1.5 mm wide, membra- 
nous, pale-green, glabrescent or sparsely puberulent on the up- 
per half, slenderly many-nerved, cuneate at short-stiped base, 
contracted above to a short conical beak 1/2 to 1 mm long, the 
orifice 2-toothed. Achens tightly enveloped, rhombic -fusiform, 
2.5-3 mm long, triquestrous , 1.25-1.5 mm wide, cuneate at base, 
gradually narrowed above the middle to transversely truncate 
apex with annulate margin ca. 1 mm in diameter; style short. 



U06 



PHTTOLOGIA 



Vol. 17, no. 5 




J fCff-^a^HiyO., dtt. /fUf. 



Plate 15. Carex jackiana Boott 
subsp. parciflora Kukenthal 



]_968 Koyama, Iconographia Cyperaceanira UO? 

often with thickened base; stigmas 3. 

Voucher specimen: Formosa, Taipei-hsien, Shirin, Ohwi Hb^ 
TNS 55,925 (TNS) . 

Undergrowth of dense forest in the tropical high mountain 
zone and on hills in the subtropics. Distribution. From Cey- 
lon through Malaysia eastwards to northern Queensland of Austra- 
lia, and northeastwards to Annam, Formosa and the Ryukyu 
Islands . 

This species is recognizable at once by the slender short 
culms hidden in the well-elongated leaf blades as well as m 
the peculiar lageniform perigynia. 

Plate 14. A. Total plant; B. Pistillate glume; C. Dorsal 
view of perigynium; D. Lateral view of perigynium; E. Trans- 
verse section of perigynium at the level marked A - B m Fig. 
D; F. Dorsal view of achene; G. Annulate apex of achene. 



Plate 15. CAREX JACKIANA Boott subsp. PARCIFLORA KUkenthal 



Carex jackiana Boott subsp. parciflora (Boott) Kiikenthal, 

Pflanzeru. 4(20), Cyper-Caric. 638. 1909. Incl. 
forma ochrolepis (Franchet) Kiikenthal. 

Synonymy. Carex par