Annals
of the
Missouri Botanical
Garden
Volume XLIII
iD 5:6 ¡A
With 11 plates, 91 figures, and 4 maps
Published — at Galesburg, Illinois, by the Board of Trustees of
the Missouri Botanical Garden, St. Louis, Mo.
Entered as second-class matter at the post-office at Galesburg, Illinois,
under the Act of March 3, 1879.
Mre@eva! "ToTANIEXT
GARDEN LIONARY*
Annals
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Missouri Botanical Garden
A Quarterly Journal containing Scientific Contributions from the
Missouri Botanical Garden and the Henry Shaw School of Botany of
Washington University in affiliation with the Missouri Botanical
Garden.
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The ANNALS OF THE MissourRI BOTANICAL GARDEN appears four times
during the calendar year: February, May, September, and November. Four
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Single Numbers ___.___-_-__ 2.50 each
Contents of previous issues of the ANNALS OF THE MissourI BOTANICAL
GARDEN are listed in the Agricultural Index, published by the H. W. Wilson
Company.
TABLE OF CONTENTS
A Revision of the Genus Erechtites (Compositae) SA Sec into
Senecio and A E. Belcher
The Australian and New Zealand Species of Pittosporu
Ro
bert C. Cooper
The Cultivated Beans of the Prehistoric Southwest
Lawrence Kaplan
A ee of Rauvolfia, with Particular Reference to the American
pe Aragula Sathyanarayana Rao
A New Fossil Plant from the New LT Shale, with Some Comments
on the Origin of Land Vascular Plants
Part 1. Crocalophyton, A Soest Transitional Sea-Land Plant...
ry N. Andrews and Karen S. Alt
Part 2. Some Comments on the MR of Land Vascular Plants
and the Taxonomic Position of Crocalophyto Mes SES
ry N. Andrews
A Note on the Nodal Anatomy of Ankyropteris er eae EE
ry N. Andrews
Some Lichens of Tropical Africa. IL. Ussea Carroll W. Dodge
General Index to Volume XLIII
PAGE
I= 85
87—188
189-251
253—354
355-363
363-378
379-380
381-396
397-399
The publication of this paper was made pook through a special subvention from
the od fund of the National Academy of Scienc
Errata in Vol. XLIII:—
Page 299, line 8—Insert "T:" before Stork. ` `
Page 309, caption of fig. 11—Castaneda's number should be goo.
Page 316, line 8—After “RB,” insert "S,".
Page 347, line 1—For rbombofae, read rbonbofae.
Page 354, last line, "illustration" should read "specimen".
Page 382, 14 lines from bottom, Pereoensis should read hereroensis.
Page 385, line 28, “mm.” should read "cm."
Page 394, line 5, "annulatus" should read "annulata".
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Volume XLIII Number 1
Annals
of the
Missouri Botani
Garden
Annals
of the
Missouri Botanical Garden
A Quarterly Journal containing Scientific Contributions from the
Missouri Botanical Garden and the Henry Shaw School of Botany of
Washington University in affiliation with the Missouri Botancal
Garden.
Informaton
The ANNALS or THE Missournr BOTANICAL GARDEN appears four times
during the senem: year: February, May, September, and November. Four
numbers constitute a volume.
ipm Price rS per volume
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Contents of re issues of the ANNALS OF THE Missourr BOTANICAL
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Company.
Annals
of the
Missouri Botanical Garden
Vol. 43 FEBRUARY, 1956 No. 1
A REVISION OF THE GENUS ERECHTITES (COMPOSITAE),
WITH INQUIRIES INTO SENECIO AND ARRHENECHTHITES *
ROBERT O. BELCHER
INTRODUCTION
This paper is an outgrowth of a study of the botanical collection of the United
States of America Typhus Commission, India-Burma Field Party, which collection
is now part of the U. S. National Herbarium. It was made by the author and others
from June to November, 1945, in the vicinity of Ledo, Assam, and Myitkyina,
Burma, mostly from the immediate environs of military camp-sites as part of an
investigation of the epidemiology of severe outbreaks of scrub typhus in those areas.
Although a preliminary report of the botanical studies was made to the Com-
mission in 1946, it was never published and the final evaluation of the role of plants
in these epidemics has been deferred pending more accurate identification of the
species involved, This has been much more difficult than anticipated, because there
is no adequate modern floristic treatment of the area from which the plants came.
A further complication has been the unsatisfactory nomenclatural status of
many of the species which fall into the category of so-called pantropical weeds and
are regarded by many botanists as too well known and widespread to be worthy of
serious consideration. Detailed examination of several such “species” has led to the
conviction that such weedy genera are quite generally in need of critical review.
paper is part a dissertation presented to the Rackham School of Graduate Studies,
doctor
* This
University of Michigan, in a paral fulfillment of the requirements for the degree of
science. e gui of Prof. H. H. Bartlett, Dr. Rogers McVaugh, and Ge of the doctoral
The able guidance
ittee is per te ti
mare ch of the examination of herbarium material was carried out during the academic year
1951-52, ES ma author was a ¿De me at Bedford na a London, and the Royal
of the rich experiences of that year was published in 1953
Botanic Gardens
(Asa oo “Bull. ie SCH 13- 16).
bed April 15, 1956.
(1)
[Vol. 43
2 ANNALS OF THE MISSOURI BOTANICAL GARDEN
The present paper deals with a complex of genera centered around the genus Erech-
tites Raf. in the subtribe SENECIONINAE of the Compositae
The Compositae are represented in the weedy flora of the tropics by numerous
species, among them Erechtites hieracifolia Raf. ex DC. The fact that numerous
specimens of the African Crassocepbalum crepidioides (Benth.) S. Moore in the
U. S. A. T. C. collection were originally determined as Erechtites hieracifolia led to
the discovery that most Asiatic specimens were in fact wrongly determined as the
latter species in the major herbaria studied. An effort to clarify the status of these
two species in the weed flora led to a critical study of the genus Erechtites in the
New and Old World. An attempt was made to locate and compare type material
for every binominal published in the genus, and to collate the major treatments of
the genus. This revision has also necessitated inquiries into related genera, notably
Senecio L., Arrbenecbtbites Mattf., Crassocephalum Moench and Gynura Cass.
The status of the latter two genera was discussed by me in a recent paper!.
ERECHTITES
HISTORY OF THE GENUS
Erecbtites was originally published by Rafinesque? in 1817, based on an un-
named plant described by C. C. Robin,® and it contained one species, E. prealta
Raf. Sprengel* in 1826 in effect reduced the genus to Senecio by reducing the
species to S. bieracifolius L. Cassini? in 1827 placed Erechtites Raf. in section
OTHONNÉES of the SENECIONEAE, with "péricline uniserie, tres-simple, nu à la
base", because of Rafinesque’s description: ‘Perianthus . . . denudatus .
perianthe neither caliculate nor sphacelate". Cassini had earlier (in 1820 and 1825)
based the well-defined genus Neoceis on Senecio bieracifolius L.
In 1831 Lessing? identified a Schiede and Deppe specimen as "Erecbtbites
pracalta”, and cited "Senecio bieracifolius L.” and “Sonchus agrestis Sw. fl. ind.
occ.!” as synonyms. In 1832 he published” a thoroughly revised diagnosis for
“Erechthites Raf.”, in which the distinctive characters, including the appendaged
style-arm apex, are clearly brought out, and reduced Neoceis Cass. to synonymy.
He recognized two species, E. praealta Raf. and E. cacalioides (Fisch.) Less.
De Candolle? in 1838 published “Erechtites Raf. fl. ludov. (excl. car.) ", with
a à diagnosis much more resembling Lessing’s and Cassini’s than that by Raf-
inesque. Neoceis became a section of Erechtites containing six species from the
Americas. Three new sections were added by de Candolle to receive a total of thir-
1 Kew Bull. 1955: 455—465.
Martinique Se ra Saint Wiese pier t les années 18
T!
le,
02, 1803, 1804, 1805, et 1806. Vol. III.
* Caroli Linnaei Syst. Veg. ed des 2 565. 1826.
5 Dict. Sci. Nat. E
$ Linna 1831.
T Syn. Gen. Comp. e 1832.
Tech 6:295. 18
1956]
BELCHER— REVISION OF GENUS ERECHTITES 3
teen species from Australasia, including two treated by Cassini under Neoceis and
a number previously published as Senecio.
It is significant that although botanists in general, other than Schultz Bipon-
tinus and Mueller, have not openly questioned de Candolle's treatment of Erech-
tites, de Candolle himself included in the diagnosis of each of the three Austral-
asian sections a query as to the propriety of including it in Erechtites! Those doubts
were fully justified. The indigenous Australasian and Indonesian species must be
excluded from the genus Erechtites as here defined.
Comparison of the types and other material of the American species with those
of the Old World has led to the conviction that they are not really congeneric,
just as de Candolle suspected. The American species, with the exception of two
which clearly are discoid species of Senecio, agree in having the style-arm apex
prolonged into a tuft of fused papillose hairs, which is quite similar to the somewhat
longer appendage that helps to characterize Crassocepbalum. The native Old
orld species, with the exceptions noted below, all agree in having the style-arm
apex bluntly truncated or rounded, and nude or with a marginal fringe of diverging
papillae; that is, indistinguishable from the style-arm apex characterizing the genus
Senecio. The American material is heterogamous. Within the Australasian material
occurs every degree of transition from the outer two rows of florets pistillate and
filiform to all florets perfect and infundibuliform, or to outer florets ligulate and
pistillate; in other words, there is intergradation without a break into both the
discoid and the radiate groups of Senecio
The name Erechtites was first ES to and must be typified by American
material. Therefore the non-congeneric Old World species must receive another
name. In the absence of any clear-cut distinction between the majority of these
species and other Australasian species which have universally been referred to Se-
necio, I have transferred them, except for six species, to that genus, to which, indeed,
most of them were originally ascribed. The only alternative to broadening some-
what the limits of Senecio is to set up a new and ill-defined segregate, necessarily
impossible of separation except on minute, arbitrary, and unreliable features, a
course which I reject.
The return of all species of Erechtites to Senecio has twice been briefly ad-
vocated. Schultz Bipontinus? found in a specimen of his Senecio flavus (S. Decais-
nei DC.) from the Canary Islands some slender non-ligulate pistillate margi
florets. On the mistaken assumption that the sole difference between Erechtites
and Senecio was in the filiform pistillate marginal florets of the former, he swept
s the species of Erechtites and of certain other related genera back into cio.
Yet his figure of S. flavus in Barker-Webb and Berthelot's Flora of the Canary
Islands!? clearly shows the style arm bluntly truncated with a corona of divergent
hairs, indicating that it is a Senecio and not an Erechtites! Furthermore, two of the
three otherwise identical specimens of S. Decaisnei in the Prodromus Herbarium,
9 Flora 28:497-498. 1845.
10 Hist, Nat. Iles Canaries 2:319. £. 107. 1836-50.
[Vol. 43
E ANNALS OF THE MISSOURI BOTANICAL GARDEN
duplicates of those cited by Schultz, have ligulate marginal florets whereas the
third does not, and all have senecionoid style arms.
A somewhat similar position was taken independently by Baron Ferdinand von
Mueller*! at one time for the Australasian species of Erechtites. He listed nineteen
species of Senecio under cultivation in the Melbourne Botanic Garden, of which
two, S. bispidulus A. Rich. and S. quadridentatus Labill, were revived names for
species which both earlier and later he treated in Erechtites. His argument was
presented by Bentham!”: “F. Mueller proposes to unite the two genera Erechthites
and Senecio on account of those supposed intermediate species forming de Candolle's
section Plagiotome". Bentham rejected this view, and Mueller resumed use of the
name Erechtites. But later!?, in describing Senecio baplogynus from New Guinea,
he stated:
ntham and J. Hooker observed eiit hong decns II. 208), that occasionally some thin
solely D grits occur in specie: ; hence the only characteristic which separates
¡Erec s from that genus is enrehsble, A pss the present plant ma y be placed in either
csv
This applied to the Australasian material, but overlooked the nature of the style-
arm apex which further distinguishes the American species
Exceptions to this reduction of Old World species to Senecio are the six species
to be referred, one for the first time, to Arrhenechthites Mattf. These species, in-
cluding Senecio baplogynus, are well distinguished from Senecio, Erechtites, Crasso-
cepbalum, and Gynura by their functionally staminate center florets with short
style arms which are densely papillose on their outer faces and are not appendaged.
The range of Arrbenecbtbites, formerly known only from the mountains of New
Guinea, now includes the Blue Mountains of southeastern Australia
All this leads to the problem in the Candollean treatment of Erechtites posed by
Merrill:!*
is generic name has been accepted by all authors, and yet i rather ous to note that
Candolle, Prodr. 6: 294. 1838, in his full eg of the genus, Pee "Erechtites Lm fl. ludov.
(1817 p. 65 (excl. char.)’. = grand should be interpreted to Be E dx
ae aha type E. prealta Rat. placed as a variety of E. HG er 2 "Raf. ex DC."
In undertaking the EE of the ‘Prodromus’, de Candolle had solicited
specimens from all and sundry, promising to give full credit for either specimens
or names. The scrupulousness with which he carried out this promise is a chal-
lenge to his successors and a reproach to more than one. But in this case it led him
into difficulties. There is clear evidence in the Prodromus Herbarium that de
Candolle originally considered the American species which he finally placed in
Erechtites to belong to Neoceis Cass. This name in his writing appears repeatedly
11 Catalogue of plants under es in the nem Botanical Garden, October, 1858, in
General eebe of the Government Botanist for 1858. p. 1 is is the Cat. Hort: Melb.
cited by am (Fl. Austral. 3:659) for | Sach Lessonii F. "Muell.
SR E 3:659. 1866.
13 Trans. Roy. Soc. Victoria 17:14-15. 1889.
14 Index Rafinesquianus, p. 235.
-
1956]
BELCHER— REVISION OF GENUS ERECHTITES 5
on the sheets of the first six species. In some instances there is no other generic
designation except on the outside label of the cover, which of course was collated to
agres with the published text of the ‘Prodromus.’ He probably also — Cas-
sini’s relegation of the faultily drawn Erechtites Raf. to section OTHONNE
This nicely settled state of affairs was rudely upset in 1830 by a Ae from
Rafinesque, in which he placed S. hieracifolius in a new subgenus. This note, of
crucial importance in evaluating both Rafinesque’s and de Candolle’s interpreta-
tions of Erechtites, is reproduced here for the first time. It is found in a lengthy
folio letter entitled: "Serie de Lettres Botaniques addressées au Prof. Decandolle de
Genéve, par le Prof. Rafinesque de Philadelphie. N. 5. Octobre 1830 philadelphie,”
and preserved in the library of the Conservatoire de Botanique, Genéve. The
writing has faded somewhat, so that some words were only doubtfully deci-
phered. These are identified by bracketed queries or reconstructions. I am greatly
indebted to Dr. Charles Baehni, Director of the Conservatoire, for locating the
letter for me, to him and Dr. C. E. B. Bonner for help in transcribing the passage
in question, and to Prof. H. H. Bartlett for suggestions on deciphering it. It ap-
pears to read as follows:
"13. Senecio L. Il faut restituer le G. [genus] Jacobea T. pour less S. de: Së Esp. [spec cies] ont
z rapas 3 Ae u]s [?, aopa > pieni GE 3 fids, S. G.??—Les vrais ee
p. bieracifolius, ierg Raf. & elon ormant mon S G. [subgenus]
Boch SZ Gen. fl. lud) Per. fol. ovals ^ aia 51 vix calic.-flosc. fem. vix [radiis
[?, radiatis ?] filiformi[bu]s vix L5 dentat
The intention here seems clear Bees The radiate species of Senecio are to be
returned to Jacobea Tourn. (this is an old and recurrent argument), with S. bal-
samitae perhaps the type of a subgenus on account of its trifid rays. The true (ie.,
discoid) species of Senecio in the United States are sufficiently distinct from the
Old World species (as exemplified by S. vulgaris) to be placed in a separate sub-
genus containing three species. This subgenus Erechtites of the genus Senecio as
here redefined is so changed from the 1817 version as to be scarcely recognizable.
Formerly it was ecalyculate, with florets hermaphroditic; now it is scarcely calycu-
late, with feminine florets filiform. That it is still applied to the same material as
was the original genus Erechtites is supported not only by his explicit reference to
its former status but also by the change of gender of the original specific epithet
to agree with the new generic name.
This letter, in my opinion, is the one referred to by de Candolle "7 in a footnote:
"Cl. Rafinesque in litt. 1832 [i. e., 1830] forte non immerito distinguit Erech-
tites sp. A, hieracifolia, cds fl. lud. et elongata, sed descriptiones nullas
edidit." The letter would seem. to be the authority for the new combination,
"Erechtites hieracifolia (Raf. in = )”. That Rafinesque distinguished three
species of Erechtites; namely, hieracifolia, praealta, and elongata, at least in the
1830 letter, is scarcely accurate. What he actually did was to designate three dis-
coid species which he regarded as together forming a subgenus of Senecio. That he
15 Prodr. 6:294. 1838.
[Vol. 43
6 ANNALS OF THE MISSOURI BOTANICAL GARDEN
wrote nothing of their descriptions was but natural, since all three had been de-
scribed previously, by Linnaeus, by Rafinesque (as Erechtites prealta), and by
Pursh, respectively.
Now that Rafinesque had identified his genus with S. hieracifolius as a sub-
genus of Senecio, the revised diagnosis, while still leaving much to be desired, was
recognizable as applying to the Linnaean species. I believe that de Candolle wished:
(a) to maintain Erechtites (Neoceis) as a genus, rather than a subgenus; (b) to
accept Erechtites in sensu 1830 rather than 1817; and (c) to credit the name to
the original author. But the Erechtites of 1830 was unpublished. De Candolle's
solution was to adopt the earlier and validly published name but to exclude the
faulty diagnosis, substituting for it a new one based on that of the now super-
fluous Neoceis Cass., which he demoted to a section. I believe de Candolle meant
"Erecbtites Rafin . . . (excl. car.)" literally, and with justification. About the
only feature in common between his diagnosis and the one published by Rafinesque
is that in both the heads are described as multiflorous!
Early references to Erechtites in floristic works for North America were erratic.
W. J. Hooker in 1834 and Darlington in 1837 both cited “Erecthites [sic!] praealta
Raf.—Less." as a synonym of S. bieracifolius, which Darlington, however, correctly
described as having "heads discoid, branches of the style conic at the apex". Torrey
and Gray in 1845 took up the Candollean treatment of Erechtites, with “E. bieraci-
folia (Raf.)” the only species. Darlington in 1853 gave “Erechtites Rafin.”, but
followed his own and other previous treatments in describing the capitula as discoid
but the “florets all fertile".
Grisebach, who, in 1861, was first to bring both the temperate and the tropical
forms of E. hieracifolia together under one specific epithet, gave in his generic diag-
nosis an accurate characterization of the two types of florets in the capitulum. His
description of the style branches as conical at the summit would restrict the cover-
age to the American species only, and indeed he did not mention the many Austral-
asian species added by de Candolle and others.
Baker in the ‘Flora Brasiliensis’, in 1884, gave a generic description in many
respects excellent, but unfortunately maintained the genus in the extended sense,
with "styli rami elongati apice truncati vel obtusissimi”. He adduced several new
synonyms for E. hieracifolia, maintained Grisebach's varieties, but with new di-
agnoses, and described a new species, E. ignobilis, as well as treating E. valerianae-
folia. Gardner had earlier (in 1845 and 1848) described new species of Erechtites —
and of Senecio from his Brazilian collections without defining the generic limits
explicitly, His two species of Erechtites were later reduced, one to a variety, the
other to synonymy.
Fernald in 1917 described E. megalocarpa from Cape Cod, and at the same time
revised Erechtites in temperate North America, establishing three varieties of E.
bieracifolia. Cronquist in 1946 reduced E. megalocarpa to a variety of E. hieraci-
folia, a change not recognized by Fernald, Standley and Steyermark in 1947
1956]
BELCHER—REVISION OF GENUS ERECHTITES 7
published without comment E. agrestis (Sw.) as a new combination for E. caca-
lioides Less.
In the southern hemisphere, meanwhile, Malme in 1899 described E. missionum
from northern Argentina, to which Chodat and Hassler soon added a var. lanceolata.
Cabrera recently transferred Senecio leptantbus Phil., S. goyazensis Gardn., and S.
valerianaef olius Gardn. to Erechtites.
Thus, since 1838 numerous new species have been described as Erechtites, but
without critical review of the whole genus. During this time considerable con-
fusion has developed in the application of names, circumscription of species, and
delimitation of the genus. As a further complication, there is evidence of introgres-
sion among certain of the supposed species, the extent of which can only be deter-
mined by field and laboratory studies yet to be made.
. Thesearch for typifying material was mostly successful, although a few critical
specimens are apparently lost. The lack of a type for E. prealta Raf. posed a major
problem. The many Australasian sheets in Kew Herbarium which were annotated
y Hooker, or Bentham, or both, in the preparation of their treatments of this
group, were gone over carefully, and every effort was made to collate them with
the types, particularly of A. Richard and de Candolle. Several misapplications of
names and erroneous circumscriptions which have passed from the works o
Hooker and of Bentham very generally into the taxonomic literature of Australasia
have thus been detected and corrected. Numerous other adjustments in the nomen-
clature of this long-neglected group have had to be made.
The proper spelling of the generic name has long been debated. Rafinesque
states that the name “Erechtites” was "one of those given by Dioscorides to the
Senecio". But the spelling used by Dioscorides in the ‘Materia Medica’ was
"epexÜiris" and the arguments by Bentham!? and Hegi” for using “Erechthites”
are perfectly valid. Under the International Rules a case could be made for cor-
recting Rafinesque's spelling as an unintentional orthographic error. If the genus
were to be maintained in the extended Candollean sense, then the Benthamian
spelling, widely adopted in works on Australasian botany, might well be retained.
But since the genus should be restricted to the American species, it seems better
to retain the original spelling of Rafinesque, used by most American authors, the
"Index Kewensis’ and the Gray Herbarium Index, and Dalla Torre and Harms.
— CONDENSED KEY TO THE SENECIONINAE
The tribe sENECIONEAE may be characterized as having no latex, corollas of
the disc florets actinomorphic, anthers ecaudate, pappus capillary (or rarely set-
aceous) , and style arms usually with a crown of diverging pollen-presentation hairs
near the truncated, obtuse, or papillose-appendaged apex (but not in Ligularia,
16 Fl, Austral. 3:657. 1866.
17 fll, Fl. Mittel Europa 6?:701. 1929.
[Vol. 43
8 ANNALS OF THE MISSOURI BOTANICAL GARDEN
Homogyne, Gynura, Cacalia (Adenostyles) , Psacalium) .
The subtribe sENECIONINAE O. Hoffm. is further distinguished by having the
phyllaries free or very nearly so, and the involucre uni-or biseriate, or if pluriseriate
then with the style arms of the perfect florets truncated and penicillate or with a
distinct crown of longer hairs.
The following key is based on that given by Hoffman,!? but has been somewhat
condensed and extensively modified to accommodate the following groups: Arrben-
echthites Mattf.; Crassocephalum Moench confused with Gynura by Hoffman;
certain erechthitoid species of Senecio, considered by him as included in Erechtites;
and four genera revived by Rydberg!? for the North American species commonly
referred to Cacalia: Psacalium Cass., Pericalia Cass., Mesadenia Raf., and Odonto-
trichum Zucc. emend. Rydb.
KEY TO THE SENECIONINAE
A. Receptacle with paleae Schistocarpha, N.
(cf
rolae
esoe spp. >)
AA. E without paleae.
. Disc
S st erile
we: GE not Pa alike in floral composition, plants being. more or less
d
or polygamo Petasites, Robinsonia, re
CC. etiim all “alike in floral composition, plants being neither dioecious nor poly-
L
D. "Herbs with leaves radical or decur
E. — sticky-haired herbs with Ce lavet 0 cs S oer amd
EE. s with leaves radical; scapose stems beset with scale-like leaves.............. Tussila
DD. Under or perenn =A her bs with cauline leaves well developed, not renkon Ze F
F. rous ius shru eege ëch fleshy
FF. ege csset pu herbaceous or suffruticose; leaves not fleshy
ARRHENECHTHITES
BB. Disc florets fe OS G
ipitula wei? mm florets perfect; margin florets pistillate, with corolla tubular,
rm, or irregularly eem A ue lat
H. Leaves radical, cau = e lea coles Homogyne, Stilpnogyne
HH. Leaves cauline, n ale- "ike : I
Leaves entire, thickly crowded; branches leafy to base of capitulum..................
E Leaves es more or less toothed or divided, scattered -reduced-to bracts in
orescence ..............
BE Pistillate te florets SE 4-fid; shrubs
JJ- pues ae filifo.
tyle
abes crown re divergent hairs surrounding appendage % fused
T
J
Faujasia
i ose
St Sie arm pes crown of divergent hairs one truncated or roma tly
Dein d, not pne se echthitoid species a SENECIO
GG. Capitals "Esco with all florets perfect ee or radiate, with marginal
tres unes ^ dor s or dy "bilabiate
Pappus
=
Raillardella
(cf. Dubautia and Raillardia
EL ue menie dine
. Phyllari any-seria Alciope, Culcitium, Lepidospartum
MM. Phyllaries i or SC Ge posesa often 1-seriate, ani shorter calyo ulus "
18 Hoffman, O., in Engl. & Prantl’s Die Nat. Pflanzenfam IV®: :286-289. 1894.
19 Bull. Torr. Bot. Club 51: 369-378, 409—420. 1924.
EE Erotbrix ee
1956] ;
BELCHER—REVISION OF GENUS ERECHTITES 9
N. Leaves opposite, or only the uppermost alternate
Mallotopus, Haploestbes, Arnica, x gai
NN. edis mem or radical
O. Receptacle hemispherical, gie "gite arched, or flat with the center
mër pesa al prolon
P; SAPE radiate, LR various but not du and rees central
ocidium, Bartlettia, Doronicum
PE; oan discoid, receptacle flat, with center produc
prolongation; corolla white or whitish,
throat and long lobes; achene terete, UM
Mesadenia (Cacalia $ Conophora DC.)
n a conical
with Perrin campanulate
OO. Receptacle flat
. Phyllaries with leaf-like appendage along midrib ........................ Lopbolaema
QQ. Phyllaries Mire leaf-like appendages R
R. Capitula radia S
$. a: SE bilabiate, inner lip short, outer strap-like........
ipid
SS. Marginal florets ligulate, without inner lip, Raid entire or 2- t
3-dentate e $
gU Dezem weng ere sed Cineraria
TT. Achen sed but prismatic or cylindrical, 4- or 5-
add or md to Gees ed
U. Style arms virtually verus or at most T papillose
below the truncated or obtuse apex, with crown of diver-
gent hairs but See appendage ........ e speci SE Senecio
UU. Style arms thickly hairy on most or all of the outer fac V
V. Capitula in racemes or racemose-paniculate reen
Ligularia
VV. Capitula solitary, nodding (in one species discoid)
Cremanthodium
RR. Capitula discoid (marginal florets sometimes with anthers variably
abortive)
Y
W. Style-arm apices with greatly prolonged, non-stigmatic, fre-
quently recurved appendage x
X. Appendage va Manet densely hairy on outer face; leaves
i abire crowded basally and subpetiolate or sessile; stem
Mise
ot vascularized; basal leaves peltate .............-.. sacalium
ain P.
WW. v E apices with appendage not exceeding arm in length,
or not appenda
Y. Style-arm apex obtuse, unappendaged, lacking crown of
divergent hairs Z
Z. Style arm not hairy, but oblong and rem adus
ia L. (Adenostyles)
ZZ. Style arm uniformly beset = “ang D uaa T
acaliopsis, Luina, Peucephyllum
YY. i finis apex penicillate Sc crown Ze am papil-
rs which is sometimes indistinct re florets S
a. Achene i cae dbge? or obovoid b
b. Achene inverted- -pyram Psatbyrotes
bb. Achene id. elliptic in cross-section, 10- to 15-ribbed; corolla white or whitish
with tube long and slender, throat obsolete, lobes long-linear and extending to t throat;
basal leaves long-petiolate
indrical €
c. Capitula axillary or in axillary panicles Bedfordia
c. Capitula terminal or in terminal and axillary inflorescences d
[Vol 43
10 ANNALS OF THE MISSOURI BOTANICAL GARDEN
d. Style arms thickly hairy on all of outer surface; capitulum solitary, nodding
E, "m "
sp.
dd. Style arms glabrous or at most minutely papillose on outer surface e
. Style-arm apex truncated or low-domed, not appendaged, crown of divergent
hairs usually well developed f
f. Corolla white or whitish, throat elongate and narrowly funnelform, lobes
conspicuously revolute after anthesis, pappus barbellate, slightly clavate........ Pericalia
ff. Corolla yellowish, infundibuliform, lobes usually erect, pappus not clavate
discoid species of Senecio
ee. Style arm acute with more or less prominent non-vascularized appendage of
fused papillose hairs, crown of diverging papillae sometimes imperfectly formed
or nearly wanting g
g. Capitulum ecalyculate
h. Apex tufted with a multiseriate crown, the conical apex slightly pro-
longed in appendage of 3 to 6 short fused papillae; corolla deeply lobed;
low canescent shrubs
b Tetradymia
hh. Style apex with essentially uniseriate crown, conical apex prolonged in
appendage up to half as long as the style arm proper; corolla not deeply
Emilia
obed; herbs
gg. Capitulum calyculate i
i Style-arm appendage about as long as or longer than the style arm
fal TS:
ii. Style-arm appendage not more than half as long as the style arm
j. Appendage short, conica
Jj. Appendage ovoid, capitula large
EAS
SYSTEMATIC TREATMENT OF ERECHTITES
ErecHTITES Raf. Fl. Ludovic. 65. 1817. Type: E. prealta Raf.
Erechthites Raf. emend. Less. Syn. Gen. Comp. 395. 1832
32:
Erecbtites Raf. sensu extenso DC. Prodr. 6: 295. 1838; Endl. Gen. Pl. 455. 1836-40; Dalla
orre & Harms, Gen. Siphon. 561. 1906.
Erechthites Raf. sensu extenso Benth. & Hook. Gen. Pl. 2:443. 1873; Hoffm. in Engl. &
Prantl, Nat. Pflanzenfam. IV®: 291. 1894.
Neoceis Cass. Bull. Sci. Soc. Philomat. 1820:90. 1820; Dict. Sci. Nat. 34: 386, 1825 (pro
majore parte).
Senecio spp. L. et auct.
Ptileris Raf. Am. Month. Mag. 268. 1818, nom nud.; ex Jacks. Ind. Kew. 4:657. 1895,
nom. nud.
Annual or with perennial rootstock; roots fibrous; aerial shoots herbaceous or
slightly woody at base, erect, sulcate, leafy, glabrous or variously pubescent. Leaves
alternate, subpetiolate, decurrent, or semiamplexicaul, rarely petiolate, subentire,
serrate, variously incised or lobed, or pinnatifid, acute, glabrous or variously pubes-
cent. Capitula calyculate, heterogamous. Involucre uniseriate, more or less flask-
shaped; phyllaries linear or lanceolate, equal, plurinervate with margins scarious,
connivent in aestivation and anthesis, more or less divergent in fruit, finally diver-
gent and strongly xed. Receptacle always without paleae, inconstantly plane,
alveolate, or fimbrillate. Marginal florets filiform, 4- or 5-fid, pistillate or some-
times with rudimentary stamens in some, style-arm apices without marginal fringe
1956]
BELCHER—REVISION OF GENUS ERECHTITES 11
of pollen-presentation hairs but with apex conic-appendaged; central florets slen-
derly infundibuliform, 5-fid, perfect with functional ovary, style-arm apices diver-
gent with a semi-circular crown of pollen-presentation hairs surmounted by a
terminal appendage of fused papillose hairs more or less prominently developed,
sometimes nearly wanting in section GOYAZENSES. Achene subcylindric to sub-
fusiform, base inconspicuously callose-annulate, apex conspicuously callose-annu-
late, slightly constricted below the annulus but not attenuate-rostrate, about 10-
ribbed, ribs prominent, cartilaginous, light brown, glabrous, intercostal planes dark
brown, sparsely puberulous. Pappus pluriseriate, subequal, capillar
Distinguished from Crassocepbalum, Gynura, Cacalia (sensu strictiore), and
discoid species of Senecio by the heterogamous rather than homogamous capitula,
from radiate species of Senecio also by the corollas of the filiform marginal florets
more or less regularly 4- or 5-fid rather than ligulate, and from erechthitoid (and
all) species of Senecio by the style arm appendaged with fused papillose hairs rather
than truncated or bluntly rounded; and from Arrhenechthites by the disc florets
being numerous and developing viable embryos, rather than few or solitary and
abortive.
Includes five species: one highly polymorphic and widespread in the less-
elevated or more humid parts of both Americas and adventive in central Europe,
Hawaii, Indonesia, and southeastern Asia; one widespread in continental tropical and
subtropical America and adventive through many of the tropical Pacific Islands to
northern Australia, Sumatra, the Malay States, and southeastern China; and three
restricted to South America; not known from Africa. Formerly considered to
include several species with filiform pistillate marginal florets, indigenous to
Australasia, but these, without exception, lack the terminal appendage of fused
papillose hairs on the style-arm apex and are accordingly here excluded from Erech-
tites, and treated as belonging in Senecio or (in one instance) in Arrhenechthites.
SECTIONS AND SPECIES OF ERECHTITES
Erechtites may be divided into two sections:
ERECHTITES sectio Hieraciifoliae Belcher, sect. nov.
Annuae, herbaceae; foliis caulinis non ultra octies longioribus quam latioribus;
corolla lobata, non profunde fissa.
Type: Erecbtites bieracifolia (L.) Raf. ex DC.
ERECHTITES sectio Goyazenses Belcher, sect. nov.
Perennes, suffruticosae; foliis caulinis decies longioribus quam latioribus vel
eene appendicibus styli florum marginalium non obviis; corolla lobata,
profunde
Type: pube goyazensis (Gardn.) Cabr.
[Vol. 43
12 ANNALS OF THE MISSOURI BOTANICAL GARDEN
KEY TO SPECIES OF ERECHTITES
A. Annual, herbaceous; length of cauline leaves usually not over eight times their width;
marginal florets with style-arm appendages well developed; corolla shallowly cleft
$ HIERACIIFOLIAE
B. Leaf sessile, or if briefly petiolate then petiole conspicuously alate; diameter of capi-
tulum about Ya its length; marginal florets bi- or pluriseriate; pappus white 1. E. bieracifolia
BB. Leaf petiolate or inconspicuously alate at base.
C. Diameter of capitulum less than V4 its length; marginal pistillate florets uniseriate
or sub-biseriate; style-arm appendages long; corolla and pappus usually more or less
reddis E. valerianaefolia
CC. Diameter of capitulum Y, its length or more; marginal pistillate florets bi- or
pluriseriate; style-arm appendage short; corolla and pappus whitish............ E. missionum
inear-lanceolate . goyazensis
DD. Capitulum thick, 2 cm. long, 1 cm. wide, or larger; corolla lobes deltoid
5. E. ignobilis
$ HIERACIIFOLIAE
1. ERECHTITES HIERACIFOLIA (L.) Raf. ex DC. Prodr. 6: 294. 1838.
Annual herb, 4 cm. to 2 m. tall, simple or much branched above; stem glabrous,
setose, or pilose. Leaves alate and attenuate at base or semiamplexicaul, acute, with
8 to 25 indistinct pinnate veins on either side, each diverging from the midrib at
about 45^, running irregularly forward and terminating in an acute callose tooth;
oblanceolate, oblong-lanceolate, lanceolate, or linear-lanceolate; margins subentire,
irregularly serrate, incised-serrate, sinuate-dentate, irregularly and coarsely lobate-
dentate, or subpinnatifid; glabrous or minutely pubescent on nerves beneath or
sparsely to densely clothed with fine unicellular or coarse multicellular hairs up to
5 mm. long in pilose states of var. cacalioides; membranous to subcoriaceous; lower
ones from 1 to 10 to 30 cm. long, 0.5 to 2 to 7 cm. wide, gradually or abruptly
reduced upwards, Inflorescence varying from a single terminal capitulum in
depauperate specimens to decompound terminal and axillary corymbose panicles of
50 or more capitula; branches glabrous to pilose, bracts ranging from broadly
amplexicaul to attenuate. Capitula with bracteoles of the calyculus linear, variable
in number and length, and glabrous or ciliolate with unicellular or multicellular
hairs and either with or without multicellular hairs on their outer faces. Phyllaries
8 to 11 (to 15 to 15) mm. long, 0.5 to 1.0 (to 2.0) mm. wide, broadly linear or
oblong, entirely glabrous to sparsely setulose; apices bluntly acute to subobtuse,
often darkened but not sphacelate, usually minutely papillose-cilioclate. Florets
numerous, varying from 20 to 25 (to 150 or more) ; corolla of marginal florets in
var. bieracifolia and var. cacalioides 7.5 to 11 mm. long with throat 0.5 to 1 mm.
long and only slightly dilated to diameter of 0.5 to 0.75 mm., 4- or sometimes 5-
fid, lobes 0.4 to 0.5 mm. long, in var. megalocarpa similar, 10 to 12 mm. long;
corolla of disc florets in above varieties 8 to 12 mm. long, throat 1.5 to 2 mm.
long, slenderly campanulate, with circumference of 1.5 to 2.0 mm., 5-fid, 10-
nerved, lobes 0.5 to 0.7 mm. long; corolla lobes in disc florets of var. megalocarpa
1956]
BELCHER—REVISION OF GENUS ERECHTITES 13
11 to 13 mm. long, very slenderly campanulate, throat 1 mm. long, 5-fid, lobes
0.75 mm. long, 0.4 to 0.5 mm. wide. Achenes strongly costate
Species highly polymorphic, but divisible into three varieties, Widely distrib-
uted from southern Canada to northern Argentina, but absent from the prairie,
high montane, and Pacific coastal regions. Unknown from Africa, the Near East
and Australasia. Its reputation as a pantropical weed rests largely on confusion
with Crassocepbalum crepidioides (Benth.) S. Moore.
Several departures of the foliage from the narrowly defined type have been
given varietal or even specific rank. In fact, Erechtites itself was founded on one
such variant. After examining a large suite of specimens, both dried and living, I
have concluded that foliar characters do not offer a reliable basis for separation of
varieties (nor for the establishment of formae) within this species. I have seen
every degree of variation in the leaf from the subpetiolate oblanceolate to the
broadly amplexicaul lanceolate state without finding at any point an obvious and
usable discontinuity. Furthermore, I have been able to duplicate much of the
spectrum of variation in greenhouse culture by manipulation of the environment.
The same series of foliar variation is clearly discernible in plants both of tropical
and of temperate habitat.
I have based the two main varieties maintained here on features which do appear
to be discontinuous and to show a geographical pattern. The most accessible of
these features are the length of the calycular bracteoles and the type of pubescence.
That it has been necessary to use so slender a distinction will perhaps in itself be
sufficient answer to those who, without extensive examination of the full range of
variation, would establish separate species for the extremes of the temperate and the
tropical zones.
KEY TO VARIETIES OF ERECHTITES HIERACIFOLIA
A. Bracteoles of the calyculus extending less than 1⁄4 the length of the involucre; brac-
teoles of the peduncle much shorter than the involucre; all bracteoles glabrous or beset
with clas hairs only.
B. Ge pitula with denuded ‘receptacles less than 1 cm. in diameter; achene 2.5 t
long, 10-costate ] a. var. ' bieracifolia
BB. Capitula wis desde e more than 1 cm. in diameter; achene 4 to 4.5m
long o 20- var. Siet
AA. Bracteles of E Pha a extending more than Y, the length of the inv volucre; bra
teoles of the peduncle approximately as long as the involucre; all brio ciliolat
with: ier hairs var. wf R
la. ERECHTITES HIERACIFOLIA Var. HIERACIFOLIA
ets po L. Sp. Pl., ed. 1. 866. 1753; ed. 2. 1215. 1762; ed. 3, 1215. 1764;
L. Man «DT, ek i is var, cacalioides; non Forssk. Fl. Aegypt. 73. 1775;
= » Walt. Fl. Carol 208. 1788, a radiate species; non Herb. Labill., which is Senecio
inimus Desf. e E
eren laevis Sloane, C t. Pl. Tam. 122. 1697; Nat. Hist. Jam. 255. 1707 (vol. 5, p. 1
Sloane Herb. BM!, Rot record of temperate var. ‘hieracifolia i in the Caribbean, surely
not in Madeira
Anonyme Robin, Voy. Louisiane 3:435. 1807, basis of E. prealta Raf.
(Vol. 43
14 ANNALS OF THE MISSOURI BOTANICAL GARDEN
Erecbtites brealta Raf. Fl. Ludovic. 65. 1817
enecio seminudus Bory, Ann. Gén. Soc. Phys. 1 : 303-307, pl. 12. 1819; figure pra
Lee CS non- ica duced upper leaves. described by Fernald "for var. inte
Neocei ‘Bieracifli (L.) Cass. eng Sci. Soc. Philom. 1820: 91. 1820; Dict. Sci. Nat. 34:
87. 1825; type of Neoceis Cas
Mus oie Cass. 1. 6: by desc) also e ieu to var. intermedia F
Senecio bieracifolius E. var. gigantea Raf. Med. Fl. 2:262. 1830; by dag a very robust
Senecio Vukotinovici Schloss. Oesterr. Bot. FR 31: 5. 1881; Schloss. ex Vukot. Rad.
Jugo slav. Akad. 58: 85, 145. 1881
gap sonchoides => Fl. Exsicc. So pe n. 658. 1881 UA ae on Schlosser’s
ous; non Kunt ov. Gen. & Sp. 4:
Erechthites —— (L.). DC. var. glabrescens O. Ktze. tea tie = 1:335. 1891,
Brechtbites bierecifolia Raf. forma minor Waisb. Oesterr. Bot. Zeit. 45:109. 1895; said to
he it o the typical state by “Uebergangsformen”; Waisbecker (US!) agrees
ct é
Prileris hirracifolia Raf. ex Jacks. Ind. Kew. 2:657. 1895; nom. n
Erecbtites bieracifolia (L). Raf. in DC. var. typica, var. byHWlie Fern., and var. praealta
Raf.) Fern. Rhodora 19:27. 1917; Gray's Man., ed. 8. 15 1950.
yt y: Coope e 936.
Seedling Morphology: Kumner, Weed S dip 381-382, 1951.
mmon names: Fireweed; Butterweed (Ken tucky Mountains).
Lectotype: “Senecio hieracifolius,’ 996-1 (LINN!) ?°
Habit, stem, and foliage as in the species. Bracts subtending the peduncles and
peduncular bracteoles usually not as long as the capitulum, often not half as long;
calycular bracteoles linear, very short, glabrous or sometimes minutely ciliolate with
unicellular hairs, never having coarse multicellular hairs; phyllaries (7 to) 11 to
16 (to 21), linear, glabrous or minutely glandulose-papillose. Denuded receptacles
(3 to) 5 to 8 (to 9) mm. in diameter. Achenes 2.5 to 3 mm. long, 0.5 mm. in
diameter, 10-costate.
Prince Edward Island to southeastern Saskatchewan to Minnesota, south to
southeastern Texas, to Florida; of scattered occurrence in the West Indies from
Cuba to Puerto Rico and possibly through the Lesser Antilles to Venezuela and the
Guianas; an introduced weed in the Hawaiian Islands and Central Europe; pri-
marily an occupant of recently disturbed areas in forest zones.
CANADA. PRINCE EDWARD ISLAND: Bunbury, Aug. € ders: Long & St. John
8242 (K). QUEBEC: St. Se Estuary, Portneuf, Aug. arie-Victorin 28309
uses Victorin 28657 e S) Ge Park Skyline Trail,
July 1941, Senn 2078 (S). ONTARIO: Battersea, Aug. 1898, Fowler (F) ; Galt, bogs, Aug.
1899, Umbach m a Pak. Aug. 1900, an 21827 (F) SASKATCHEWAN:
Alameda, Drummond (K).
a MAINE: Mt. Megunticook, Camden, Aug. 1930, Steyermark 2181 (F);
Megunticook Lake near Tsuga Ledges’ Sept. 1946, Friesner 20982
Cambridge, waste Get: e 22, — Smith & Steward in Pl. Exsicc. Gray 899
var. intermedia CH, S) ; Cape Cod near Nine Mile Pond, Sept. 1898, Green-
an 399 (K); Vastu Ter Woods. S Sept. 1915, Ripley 15816 (S); S. Dartmouth,
Herbaria in which cited specimens are located are indicated by abbreviations recommended by
Lanjouw alios onal Code of Bam Nomenclature adopted by the Seventh International Bo-
ca Congress, Stockholm, 1950).
1956]
BELCHER—REVISION OF GENUS ERECHTITES 15
Salter's Point, Sept. 1917, King 146 (F); — sandy field, Aug. 1930, Beattie (K, det.
as var. praealta Fern., very depauperate). CONNECTICUT: New Haven, 1858, Eat on, (S);
(S). i C
arsh, Dor, 6, H Geméen 7999 (F). rithis dnt Bethlehem, pag 1832, Moser
E FI); Philadelphia, Sept. 1849, Prior (K) ; West Chester, Darlington (K, FI) ; Dauphin,
Sept. 1887, Small (F); Lancaster Co., near teri — 1892, Heller 8 gren i
Strattonville, Oct. 1928, Eggleston 22838 8 (US). onto: Elyria, Gbale banks of B
CH .
Mosely (F); Lawrence Co., clay pit south of Blackfork, Sept. 1952, Belcher 1115, 1116,
1117; same location, Oct. 1952, Belcher 1122 to 1140 inclusive (series to show range of
variation, with achenes for planting). INDIANA: u 1890, Hill 116 (F);
ells Co., near Bluffton, Aug. 1898, Deam (F); Miller, base of dune, Sept. 1900, Lansing
119 , 2); mora, Sept. 1935, Friesner 9 Mots: Lake
we View
(Chicago), Sept. 1884, Oblendorf (F). MICHIGAN: New Richmond, Aug. 1910, Kauffman
(MICH); Benton Harbor, near St. Joseph Riv., Sept. 1910, Lansing 2877 (F); o
Sept. 1838, Houghton (MICH); Prairie Rhonde edge of rich maple woods, Sept. 1
tows 287 (F); EN ac Co., Prentis Bay, Sept. 1916, Ehlers 22 (MICH) ; Menominee
Co., between Cedar Riv and. Stephenson, Aug. 1933, Grass] 3005 (MICH); Drayton
Plains, Aug. 1922, Farwell 6354 (MICH); St. Clair Co., near Port Huron, Aug. 1892,
Dodge, (MICH); Flowerfield, Burgess 424 (F); Ann Atbor; Aug. 1862, Almendinger
(MICH). wisconsin: Raukana, Aug. 1879, Schuette (F, upper leaves broadly amplexi-
e most westerly gëf noted for this variant); Dane Co., N. of Cross Plains, Sep
1916, Heddle 2667 (F); Trempealeau, Aug. 1927, Fassett 4496 (MICH). MINNESOTA:
Center City, Aug. 1892, Taylor (S). 10wa: Decatur Co., Aug. 1898, Fitzpatrick (F).
MISSOURI: St. Louis, 1832, Drummond (K, 2); Davis Spring, Aug. 1937, Moore
Shelby Co., N. W. of Emdem, Sept. 1948, dk '66595 (F); N. W. of Chillicothe,
Aug. 1951, Sparling 1325B. (F). DELAWARE: dune region between Cape Henlopen and
Rehoboth go. Snow 72, 192 (F). MARYLAND: Relay, Aug. 1910, Jones (F). DIST. OF
COLUMBIA: near Georgetown, Oct. 1858, Schott (F); Washington, Sept. 1897, Pieters
(MICH, dd plant 8.5 cm. tall, one capitulum). WEST VIRGINIA: Smyth Co., Pine
Glade Mtn., Aug. 1892, Small (F). NORTH CAROLINA: Statesville, Hyams (MICH); Hot
Springs, Aug. 1924, Webmeyer 694 (MICH, 2). KENTUCKY: Bell Co., near Wasioto, Sept.
1893, Kearney 502 (F); Meade Co., Otter Creek, Sept. 1950, Fen (S); about 2 mi. west
of om on weathered shale in highwiy cut, Sept. 1952, Belcher 1118, 1118A; roadside
nark 7 mi. E. of Vanceburg, by edge of woods, Sept. 1952. Belcher 1119 (sisde form.
leaves ptr ARKANSAS: Bradley Co., Jersey, “bottom thickets, 8 ft. high”, Sept. 1938,
‘Demaree 18313 (F); de banks of Saline River, Sept. 1938, Demaree 18388 (F).
TEXAS: Harrisburg, Sept. 1875. Joor (MICH); Hocklev, 1890, Thurow (F). FLORIDA:
i u
); Ft. Myers, 1900, Hitchcock 142 (F); Clarcona, Nov. 1899, Meislabn 126 (US);
Palm Beach, 1896, Hitchcock 925 e
West INDrEs. cusa: Havana, Guanabacoa, May 1914, Ekman 618 (S); Oriente,
Bayate. Nov. 1914, Ekman 3403 (5); Sierra Maestra, above oer Oct. 1916, sar
8063 (S). jamaica: St. Louis, roadsides, 1850, Prior (K); Glasgow, near Tro
1917, Harris 12637 (F). par: Dept. du Nord, cultivated field north of St. Michel, Dec.
1925, Leonard 7741 (F); Massif pP Nord, May 1927, Ekman H8305 (S). DOMINICAN
REPUBLIC: Santo Domingo City, at Rio Ozama, Jan. 1929, Ekman H-11375 (S). PUERTO
RICO: waste ground, Martin Pena, March 1924, Dale (MICH); Cayey, i in ee Llano,
Sept. m Sintenis 2412 (F, left and center specimens only, right is Sonc.
EuRoPr. CZECHOSLOVAKIA: us prope urbem Pferov, Aug. DE rinde. FI.
Exsic. Reipub. Bobem. Slov. 951 (S, US); Olmüvzl, Pferov, Sept. 1938, Laus (K).
AUSTRIA: prope Purkersdorf, Sept. 1909, Keller 5102 (S); Sóchau, Waldblóssen, Aug.
[Vol. 43
16 ANNALS OF THE MISSOURI BOTANICAL GARDEN
1909, Sabransky (S), Waldrodungen, Sabransky (US); Wurmberg prope Pettau, Sept.
1915, Maly (K, 2). HUNGARY: "ese Arad. in silvis caeduis montis Piliske, Aug. 1916,
Kiimmerle ES Jávorka, Fl. Hun . Exsic. 588 (F, K, S, US); Reposmere, Sept. 1922,
e" :
silvis caediis ad Zag ormastiny, Fl. Exsic. Austro-Hung. 658 (K). RUMANIA:
ransylvania, ad Gürscuysentinire, ie 1913, Heuzyel (S).
AWAIHAN ISLANDS. KAU ar Hanalei, Sept. 1922, Degener 1521 (NY). o
Honolulu, * de Ber DR 1916, Hitchcock 13716 (US); Honolulu, Tantalus
Mountain, June 1923, Degener 1520 (NY). Morokar: Tukoo, Oct. 1916, Hitchcock
15034 (US). west MOLOKAI: Hauakea, Pali, arid plain, April 1928, Degener 18134 (NY).
HAWAII: Flow of 1840, Puwa [?], July 1915, Pots 1055-H (NY).
Fernald concluded that “Erechtites hieracifolia is a very polymorphous plant
and that much tropical American material referred to it belongs clearly to some
other species. In the eastern United States and Canada the plant, although appar-
ently all of one species, is very variable.” He defined Senecio bieracifolius L. as
having auriculate-based clasping leaves, on the basis of Hermann's figure and the
diagnosis in the ‘Species Plantarum’; and Erechtites praealta Raf. as having all
leaves basally narrowed, on the basis of Rafinesque's description. He then stated
that these two plants are "in their involucres and achenes clearly extremes of one
species, but so different in foliage that they should be designated as forms or vari-
eties”. Fernald next stated that the most common variation appeared to be neither
of the above, but one “with the upper leaves sessile and broad at the base but very
rapidly decreasing to small bracts below the inflorescence", which he described as a
new variety, infermedia. From an examination of herbarium material he concluded
that "the three plants seem to have somewhat different ranges and they are there-
fore here treated as geographic varieties"; namely, var. fypica, var. intermedia
Fern., and var. praealta (Raf.) Fern. This treatment he maintained without mod-
ification in the 8th edition of ‘Gray’s Manual.’
Fernald's paper afforded an excellent starting point for my investigations, but
I have to disagree with his conclusions. The tropical material is not specifically
distinct from that of the temperate zone. His new variety intermedia has previ-
ously been described as a species. The three states he named do not in fact have
clearly distinct geographical ranges, but may exist side by side in the field and may
occur in the progeny of a single plant during greenhouse culture. Finally, he
formed a very inadequate concept of E. prealta.
The earliest name applied to the “intermediate” state is Erechtites prealta Raf.!
As there is no specimen surviving, it must be typified by Robin's description. This
is of a discoid Composite some 6 or 7 feet tall; with sessile leaves 1 foot long and 4
inches wide, oblong, attenuated toward the base, irregularly toothed, acute, glab-
rous with the nerves underneath covered with down; the involucre "monophyllous"
7 to 8 lines long, 3 to 4 lines in diameter, fleshy, "glanduleuse", striate, terminated
by a score of little teeth; florets white, 5-toothed; growing in the open near woods
and flowering in September.
This description undoubtedly E to an Erechtites, despite omission of several
details. It also obviously refers to a very robust specimen. The description of the
1956]
BELCHER—REVISION OF GENUS ERECHTITES I
leaf is hardly definitive. The measurements of size certainly do not apply to all of
the leaves indiscriminately, but surely only to the lower ones. It then is reasonable
to assume that the phrases which immediately follow these measurements; namely,
"oblongues, plus rétrécies à la partie inférieure", also apply to these same large
lower leaves and therefore not necessarily to those higher up which are of course
smaller. It is a general feature of every complete dried plant of Erechtites which I
have seen, and of all those which I have grown from seed or seen in the field, that
the lowest leaves are basally narrowed, even to the extent of being briefly petiolate.
This intimate association of the description of the size and of the base of the
leaves was, however, severed by Rafinesque, who also modified the description in
other unfortunate ways. Thus Fernald, apparently without referring to Robin's
original description, construed Rafinesque's "foliis . . . basi attenuatis" as applying
to all the leaves and particularly to the upper ones. He therefore placed the taxon
praealta in varietal status, containing the plants which do have all the leaves atten-
uated; that is, those depauperate specimens called f. minor by Waisbecker. Although
Fernald cited no specimens for his var. praealta, all of the sheets distributed from
the Gray Herbarium under this name which I have seen are of this depauperate
state, scarcely one foot tall, so much at variance with Robin's description and so
contradictory of the epithet.
In several hundred sheets of E. bieracifolia var. bieracifolia from a dozen major
herbaria, I have seen virtually every conceivable degree of intergradation among the
extremes recognized by Fernald. And, like Deam,”* I have seen and collected not
two but all three of these extremes plus other intermediate states in the same loca-
tion, growing virtually side by side, with the variation in leaf form correlated with
over-all size and that apparently directly related to soil fertility and moisture
supply.
This field experience has been substantiated by four years of growing these
plants in the greenhouse under varying conditions of moisture, fertility, and photo-
period. These studies, still incomplete, seem to indicate that the lower nodes always
produce leaves with attenuate, even subpetiolate, bases; that expansion of the basal
region of the leaf begins rather abruptly about the 30th to 35th node, and reaches
the broadly amplexicaul condition only if the sudden bolting which marks the
transition from the vegetative to the reproductive phase is delayed beyond the
maturation of this region. If vegetative growth is retarded by limiting either
water or nutrients, or terminated by the onset of short days, floral initiation ap-
pears before leaf-base expansion can occur. By varying the degree of stress applied,
sister plants grown from achenes taken from a single capitulum of a plant with
broadly amplexicaul bracts have been made to yield a spectrum of variants ranging
from minute specimens only 6 cm. high and bearing four or five subpetiolate leaves
and a single tiny capitulum to specimens over 2 m. tall and bearing hundreds o
21 Fl. Indiana, p. 994. 1940.
[Vol. 43
18 ANNALS OF THE MISSOURI BOTANICAL GARDEN
large capitula, the lower ones on extensively ramified branches subtended by large
broadly amplexicaul bracts.
The "intermedia" condition arises readily in summer plants restricted in access
to soil nutrients by being retained in 3-inch pots without additional fertilizer but
well supplied with water. Such plants commonly show signs of nitrogen starvation
in the lowest leaves, and fail in various degrees to develop complete tops. Their
growth is usually good at first but later it slows down so that the critical day
length is reached, and bolting, sharply marked by a maximal internodal length,
occurs just after leaf-base expansion has begun. Once bolting starts, the expansion
of leaves at the superior nodes is severely and progressively retarded, resulting in the
familiar series of bracts and bracteoles. Furthermore, in plants undernourished at
and after bolting, the development of floriferous axillary branches is greatly in-
hibited and may not descend below the 40th node, whereas in well-nourished plants
such branching may extend to or even below the 20th node. This results in the
condition described by Fernald, of the "upper leaves sessile and broad at the base
but very rapidly decreasing to small bracts below the inflorescence". A more
accurate statement would be that the lower part of the inflorescence fails to de-
velop properly, for minute capitula can usually be discerned in the axils of these
supposedly reduced leaves, which are indeed bracts.
The common name of var. hieracifolia is well explained by Pursh??: “This is
one of the plants which spring up in the most remote western counties when the
land is cleared of timber, particularly when the brushwood is burnt on the ground;
from which circumstance it is generally known by the name of fireweed." The
logic of the name "butterweed", by which this plant was well known to me as a
boy in the hills of Kentucky, has eluded me.
1b. ERECHTITES HIERACIFOLIA var. MEGALOCARPA (Fern.) Cronq. Rhodora 48:
122. 1946.
Erecbtites megalocarpa Fern. Rhodora 19:24. 1917.
eaves large, 10 to 15 cm. long, 3 to 7 cm. wide, fleshy; capitula 1 cm. or more
in diameter (pressed) and to 2 cm. long including pappus; phyllaries 16 to 21 or
more, lanceolate, 13 to 15 mm. long; florets 100 or more, corolla lobes with brown
margins and nerves; achenes (3.5 to) 4 to 5 mm. long, up to 1 mm. in diameter
(i. e., twice the diameter of the achene of var. hieracifolia), subturbinate, 16- to
20-costate.
TYPE: Upper and middle regions of sandy sea-beach, West Yarmouth, Mass.,
Oct. 8, 1916, Fernald, Butters & St. John 15468 (G).
Cosst of southern New England, where it occurs along the beaches and in salt
marshes.
— ade Groton, seashore, Sept. 1927, Janssen (S); Oct. 1927, Janssen (S).
ETTS: Yarmouth, sandy beach of Nantucket Sound, Oct. 14, 1916, Fernald es
[ras E [nis Gray 299 (F, K, MICH, S, W; paratypes) ; x salt marsh Sion Dess of Bass
River Light, Dennis, Sept. 1918, Fernald S Long 17636 (F, K, M 2); Bourne, inner
edge of Mona Bod; Sept. 1929, Blake 10975 (F, K, S); Warham, Blake 1 0962 (K).
22 Fl. Amer. Sept. 2:529. 1814.
1956]
BELCHER—REVISION OF GENUS ERECHTITES 19
In reducing Fernald's species to varietal rank, Cronquist stated that he was
unable to see any real specific discontinuity, with which I agree, and that he re-
garded it as a "saline marsh ecotype”. Fernald apparently was not convinced, since
he continued to treat it as a species. It may well be that this large-headed state is
a polyploid, perhaps a simple tetraploid, of var. bieracifolia.
Most of the specimens of var. megalocarpa which I have seen strongly resemble
the broad-bracted state of var. hieracifolia'but have the bracts or uppermost leaves
just below the inflorescence very broad at the base, up to as much as 5 cm. wide, It
is therefore of particular interest that one of the specimens examined from Con-
necticut, Janssen, Oct. 3, 1927, shows a distinct departure from this pattern, even
as ase: by Janssen, Sept. 3, 1927. In the former, the leaves immediately
low the inflorescence, instead of being very broadly semiamplexicaul and large,
are much reduced in all dimensions, with the semiamplexicaul base only 1 cm. wide,
and the medial leaves are basally attenuated instead of being broadly clasping.
Although the specimen lacks the basal portion, it seems probable from the reduc-
tion in the degree of branching and in the size of the branches that this October
gathering was from a plant less robust than the others. This would accord well
with the thesis that foliar form is largely under environmental control, not only in
var. hieracifolia but in var. megalocarpa as well. It also reinforces the interpreta-
tion of this taxon as being of varietal rather than of specific rank.
lc, ERECHTITES HIERACIFOLIA var. CACALIOIDES (Fisch. ex Spreng.) Griseb. emend.
Belcher.
M ee bi Ce deng P DC. var. ben zé Sieg? ex Spreng.) Griseb. Fl. Brit. W.
61 cies incorrectly attributed t . and var. to Less.), and var.
se ik. (Cas) eech be cir ad to DC.).
Senecio Juve opus . Mant. Pl. 469. 1771, Type: P. Browne (LINN 996-2), non Sp.
Pl. 886. 1753; Willd. Sp. Pl. ed. 4. 3:1974. 1800, pro parte ;
Senecio cacaliides Fisch. ex Spreng. ëch Prov. 37. 1819; Syst. Veg. 3:565. 1826; Link,
num. Pl. Hort. Berol. 2:325. 182
oncbus agrestis Sw. Prodr. 110. cem excl. syn.; Fl. me Occ. 3: Eo 1806, excl. syn.;
Willd. Sp. Pl. ed. 4. 3:1513. 1800; Spreng. Syst. Veg. 3:648.
Neoceis carduifolia Cass. Bull. Sci. Sot. Philom. Paris 1820:91. E Dict. Sci. Nat.
34:386.
Soncbus etd ii Spreng, Neue Entdeck. 2:143. 1821; Syst. Veg. 3:648. 1826.. Type:
Sprgl. Herb. n. 1985
Senecio Ger Ce GER Desf. Cat. Hort. Paris. ed. 5. 177. 1829 (not 1819, as cited by
de Cando
i ee KSE (Fisch. ex Spreng.) pe Syn. omg Comp. 395. 1832; DC. Prodr.
ue
d
6:295. 1838 (as lie atte as to descr., not specim
Brechiites Ar (Cass.) DC. Prodr. 6: 294. 1838; "Benth. Vidensk. Meddel. Natur-
Forening Kjoben. 1 t 1852; Walp. Ann. Bot. Syst. 5:290.
Erechtites Dame Walp. Rep. Bot. Syst. 2:651. 1843; Cabr. Rev. Mus. La Plata 4: 286,
: 90. 1941; non DC.
TAM a eren & Walp. Nov. Act. Acad. Caes. Leop. Carol. Nat. Cur. 19,
ppl. 1:2
Senec heben Sch. Bi >. Flora 28:498. 184
lor roni esp Gar de. L Lond. Jour. gue 3 419. 1848. Type: Goias, Gardner 3868 an
qs Dit ee L.) DC. var. cacalioides Less. f. pubescens O. Ktze. Rev. Gen. P
91, nom. qi Of 4 co os cited, one at least, “Java, Sindanglaja, pro
4488 (NY), is typical var. cacalioides.
[Vol. 43
20 ANNALS OF THE MISSOURI BOTANICAL GARDEN
— carduifolia DC. var. latifolia Klatt, Bull. Soc. Roy. Bot. Belg. 36:291. 1896,
Cb ee Rica, San Rafaél, Pitt. 6968, not seen.
Fe Awe at eylan var. malasica Ridl. Jour. Str. Br. Roy. Asiat. Soc. 61:24
Gynura erte dn ei? Fl. Malay Penins. 2:190. 1923, excl. Griffith's specimens
from Malacca, which are true Gynura.
Gynura aspera Ridl. Jour. Male Br. Roy. Asiat. Soc. 1: Ze 1923. Type: Sumatra, Beras-
i, Feb. 8 ("dwarf form"), Feb. 10, 1921, Ridley (K
a agrestis (Sw.) Standl. & Steyerm. Field Mus. eu Bot. 23:265. 1947; Rydberg,
n herb.
Bracteoles of the calyculus long, extending to one-third to one-half the length
of the involucre, typically ciliolate with coarse multicellular hairs; phyllaries 12 to
14 (to 21), glabrous or sparsely and irregularly beset with multicellular hairs.
Holotype (?): "Senecio cacalioides, Fischer, 1818, Sprgl.!, herb. n. 1687, Syst.
III, 565, n. 176." (P!
Very variable in habit and leaf form, showing much the same series of varia-
tions as in var. bieracifolia, with the variations intergrading in every degree. A
characteristic state in Florida is intermediate between var. cacalioides and var. hier-
acifolia in that the bracteoles are long but lack multicellular hairs. Especially
variable in Brazil, one particularly interesting state from Paraná having a distinct
basal rosette of leaves.
MExICO. TAMAULIPAS: Tampico de ee Bee k^ Berlandier (FI, 2); vicin-
ity of Tampico, Jan. 1910, Palmer 107
34
1899, Pringle 8187 (BM ; 9) born, Apri 0, Orcutt 3018 (F); vicinity of
ruz, W (W). Jalisco: “Jalisco, 1886”, eque (BM); Quimixto, Nov.
es pe 1193 (BM, F IAP. Mt. Ovando, is seh oo 1 DS
26, ,
Matuda 16169 (F); Mandolopez, Des 1947, Matuda 1 6647 (F). TABASCO: Boc
ber July 1939, Matuda 35 MICH). cAMPECHE: Palizada, July 1939, ‘Matude
I SH YUCATAN: pem Vista, Xbac, Gaumer 1438 (F, S); Pocobach,
Cae 2394 deg F).
G : La Libertad & bizie 1933, Aguilar 207 (MICH); Lake
Zotz, May 1933, age sis (MICH). a AZ: Coban, Jan. 1908, von Türck-
beim 1I 1319 (BM, CH, S); dus Lee N. È of Carchá, April 1939, Standley
(F Soso near Amatitlán, Dec. 8, Standley 61433 (F). IZABAL:
Livingston, sandy beach, Feb. 1905, Duos 244 (M ICH). SE ATEP
ate, N. of Antigua, Feb. 1939, Standley 64698 (F). sucHiTEPÉQUEZ:
March 1928, Morales 1039 (F). EscuINTLA: S. R
89624 (F). saNTA ROSA: Volca eg ecd 8000 ft., Jan. 1893, Heyde & Lux 4227
(F), 6000 ft., Dec. 1892, ibid 4247 (K, 2). JALAPA: betwe
Nov. 1940, Standley 76933 (F). zacapa: Sierra de las Minas, Oct. 1939, ves tip
29638, 29719 (F). D ue Lago Retana, between Ovejero and Progreso, Nov.
Hoops: Morazá
e a
4 e wur valley near Las Mesas, Feb. 1947, Williams t$ Molina 12053 (F); hills
above Jicarito, Nov. 1950, ege 17293 (F). El Cayo: Mountain Pine ridge, Marc
1931, Bartlett 11917 (MICH ánti i
(FR: S
1956]
BELCHER—REVISION OF GENUS ERECHTITES 21
1887, Cooper en (K, F); Ferme de Frec: July 1901, Soe 14744 (BM); Galieros,
CUP var Aug. 1 (ee a K-154 (K); Ton dr San Ramón, Nov. 1927, Brener
5806 (F); Tablazo, Tu LE Valerio 1090 Te Prov. San José, vicinity El Géneral, July
1936, Skutch 2706 (K, M CH, Sy; EE de e 4500 ft., Oct. 20, 1937, “herb
i a with erect branches se 16 [sic!] Misco AI i
clesred of forest", Austin Smitb A-557 7 (P). PAN MA: Empire station, Nov. 1861, Hayes
587 (BM, K); "in ruderalis prope urb. Panama", prd 444 (BM, K, P); Canal Zone,
jue — Island, July 1931, Starry 283 (F).
DIES. BAHAMAS: Red B we Andros, April goe Northrup (F). cusa: Her-
ki
—
SS
3
Br
EC
D
D
-
D
D
GEN
EI
zi
3
e
<
eso eg 1906, Hitchcock (F); Camaguey, Tiffin, March 1909, Schafer 1090 (F);
Havana, Puentes Grandes, April 1914, Ekman 483 (S); Ces eise ad Rio Jagua, May
1919, Ekman 9610 (F, S), Sierra Maestra supra Daiquiré, Oct. 6, Ekman 8177 (P E
SLE OF PINES: Pedernales Point, Feb. 1889, Millspaugh 1418 B). JAM AICA:
(LINN 996-2); “Sonchus agrestis”, Swartz (S); Castleton Hill, July 1900, Paweh.. 8024
(F); Castleton, April 1910, Harris 10890 (F, K). Harti: Dept. du Sud, prope Civette,
June 1917, Ekman H-221 (S); Massif de la Geier pra des Bouquets, Terrelonge, March
St
27; Hess H-7810 (S). DOMINICAN RE to. Domingo City, ad Rio Ozama,
Jan. 1929, Ekman ues (S, very large leaves ess had eer bracts); vicinity of
Santiago, near La Cumbre, Jan. 1946, Allard 14587 (S); Voitea u (FI). PUERTO RICO:
Wydla, 1825,— ae Mayaguez, Oct. 1881, Sintenis 164 (K); Cayey, in monte Llano, Sept.
1885, Sintenis 2412 (F); prope Bayamon, Pob: se e Sein 865 (S). sr. THOMAs: Riedlé
(FI); St. Peter, Jan. 1881, Eggers 209 (K). rt. i “Spray garden” , July 1896,
Rickseckler 474 (F). GUADELOUPE: “E. hieracifolia Rafin. Guadaloupe” (K). GRENADA:
Ve ree St. George's, May 1905, Broadway (F). towaco: April 1913, Broadway 4463
(S); on Hill near the river, May 1913, Broadway 4507 (F); Caroni swamp, July 1931,
Willems "cres
Sou RICA. FRENCH GUIANA: Acarouany, 1854, Lagotz (S). DUTCH GUIANA
ER Dee Sipolivini R., March 1936, Rambouts 544 (K, 2). BRITISH GUIANA:
Hyde Park, Demerara, May 1922, “
ren (F);
1 ). VENEZUELA: Montevideo, C s, Ma E d (F); Bolivar,
ran Sabana, Oct. 1944, Steyermark oe CoLomBiA: Magdalena, near Lake
Zapatoza, Aug. 1924, Allen 277 (K); > Masta, Smith 666 (F, ei Ch ya de
Togoromá, June 1944, Killip © Catia 39075 (F). Valle: Rio Colima, La Trogita,
1944, Cuatrecasas 16502 (F). Cauca: la Paila, s Helton 384 (K); Popayán, Lehmann
5224 (K), Lebmann 7598 (F, K), Lebmann 7985 (F); ad pag. El Tambo, June 1938,
von Sneidern 1535 (S). Tolima: Ibagué André T 262 (K). Meta: Villavicencio, Dec.
1928, Apolinar-Maria 427 (F). Vaupés: Bocas del Carurú en Casa Alvarez, Sept. 1939,
Cuátrecasós 7009 (F). Peru. Loreto: Lower Rio Nanay, May 1929, Williams 282 (F);
Caballo-Cocha on Amazon River, Aug. 1929, Williams 2489 (F). Ambo, 7000 ft., April
1923, Macbride 3158 (F). Amazonas: Chacapayas, Matthews (K). Piedra Grande:
estación near Rio Santo Domingo, Macbride 3673 (F). Junin: Satipo, A
2841 LEL BoLivia. Reyes: Rurrenabaque am Rio Beni, 1930, Fleischmann 147 (S). La
Paz: Apolo, April 1902, Williams 182 aa "Nord-Yungas", Milluguaya, Dec. 1917,
prn ons 4072 (F). JU MM Sireapaya bis Yoinacachi", Dec. 1906, Bucbtien (W).
anta Cruz: Buena Vista, July 1924, Steinbach 6208 (K). Larecipa: t
Canale Challapampa riv.", 1860, Mandon 117 (K, det. as E. ambigua DC.; S). Casana im
Tipuani-Tal, Sept. 1922, Buchtien 7589 Gë Brazit. Amazonas: Taperinha bei San-
tarem, July 1927, Ginzberger 412 (F). Para: Béa Vista on Tapajós River, 1929, Dahlgren
& Sella 137 (F); Belém, Instituto Agronomica do Norte, Oct. 1942, Blake 7755 (K).
Pernambuco: Tapera, Sept. 1931, Pickel (F, MICH). Bahia: “Bahia in convallibus
humidis” (W). Goias: near Villa de Arrayas, Gardner 3868 (K, 2, syntypes of E. pores
Gardn.). Minas Gerais: Caldas, 1845, Widgren 211 (S, 2); Lagoa Santa, Nov. 18
Warming (S, 2). Rio de Janeiro: in locis humidis, Dec. 1831, Riedel 230 (P); ee
between Alto de Serra and Meio de Serra, Dec. 1928, Smith 1551 (S). Sao Paulo: Camp-
inas, March 1900, Novaes 134 (US). Paraná: Curityba, Feb. 1904, Dusén (S); Ponta
E
os
—
o
>
M
$
a
R
>
[Vol. 43
22 ANNALS OF THE MISSOURI BOTANICAL GARDEN
Grossa, March 1904, Dusén (S); lacarehy, Sept. 1908, Dusén 6583 (S); Jaguariahyva,
June 1914, Dusén 15110 (S). Rio Grande do Sul: pr. Sio Martinho, Apr. 1893, Malme
826 G (S); Ijuhy, April 1893, Malme 746 (S); Sao Leopoldo, 1941, Eugenio 63 (F).
PARAGUAY: Chaco-y pr. Concepción, Hassler 7267 (S); in viciniis Caaguazú, Feb. 1905,
Hassler 8886 (K, S); "Villa River," Jörgensen 3512 (F, S); Hassler 1405 (K); "in Para-
guay legit Fleischer” (P). ARGENTINA. Jujuy: Quinta, prope Laguna de la Brea, June
1901, Fries 100 (S) ; Rio Grande, Feb. 1924, Venturi 3418 (US). Tucuman: April 1928,
Venturi 6095 (F). Sierra de Cordoba: March 1876, Hieronymus 499 (F). Misiones:
osadas, prope "La Granja", Nov. 1907, Ekman 1106 (F, S); Apostales, San Jose, Feb.
946, Bertoni 2722 (F). Urucuay: Dept. de Colonia, Riachuclo, Charca, April 1935,
Cabrera 3319 (S, several plants; most southerly specimens seen).
: INA: Kwangsi, Shang-sze Dist., Shap Man Taai Shan, May 1933, Tsang
22219 (BM, GH, S). Siam (all det. as Gynura malasica Ridl.); Surat, Panam, March
1927, Kerr 12413 (K); Kurabi, Kao Sataw, Kerr 12424 (K); Pattoni, Kao Kalakino, March
1928, Kerr 14868 (K); Takinapa, Kapang, Feb. 1929, Kerr 17551 (K); Kurabi, Tambur
Kao Panom, April 1930, Kerr 18830 (K). MALAYAN FEDERATION (all det. as G. malasi ):
Selangor, Kuala Lumpur, Damansara i
Tanga Pass, Ridley (K); Johore, Sungai Tukong, July 1930, Spare F802 (K). BRITISH
BonNro: Myburg Prov., Sandakan, 1921, Elmer 20159 (K, det. as Soncbus arv-
7395 (GH, US); Island of Siberoet, Sept. 1924, Boden-Kloss 14556 (K). Java: Sin-
danglaja, May 1875; Kuntze 4488
» 2); Gedeh, Tjibodas, Feb. 1897, Móller (S, right-hand specimen only; left is E.
valerianaefolia).
Grisebach was apparently the first to propose combining all of the Caribbean
material under E. hieracifolia, and to use differences in bracteole length to distin-
guish varieties. By considering length and width of phyllaries, length and number
of calycular bracteoles, and general pubescence, he established three varieties, in-
cluding the typical one. Unfortunately, the number of bracteoles, upon which he
based the tropical varieties, is not consistent, even from capitulum to capitulum on
the same inflorescence, even on the very specimens he cited. The description of
var. carduifolia as glabrescent is not generally applicable, the description as pubes-
cent or as setaceous, given by Cassini, Desfontaines, and de Candolle, being much
more appropriate.
Despite the artificiality of his varietal limits, the epithets which Grisebach used
for his varieties have clear priority in that rank, and at least one must be main-
tained. Since I could not locate any authenticated material of Neoceis carduifolia,
I have taken up cacalioides, The epithet agrestis, although clearly the first unpre-
occupied name to be applied to the tropical material, was used only in specific rank,
where it has priority for those that might wish to continue to separate the temperate
and the tropical states into different species.
Swartz described Sonchus agrestis from Jamaica.. There is no type in the strict
sense, for Dr. Asplund informs me that there are two specimens of E. hieracifolia
at Stockholm which were collected by Swartz in the West Indies but that the prin-
1956]
BELCHER-—REVISION OF GENUS ERECHTITES 23
cipal annotations are by J. E. Wikstróm. I have examined both of these sheets,
together with a third Swartz specimen labelled by Banks as Sonchus agrestis and
now in the herbarium of the British Museum (Natural History). Iat first believed
that Swartz applied Sonchus agrestis to the smaller of the two specimens said to
have been collected by him. This specimen, closely resembled by the one in London,
has the phyllaries glabrous but only 7 or 8 in number, the bracteoles of the caly-
culus only 2 mm. long, and the leaves attenuate and non-amplexicaul at the base.
These lesser specimens, however, do not have tomentose, or even sub-tomentose,
branches, nor incised-serrate leaves, nor many capitula, as described by Swartz.
Particularly because of their glabrous phyllaries and short bracteoles these lesser
specimens must be the typical variety, with the more depauperate specimens of
which they agree well. Although Swartz may have included some features of these
lesser specimens in the amplified description in his later work, the name should be
interpreted by the earlier diagnosis, which applies better to the larger specimen with
the multicellular hairs and other features of var cacalioides.
Fischer in 1818 at Halle raised from Jamaican seed some plants to which he
evidently gave the name of Senecio cacalioides. He appears to have sent a named
specimen to Sprengel at Berlin, who gave it an excellent description in his account
of the new introductions at Berlin and at Halle for that year. Although it is not
clear if the specimen of "Senecio cacalioides Fischer, 1818” in Sprengel’s herbarium
is the one actually grown by Fischer, it agrees in every respect with Sprengel’s de-
scription, particularly in being distinctly pubescent with multicellular hairs.
e fact that the Compositae of Sprengel’s herbarium are in the Paris Herbarium
seems not generally known, but is of considerable importance to the synantherol-
ogist, particularly in the interpretation of the “hort. berol.” synonyms used by de
Candolle now that the Compositae in the general herbarium at Berlin are destroyed.
When Sprengel’s collection was sold, the Compositae were bought by Schultz Bi-
pontinus, whose collection of this family alone was said by Alphonse de Candolle??
to have numbered 50,000 specimens. His collection, containing much type mate-
rial, was in turn acquired by Cosson, and this eventually (after publication of “La
Phytographie") by the Muséum d'Histoire Naturelle. Sprengel's sheets are iden-
tified by uniform tickets reading: "Sprgl.! herb. n.
Syst. III, [page n. ], n. ". I believe that these tickets are in the
writing of Schultz, rather than that of Sprengel. This reduces their claim to
authenticity, and I hesitate to cite these specimens as holotypes for names that
originated with Sprengel, particularly those from publications which preceded the
"Systema Vegetabilum". But for those specimens which I have examined the col-
lation to page number, species number, and description in the "Systema Veg-
etabilum” is excellent. Link also attributed Senecio cacalioides to Fischer, but
based his own description on a similar plant from Brazil. De Candolle later re-
duced Link's name to Erechtites ambigua, but wrongly, the latter actually being
the petiolate form of E. valerianaefolia.
23 La Phytographie. p. 450. 1880.
[Vol. 43
24 ANNALS OF THE MISSOURI BOTANICAL GARDEN
Sprengel first described Sonchus occidentalis as similar to S. agrestis Sw., but
with glabrous peduncles and leaves. In the "Systema Vegetabilum" this was revised
to very glabrous, in greater contrast with S. agrestis. Some authors have assumed
that these descriptions therefore apply to those virtually glabrous specimens of var.
bieracifolia which occur scattered throughout the West Indies. But the confusion
surrounding this epithet has been removed by discovery of Sprengel's type, or at
least an authentic specimen, in the Paris Herbarium. This specimen is not as glab-
rous as the descriptions indicate. With magnification the scattered stumps of
multicellular hairs can be seen on the lower surfaces of the leaves and on the stem.
The capitula have long calycular bracteoles which are ciliolate with multicellular
hairs. This material of S. occidentalis thus belongs to the tropical variety. Rydberg
apparently made a new combination in Erechtites for this species, for Degener has
used this name on tickets on certain of his collections of Erechtites from Hawaii,
attributing the combination to "Rydberg in herb". On comparing these and other
Hawaiian sheets with Sprengel's specimen, I find that the Hawaiian material uni-
formly has glabrous phyllaries, very short calycular bracteoles, peduncular brac-
teoles less than one-half the length of the involucre, and no multicellular pub-
escence on either type of bracteole. These characters exclude it from E. hieracifolia
var. cacalioides, and place it in var. hieracifolia instead.
Erecbtites bieracifolia is represented in the Orient, however, by var. cacalioides
only, and appears to be neither widely distributed nor abundant there. Most of the
specimens from this region which are so determined have proved actually to be
the African Crassocepbalum crepidioides (Benth.) S. Moore. The two are readily
distinguishable, the latter by the longer style-arm appendages, small dark-red
achene, lyrately pinnatifid and petiolate lower leaf, and usually lacking pistillate
marginal florets. Merrill ?* fell victim to this confusion when he wrote: “Two
species of the American Erechtites, both with pink flowers, may be dominant wher-
ever they have been introduced, E. hieracifolia, fig 185A, and E. valerianaefolia,
fig. 185B." True E. hieracifolia never has pink florets, whereas C. crepidioides does,
as numerous specimens of it incorrectly determined by Merrill as E. bieracifolia
testify. As for the figures cited, fig. 185A is certainly E. valerianaefolia (instead of
E. bieracifolia as stated), with its characteristic pinnatisect leaf, whereas fig. 185B
is (C. crepidioides but shows only the subentire foliar leaves. Merrill Ze of course
by no means alone in this misapprehension, as it is evident in nearly all of the her-
baria which I have studied.
Grisebach reported E. hieracifolia var. bieracifolia from Mauritius, based, I
believe, on two specimens at Kew. One is ticketed: “Senecio? fl. roseo. Growing
in high mountains, Mauritius. Senecio cacalioides Bojer. H. M. 188, Erecbtbites
bieracifolia Raf.". This proved to be Crassocepbalum rubens (Jacq.) S. Moore. The
other specimen, simply labeled “Telferin. Mauritius", and determined as E. bieraci-
folia, was immature and poorly pressed, but appeared to be Crassocephalum sarco-
24 Plant Life Pacific World, p. 143. 1945.
1956]
BELCHER—REVISION OF GENUS ERECHTITES 25
basis (Bojer) S. Moore, vel aff. I have yet to see Erecbtites from Mauritius,
Madagascar, or vicinity.
hese two major varieties of Erechtites bieracifolia are not absolutely distinct;
quite possibly they intergrade where their ranges overlap in the Caribbean basin.
Occasional specimens of tropical material have the calycular bracteoles rather
shorter than usual or have longer calycular bracteoles which are on first inspection
apparently devoid of multicellular pubescence. Occasional temperate-zone speci-
mens occur in which calycular bracteoles approach the length which characterizes
the tropical variety. Nature continues to defy our pigeon-holes.
Several subsidiary considerations, in my experience, may help establish or con-
firm the varietal determination in such cases. The width of the calycular bracteoles
is usually greater in var. cacalioides than in var. hieracifolia, to as much as twice as
wide, even in bracteoles of the same length. In determining the length of the
bracteoles the most reliable measurements can be obtained on capitula which have
fully elongated but have not yet begun to expand the floret buds, as during an-
thesis the bracteoles begin to flex. By fructescence they are quite often both
strongly flexed (and thus apparently shorter) and appreciably withered. The
lateral expansion of the receptacle which normally occurs during maturation of the
achenes also tends to distort the ratio between calyculus and involucre.
Care should be taken in examining younger parts of the inflorescence, to distin-
guish between members of the calyculus proper, whose insertions are always upon
the expanded torus, and the bracteoles of the peduncle and the bracts subtending
the peduncles, Either of the last two may, before the peduncle is fully elongated,
overlap the calyculus and appear to be a part of it. It is also true, though, that
both of these structures tend to be longer in var. cacalioides than in var. hieracifolia.
Indeed, length of these parts may be a helpful secondary characteristic if the pe-
duncles are fully elongated, although a less reliable one than length of the calyculus.
Short multicellular hairs may occur occasionally on the stems and leaves of var.
hieracifolia, and indeed do so on the Linnaean lectotype, and often occur abundantly
on the stem and leaves of var. megalocarpa. But I have not yet observed any
occurring on the inflorescence of any truly temperate-zone specimen, not even on
those Gulf Coast specimens which have the long bracteoles of the Caribbean vari-
ety. I therefore conclude that any specimen with even remnants of multicellular
hairs in the inflorescence is var. cacalioides.
2. ERECHTITES VALERIANAEFOLIA (Wolf) DC. Prodr. 6: 294. 1838.
Senecio valerianaefolius Wolf, Ind. Sem. Hort. Berol. 1825, teste Reichenb. Icon. Bot. Exot.
59, ta š
denaid herbaceous, subsimple to much branched above, glabrous or
occasionally sparsely hispidulous, striate, 0.5 to 1.0 (to 2.0 or more) m. high.
Lowest leaves petiolate, ovate-lanceolate to lanceolate, entire or serrate to irregularly
dentate; medial leaves petiolate with narrowly decurrent wings, very deeply pin-
nately lobed, the lobes lanceolate and serrate to irregularly incised-dentate, or
pinnatisect with linear segments entire or minutely serrulate, or entire or subentire
[Vol. 43
26 ANNALS OF THE MISSOURI BOTANICAL GARDEN
like the lower leaves; upper leaves similar to the medial leaves but slightly reduced
in size upward, or sometimes abruptly reduced several nodes below the inflorescence.
Inflorescences terminal and axillary, forming a rather congested cymose panicle.
Capitula slender, at anthesis about 10 mm. long, 3 mm. wide, scarcely ventricose,
with linear calycular bracteoles extending to one-fourth or one-third the height of
the involucre; involucre of 12 to 14 (to 16) phyllaries; phyllaries 7 to 8 mm.
long, 0.5 to 0.75 mm. wide, linear and acute to acuminate, with keel flat and 4- or
5-nerved, glabrous or rarely minutely hairy; marginal florets uniseriate or sub-
biseriate, corolla 5-fid, with lobes 0.5 mm. long and 0.2 mm. wide, apices glan-
dulose-thickened and incurved; style-arm apices shortly conic-appendaged. Disc
florets more numerous than the marginal, the outer ones transitional in size and
shape, with corolla 7 to 8 mm. long, only slightly longer and more dilated than the
pistillate florets, the inner ones with corolla slightly longer and larger, slender,
infundibuliform, 5-fid, with lobes 0.5 mm. long and 0.2 to 0.35 mm. wide, their
apices glandulose-thickened; style-arm apices with conical appendage approximately
0.05 to 0.1 mm. long. Achene cylindric, 2.5 to 3.5 mm. long, with about 10
heavy, pale brown ribs, dark brown and entirely glabrous to minutely villous or
hispidulous in the grooves. Pappus multiseriate, slender, rose-lilac to very pale
reddish, rarely nearly or quite faded to white, subequalling the florets, exceeding
the phyllaries.
Widespread in tropical America, where it sometimes hybridizes with E. hieraci-
folia var. cacalioides; adventive as an aggressive weed into tropical Asia, many of
the Pacific islands, and northern Australia.
Separable on the basis of differences in the foliage into four fairly well-marked
ms.
KEY TO THE FORMS
A. Medial and upper cauline leaves pinnatisect or subpinnate ...... B
B. Leaves only slightly reduced in size upward below the inflorescence
C. Segments of the strongly pinnatisect leaves lanceolate, broad, serrate to inci
cised-
dentate MAP Po de EC S DNI AMEND, EAA ZA f: alerianaef olia
CC. Segments of the pinnatisect leaves linear, entire or minutely serrulate........ . f. organensis
BB. Leaves abruptly and markedly reduced in size several nodes below the inflorescence
5 2c. f. reducta
AA. Medial and upper cauline leaves entire or subentire . 2d. f. prenantboides
2a. ERECHTITES VALERIANAEFOLIA f. VALERIANAEFOLIA
Senecio valerianaefolius Wolf, Ind. Sem. Hort. Berol. 1825, teste Reichenb. Icon. Bot. Exot.
59, tab. 85. 1827; Link ex Spreng. Syst. Veg. 5:565. 1826.
Senecio valerianaefolius Desf. Cat. Hort. Paris. ed. 3. 178, 403. 1829; by descr. florets all
perfect, a discoid Senecio, but type (FI ex Hb. Desf. !) has marginal florets pistillate.
ster ope valerianaefolium (Wolf) Less. Linnaea 5:163. 1830; Syn. Gen. Comp. 3
1832
Sonchus erythropappus Meyen & Walp. ex Walp. Nov. Act. Acad. Caes. Leop. Nat. Cur.
19, Suppl. 1: 293. 1843.
Senecio valerianaefolius Gardn. Lond. Jour. Bot. 4: 127. 1845.
Gynura rosea Ridl. Jour. Str. Br. Roy. Asiat. Soc. 61: 25. 1912.
Erechthites Gardneriana Cabr. Brittonia 7: 54. 1950.
Neotype: "Senecio valerianaefolius ex. h. Raffeliano, 1825” (W, “collectio
Reichenbach fil., aqu. 1889, no. 16256").
1956]
BELCHER—REVISION OF GENUS ERECHTITES 27
Medial cauline leaves 5 to 20 (to 30) cm. long, 2 to 8 (to 15) cm. wide,
strongly pinnatisect or subpinnate with narrowly winged rachis; segments 5 to 7 on
either side, broadly lanceolate, coarsely and irregularly serrate to incised-dentate,
sometimes basally sublobulate.
idely distributed from Central Mexico to Brazil and Argentina, rare in the
Lesser Antilles; adventive in Southeastern Asia, the East Indies, Philippine Islands,
New Guinea, Queensland, Fiji Islands, Samoan Islands, Hawaiian Islands, and else-
ish in the Pacific as an aggressive wee
TRAL AMERICA. Mexico: Vera Së recent clearings SE SSC Pringle 8334
vn es ,W); Chiapas, D Siltepec, 1938, Matuda 1971 (M t. Tacana, Aug.
1938, Matuda 2451 (MICH); MacDaniels 861 (e mS gu de m erapaz, Cobán,
Nov. 1907, von e I] 1396 (F, S). EL Satvapor: Ahuachapán, vicinity
Apaneca, Jan. 1947, Standley & Padilla 2928 (F); Cerlion de 3939 m HONDURAS:
Morazan, Standley & Molina 4138 (F). NicARAGUA: Summit of Mt. mbacho, near
Grenada, Grant 329 (Ps Jinotega, Grant 7303 (F); Tate 181 (485) sce Tao
10619 (F). Costa Ric San José, vicinity of El Géneral, July 1936, Skutch 2751
(MICH, S); San Loié, 1458: Scherzer (W). PANAMA: Allen 1 367 (F); Davidson 528
(F).
West INDIES. PUERTO RICO: headwaters, Inabón River, Jan. 1941, Otero M-111
(MICH). LEEWARD ISLANDS: Montserrat: e Mtn., Jan. 1907, Shafer 333 (F).
WINDWARD ISLANDS: Dominica: Hodge 777 (BM
H AMERICA. VENEZUELA: Vogl 475 (F); Moritz e (BM). CoLomsia: Santa
Marta, Smith 508, 664 (BM, F, S). ECUADOR: Rimba ch 277 (F). Peru: Aug. 1854, -
Lechler 2461 (W, d BoLivia: Hacienda Simaco sobre el camino a Tipuani, Feb. 1920,
Bucbtien 825 (BM, F S); Mapiri, April 1886, Rusby 1671 (BM, MICH); Bang 2068 (F,
MIC no ncbet (W); Bahia i
Agricultural College lands, March 1930, Ynes Mexia 4415 (BM, F, MICH, S). Rio de
Janeiro: “in montosis,” 1832, Lund (G Prodr.) ; ad aquas circa Rio de Janeiro, Pohl 646
(W); Organ Mts., open bushy places, 3000 ft., Gardner 522 (K, diseased, type of S.
pai pls srt Gardn.). Sao Paulo: Campinas, Oct. 1904, Heimer 228 (S). Paraná:
Passo, March 1904, Dusén (S); lacarechy, Aug. 1914, Dusén 15351 (F, S). Santa Cat-
arina: Mueller 454 (K). Rio Grande do Sul: Canóas pr. Pórto we A Nov. 1892,
veh 298 (S, 5) Ijuhy, April 1895, Malme 750 (S, 4); São Leopoldo, Oct. 1936, Dutra
1 (S). Paraguay: Villarrica, ees 7489: (F, S); San "end “Hassler 3625
(BM, W); "ad ripas Paraguay", Jan. 1873, Gilbert 1066 (K, stem flexuous) ; Hassler
11815 (BM); Fiebrig 634 (F). agoe, Misiones, Posadas, Nov. 1907, Ekman 1103
(F, S); S. José dos campos, Aug. 1909, Ds 299 (S).
od JAPAN: P eun Island, Mitsume-mura, Dec. 1948, Shigetake Suzuki,
eg: Japonicae 391088 (GH), Jan. 1949, ibid. (US). CHINA: Hainan, Sha po Shan,
Taai Shui ravine, Aug. 1927, Tsang 682 (NY, US); Id 1933, How 70622 (GH, NY,
US); S. W. Seven Fingers Mtn., April 1932, Liang 61669 (F, GH, NY); near Tau Ti P'o,
April 1922, McClure 9128 (NY); Tam dist., S. of F yg. May 1929 Tsang & Fung 223
(NY); Taam-Chau dist., Nodoa, July 1927, Mires 0 (NY, US). Maayan FEDERA-
TION: Pahang, Cinbéron Highlands, April 1 T ekle ion 32659 (GH). (Numerous
n
Gynura rosea Ridl., observed at Kew and the British Museum, by oversight were not
listed). SUMATRA: Karo Highlands, Berastagi, June 1928, Hamel 428 (GH, NY); vicin-
ity of Rantau Parapat, Bila, 1932, Rabmat Si Toroes 1716 (NY, US); Adian Rindang,
Asahan, 1935, Rabmat Si Boeea 8842 (GH). Java: “Java (iter javanum secundum)”,
Zollinger 2658 (G Deles., P); "ad rivulos pr. Gadok, 1600 ft., 12 Nov., 18—,” Zollinger
I 3655 (W, 2); eet eg May 1875, Kuntze 4407 (NY); Gedé, Mey 1875, Kuntze
4807 (NY); Preanger Prov., Tjiboeroem, forested middle slopes of Mt. Gedé, April 1909,
[Vol. 43
28 ANNALS OF THE MISSOURI BOTANICAL GARDEN
Palmer & Bryant 197 (US). Sarawak: without locality, native collector 727 (US).
BritisH Nom Borneo: Mt. Kinabalu, Penataran River, July 1923, Clemens 34047
(GH). REPUBLIC OF THE PHILIPPINES: Luzon: Sorsogon, Irosin, Mt. Bulusan, Dec.
: (
Samar: Catubig River, 1916, Ramos 24422 (US). Negros: Negros Oriental, Sibulan,
Malangco So., Sept. 1948, Edaño 6799 (GH). Balut Island, Oct. 1906, Merrill 5412 (NY,
US). Mindanao: Todaya (Mt. Apo), Davao, May 1909, Elmer 10463 (F, NY, US);
Agusan, Cabadbaran (Mt. Urdaneta), Aug. 1912, Elmer 13580 (F, GH, K, NY, US);
Bukidnon, vicinity of Tanculan, July 1916, Fenix 24936 (US); Cotabato, Nutol, 1932,
Ramos & Edaño 84871 (GH). CELEBES: “P. Boeton: Kaboengka”, Feb. 1929, Kjellberg
238 (S). New Guinea: (N. E., Morobe, bei Salamaua, Malalo Mission, Nov. 1936,
Clemens, cited by Mattfeld, Engl. Bot. Jahrb. 38:288). AMBOINA: Soja, Aug. 1913,
Robinson 1836 (NY, US).
CEANIA. AUSTRALIA. Queensland: Beaudesert, April 1907, Boorman (NSW);
Eumundi, March 1915, White (NSW); base of Mt. Gravatt near Brisbane, Nov. 1930,
White 6856 (NY). New South Wales: Billinudgel, May 1911, Stephenson 3 (NSW);
Hat Head, Jan. 1953, Constable (NSW); Durimbal to Berkeley Vale, Oct. 1953, Salasoo
(NSW). Sotomon Istanps: Guadalcanal, east fork o i è j
16 (US). Fiji IsLanDs: Vanua Levu, Thakaundrove, hills south of Nakula Valley, Nov
1933, Smith 333, Viti Levu, Mba, summit of Mt. Koro anitu, May 1947, Smith 4194 (US).
T^
o
D
KI
8
<
>
E
0
oe
ka
+
2
d
—
»
e
Aug. 1905, Rechinger 1844 (W, 3). Hawanan ISLANDS: Hawaii, Kilauea region, July
1929, Degener (NY); Oahu, Tantalus Mt., Honolulu, June 1923, Degener 1519 (NY
Lanai, Kalulu, March 1916, Munro 534 (NY); West Molokai, Hauakea Pali, April 1928
Degener 18133 (NY); Maui, east of Plinda, Oct. 1916, Hitchcock 14924
CULTIVATED: “Senecio valerianaefolius Link, Hort. Berol.” ); "Senecio valerianae-
folius h. p." (FI, ex Hb. Webb. ex Hb. Desf., type of S. valerianaefolius Desf., marginal
florets 4-fid, pistillate).
>
>
There is some question as to who actually authored the 1825 seed list, whether
Wolf or Link, and this I have been unable to establish. Every effort to locate a
copy of this list has so far been unsuccessful. I have been unable to learn even
whether the first appearance of the name was accompanied by a description or not.
If not, then the first description is the brief but sufficient paragraph by Sprengel,
and the citation would be "Link ex Sprengel.” But Reichenbach and Lessing, who
cite Sprengel, both credit the name to Wolf. Therefore I accept Wolf as the
original author pending finding of the 1825 seed list. The sheet, “Senecio valeri-
anifolius ex h. Raffeliano, 1825,” is in very excellent detailed agreement with
Reichenbach’s description, and appears to be the original from which his pl. 85 was
prepared. In view of the reported destruction of any type material there may have
been of this species in the Berlin Herbarium, I designate this sheet as neotype.
Gardner gave his S. valerianaefolius the number 2 52 in the ‘London Journal,’
but the corresponding specimen at Kew is Gardner 522, Organ Mts., Brazil. It is
a monstrous specimen with the capitula mostly much distorted as if by an infection.
(The symptoms are very similar to, if not identical with, those produced by the
aster yellows virus.) A few capitula at the apex of the inflorescence are fairly
normal, however, and show the typical structure of E. valerianaefolia, including
pistillate marginal florets and a faintly colored pappus. The leaves are quite typical
of f. valerianaefolia. Gardner’s proposed species, which is really not valid anyway
since it is based on a monstrosity, thus reduces to E. valerianaefolia £. valerianaefolia.
1956]
BELCHER—REVISION OF GENUS ERECHTITES 29
Erechthites Gardneriana Cabrera, based on it, is therefore superfluous.
Lessing transferred S. valerianaefolius Wolf to Crassocephalum, as emended and
greatly extended by himself. It is true that the appendage of fused papillose hairs
which terminates the style arm of E. valerianaefolia is somewhat longer than that
of E. hieracifolia, and might be taken to indicate intermediacy between the latter
species and the widespread Crassocephalum crepidioides. The heavily ribbed. brown
achene of E. valerianaefolia, however, is scarcely or not distinguishable from that of
E. hieracifolia, whereas the characteristic achene of Crassocephalum is shorter, more
uniformly cylindric, weakly ri and short-haired on the ribs, and reddish in
color. The florets of Crassocephalum are tubulous, rather than filiform, and are
more coarsely lobed.
2b. ERECHTITES VALERIANAEFOLIA f. des cio: o gee Belcher, comb. nov.
Erechtites organensis Gardn. Lond. Jour. Bot. 7: 420,
Erechthites valerianaefolia var. organensis (Gardn.) Bien "Mart. Fl. Bras. 6%: 300. 1884.
Differing from the typical form in having the leaves very finely divided, the
segments entire or only minutely serrulate; plants usually of small stature.
Holotype: Brazil, Rio de Janeiro, "open bushy places on the Organ Mountains,
3000 ft., March 1841", Gardner 5790 (BM!).
Known only from subtropical Brazil and Bolivia.
"ear cca xxu i Seer peri (S); São João, March 1910, Dusén 9349a
(BM, K, S). Minas Ger , Vauthier 306 (G Prodr.); Caldew 1845,
Waleres- 210 (S, 2); Caldas, wa 1973, SC III 794 (S, 3). Boma: Bridges (BM);
Bang 2068 (BM; not F, K, which are f. valerianaefolia)
In addition to the finely divided leaves, dier s species was characterized by
its hairy stem, smaller capitula, and shorter achenes, hispid instead of villous. The
hairy stem is frequently found in f. prenanthoides and not uncommonly in f. valeri-
anaefolia from widely scattered parts of its range. As for the smaller capitula,
those of Gardner 5790 have the phyllaries 7.5 mm. long, not 9 mm. as given by
Gardner, but other specimens of f. organensis exceed this while numerous specimens
of f. valerianaefolia closely approach it, so that there seems to be no useful discon-
tinuity. Moreover, I have yet to see a truly villous achene, such as Gardner
ascribed to E. valerianaefolia, anywhere in the genus Erechtites proper.
Although of the three non-typical forms of the species this one comes closest
to having a distinct geographical range, it does not replace the typical state within
that range but occurs with it as a comparatively minor element. Nor is the mor-
phological distinction by which it is recognized a very great one. For these reasons
this taxon is better given the status of a form. Hasskarl” arrived at a similar
conclusion more informally: ‘‘Misschien is E. organensis Grnd. [sic!] (Wlp. Rep.
II. 906. 2) niets, dan eene oude of magere vorm." There remains, of course, the
possibility that it is ecologically very distinctive, but this can not be decided on
the basis of the very meager information which accompanies the material cited.
25 Verh. Meded. Kon. Akad. Wet. 5:100. 1857.
[Vol. 43
30 ANNALS OF THE MISSOURI BOTANICAL GARDEN
The type was obtained at an elevation of 3000 feet in a brushy opening on the
mountain, whereas Dusén collected it in the Paraná Valley in swampy places at
unspecified elevations not more than three hundred miles further south (but a
greater distance westward). Nothing is stated about the habitat of the specimens
from Bolivia.
2c. ERECHTITES VALERIANAEFOLIA f, reducta Belcher, forma nova.
Folia 4 vel 5 superiora abrupte reducta, circa 2 cm. longa et 1 cm. lata, sessilia,
pinnatifida, segmentis utrimque 4-6, lanceolatis; folia ad caulis mediam 8-10 cm.
longa, 3.5-5 cm. lata, subpetiolata, inaequaliter dentata seu lobata, lobis distinctis
plerumque 2 vel 3, raro 4.
Specimina typica (syntypi 2) legit Dusén (n. 14156) in Brasilia, Paraná, prope
Antonia opp. in locis ruderatis, 29 Aug. 1912 (S!).
Differs from the typical form by the upper leaves being sessile, markedly reduced
in size for several nodes below the inflorescence to as little as 2 cm. long and 1
cm. wide, deeply pinnatifid, with 4 to 6 lanceolate lobes on either side, often much
more dissected than the median cauline leaves, which seldom have more than 2 or
3 distinct lobes per side but are 8 to 10 cm. long and 3.5 to 5 cm. wide and sub-
petiolate. The plants are simple below the inflorescence and of somewhat reduced
stature, 40 to 70 cm. tall. Capitula are comparatively few. Occurs in southern
Brazil; also Mindanao, Philippine Islands, and has been in cultivation.
IL: Rio de Janeiro [?], Gardner 5528 (BM); Bahia, Lockhart (BM); Ceará, June
1929, Bolland 39 (K). PmumnuPPiNESs: Mindanao, Camaguin, 1912, Ramos 14452 (US).
ULTIVA "Senecio valerianaefolius H. P.
valerianaefolia DC.", ex hort. bo
This well-marked foliage state has never been described previously, as far as I
can discover. From the limited number of specimens of this form among quite
numerous specimens of E. valerianaefolia, it would seem to be very infrequent in
the field. Because of its comparatively small stature, it would seem rather more
likely to be collected by the casual collector, who often selects individuals of a size
easily pressed, than the somewhat larger and more succulent f. valerianaefolia. 1
am unable to establish any valid pattern of distribution from the limited material
seen so far. This interesting form might well be the object of careful attention
from collectors, in order to extend the data on distribution and frequency. It
should also be brought into cultivation along side the other forms for comparison
and genetical analysis.
The very diffuseness of distribution, as indicated by the quite characteristic
specimen from Mindanao, suggests that this form may simply be the product of a
recurring mutation. That it is fertile is implied by the two plants from the Petro-
grad garden, which are presumably of the same progeny, and also by the two speci-
mens from Antonina, Paraná. Both sets imply that there might be populations.
2d. ERECHTITES VALERIANAEFOLIA forma prenanthoides (Kunth) Cuatr. in
herb., comb. nov.
Cacalia prenanthoides Kunth in HBK. Nov. Gen. & Sp. 4:167. 1820, folio ed. 4: 131.
820; non A. Gray, Proc. Amer. Acad. 19: 55; 1885; E
+» J. deParis, 1828” (G Prodr.) ; "Erecbtbites
t. Petropolitano (K, US).
1956]
BELCHER—REVISION OF GENUS ERECHTITES 31
Fees ew DC. Prodr. 6: 295. 1838, excl. sen: non Sch. Bip. Bull. Soc, Bot.
Erecht d Es, nth. Pl. Hartw. 209. 1845. Type: Popayan, Hartweg 1160 (K!).
Senecio albiflorus Sch. Bip. Flora 28: 498. 184 se
88.
praia dree (Kunth) Green xs Hieron. ex Hieron. Engl. Bot. Jahrb. 29:
d o 28: 628. 1901, Type, Pod. e 5665 (K!); mon DC. Prodr.
6: on
Differs from the typical form in having all the leaves comparatively smaller,
more or less petiolate, undivided, either merely serrate or at most the upper ones
pinnately incised on the basal portion only; stem sparsely setaceous-hispid, leaves
more or less scurfy-pubescent on the nerves beneath.
Syntypes: “Cacalia prenanthoides”, Humboldt & Bonpland (P, 2 sheets, in Herb.
Humb. & Bonpl.!).
Of sporadic occurrence over much of the range of forma valerianaefolia.
BraziL: Sellow (G Prodr., holotype of E. ambigu ua DC.). Rio Grande do Sul: Cruz
Atta, April 1893, Malme 776 (S, 2). Paraná: São João, March 1910, Dusén 9349 (S, 2).
Minas Gerais: Lagoa Santa, Warming (S); Caldas, 1866-67, Regnell 1 272 (S, complete
plant, this sheet only; others of did number with later date are typical form); Caldas,
April 1874, Mosén 1421 (S). COLOMBIA. Sec? ponerse Silvio Yepes Agredo 328 (F);
ad pagum El Tambo, June 1938, von Sneidern 1531, June 1939, von Sneidern 2770 (S).
Valle: Cordillera Occidental, Hoyo del Rio Cali, WA El Abismo, Cuatrecasas 18651
a
F ecuerdo à 4
2806 (BM). Peru: Pennell 13982 (F). Ecuapor: Pichincha, Tandapi, July 1920,
Holmgren 825 (S); Tungurahua, Hacienda San Antonio pr. Bafios, Dec. 1937, Sydow 575
(S); Eastern ceci ra, here of Biver ne Rimbach 492 (S); Camp E-4275 (F).
MEXICO: Ver z, Galeotti 2242 (K,
INA: rn tung, Kao-Yao Dist., rios Woo Shan, beside stream, July 1932
20160 ( AINAN: Kau-en Disk: ., Chim Fung Ling, near Sam Mo Watt Kg
April 1934, Lau 3870 (S, right only, left is id form). Samoa: Savaii, Safune,
in rain ue along trail, May 1924, pieds 121 ( A
L ExcLupEp: Of the several numbers of Elmer widely distributed as E.
petiolate . all dix I have seen are small states of f. vids rianaefolia with the petiolate entire
lower leaves carried upward higher than usual, but with the upper leaves mince pinna
ifid. Mandon 117 (K) det. as E. ambigua is E. hieracifolia var. cacalioide
The Humboldt & Bonpland specimens are cited as syntypes because Kunth
included both in his description, as indicated by the two measurements given,
"sesqui- aut bipedalis". One specimen is 18 inches long, the other totals 22 inches
in two pieces, Kunth’s careful description of the leaves as petiolate, lanceolate-
oblong, pinnatifid-incised toward the base, and decurrent on the petioles, makes
clear that this name applies to the taxon with petiolate subentire leaves. The place
of collection of the type specimens, usually precisely stated by Kunth, is for this
species given dubiously as "Nova Hispania?" There is nothing on either syntype
to correct this or to confirm it, unless it be the number “20” which occurs on one
of the labels. Since this form has been collected from Mexico by others, in the
vicinity of Vera Cruz, it may be that Mexico is the type locality, although it
appears to be much more frequently found in Colombia. Kunth called the plant
[Vol. 43
32 ANNALS OF THE MISSOURI BOTANICAL GARDEN
a perennial, even while confessing "folia radicalia ignota". It actually appears to
be a strict annual.
Greenman and Hieronymus apparently were the first to recognize Cacalia pren-
antboides as an Erecbtites. They transferred Kunth’s specific epithet to this genus,
reasoning that since the name Cacalia prenantboides (1820) was older than Senecio
prenanthoides A. Rich. (1832), the combination Erechtites prenanthoides (A.
Rich.) DC. must fall and be replaced by E. Labillardieri Hieron.! Greenman and
Hieronymus appear to have overlooked the appropriately named E. petiolata Benth.
Erechtites ambigua DC. was reduced by Baker to Erechtites hieracifolia. This
error apparently can be traced to a sheet laid in at Kew as the "type" of E. ambigua.
This is Mandon 117, determined by Schultz Bipontinus?? as Erechtites ambigua in
the list of determinations of Mandon's plants. The sheet at Kew has four specimens
on it, all being E. hieracifolia var. cacalioides, If Baker accepted this sheet as
authentic E. ambigua, he was fully justified in the reduction, and, indeed, it seemed
quite logical in the light of de Candolle's ambiguous description of the leaves, which
actually approximate those of some states of E. hieracifolia, and the flat statement
that the pappus was white, not purpurascent. Although the pappus in the exposed
capitula on de Candolle's holotype of E. ambigua has faded to an off-white except
where concealed by the phyllaries, the capitula in the packet have distinctly pinkish
pappus hairs, and de Candolle's statement is inaccurate.
Asa Gray, on unspecified evidence, equated Cacalia prenantboides with Senecio
runcinatus Less. (E. ? runcinata DC.) and maintained Kunth's name as the proper
designation for the latter. Lessing’s species actually is a discoid homogamous
Senecio with unappendaged style arms and must be maintained as a Senecio. Its
corollas are reddish purple and do not agree at all with Gray's definition of Cacalia
as white-flowered with deeply cleft corollas, Cacalia prenantboides A. Gray is
thus a later homonym of C. prenanthoides Kunth.
After I had established the above synonymy, I found a specimen in the herb-
arium of the Chicago Museunr of Natural History, collected by Cuatrecasas in
Colombia and determined by him as “E. valerianaefolia forma prenantboides".
Cuatrecasas considers that this petiolate state, which he finds to be characteristic
of the paramos, is an ecological response to the peculiar sort of xerophytic environ-
ment found there, with intense insolation. I can not wholly agree with him,
mainly because of those specimens, probably including Kunth's syntypes, which
have been collected at lower elevations from distant and discontinuous points, but
I am glad to acknowledge his previous recognition of the identity and subsidiary
status of E. prenanthoides (Kunth) Greenm. & Hieron. ex Hieron. The possible
ecological status of this form needs to be investigated in the experimental garden.
3. ERECHTITES MISSIONUM Malme, Kungl. Sv. Vet.-Akad. Handl. 32: 73. 1899,
excl. var. lanceolata Chod. & Hassl. Bull. Herb. Bois. Il; 3: 732. 1903:
Annual herb. Stem strongly sulcate, glabrous, much branched above, to 1 m.
high (or taller), leafy. Leaves as much as 20 cm. long and 8 cm. wide, usually
somewhat less, approximately three times as long as wide, petiolate with the petioles
*6 Bull. Soc. Bot. Fr. 12:80. 1865.
1956]
BELCHER—-REVISION OF GENUS ERECHTITES 33
narrowly or not at all winged, ovate to subovate, acute to briefly acuminate, irreg-
ularly incised-serrate with acuminately callose teeth, glabrous or minutely hairy
beneath, sometimes more or less pinnately lobed toward the base of the blade, with
lobes long-triangular and directed forward with their principal veins diverging
from the midrib at an angle from 45? to 60? and the sinuses rounded, the lobes
numbering from one to four on each side. Inflorescences terminal and axillary
cymose panicles, congested in bud, becoming diffuse at anthesis, glabrous or some-
times with minute tightly appressed simple hairs on juvenile parts; peduncles sub-
erect and elongating to 2.5 to 5 cm. at maturity; bracts subtending the primary
branches much reduced in size but similar in shape to the cauline leaves, bracts of
the secondary branches filiform, bracteoles of the peduncles 2 to 4, linear, 2 to 5
mm. long. Capitula solitary, at anthesis about 15 mm. long including pappus hairs,
about 5 mm. in diameter when pressed, ventricose; calycular bracteoles few, linear,
much shorter than the involucre, glabrous or with minute simple hairs; involucre
of 12 or 13 attenuate-subulate phyllaries 11 to 13 mm. long, 0.7 to 1.0 mm. wide
at base, abruptly reduced to a width of 0.5 to 0.8 mm., then linear, acute, finely
multinervate with nerves becoming indistinct toward the apex, glabrous or mi-
nutely hairy in bud. Pistillate marginal florets in two or three rows, sometimes with
rudimentary stamens; corolla filiform, 9 to 10 mm. long, 5-fid, with slender, acute
lobes 0.5 to 0.6 mm. long. Disc florets hermaphroditic, more numerous than the
marginal; corolla slenderly infundibuliform, 11 to 12 mm. long, 5-fid; lobes 0.5
to 0.6 mm. long, slightly then abruptly tapered to an acute and slightly papillose-
thickened apex. Style-arm apex briefly appendaged with a cone of fused papillose
hairs which has a fringe of divergent hairs at its base in the hermaphroditic florets.
Achene 2 to 2.5 mm. long, subcylindric, dark brown and puberulous between
ribs. Pappus S exceeding phyllaries by about 2 mm. and equalling the
florets.
Indigenous to dis Paraná River Basin in Argentina, Paraguay, and Brazil; also
known from isolated stations in Peru and Venezuela, an extension of the previously
reported range.
BraziL: Rio Grande do Sul: Colonia Ijuhy, in "rogas" nec non juxta vias in silvas, April
1893, Malme 744 (BM, isotype; S. holotype, 3). Paraná: Cahnon, March 1910, Dusén 9317
(S, an E. valerianaefolia?) ; Iacarehý, 1914, " Dusén 15271 (BM, F, Si: Jaguariabyva, Nov.
1914, Dusén 15899 (F, S); Tres Barras, Jan. 1916, Dusén 17624 (S, very immature);
Nova Galisia, Feb. 1916, Dusén 17700 (S). PARAGUAY: ` Caaguazú, Nov. 1874, , Balansa
Ekman 1104 (S). Peru: 1835, Matthews 1739 > YE Rizoma: “prope co odism
Tovar", 1856-7, Fendler 1972 (K); “Bajo Cotiza, en la sombra al lado de rio", Sept. 1940,
Vogl 417 (F).
This species is not easily distinguished from glabrous states of E. bieracifolia,
the most usable characters being the nature of the leaf base and the length-width
ratio of the leaf blade. Other characters, such as relative length and width of
capitulum, number of florets, size of phyllaries, time of flowering, etc., are of little
value because of variation in these features within both species. There is some
[Vol. 43
34 ANNALS OF THE MISSOURI BOTANICAL GARDEN
ground for suggesting that it is the South American equivalent of E. hieracifolia
var. bieracifolia, and should be treated as no more than a variety. This possibility
requires investigation by hybridization experiments and other such analyses, how-
ever, before it can be more seriously advanced. Until such studies have been made,
the two species should be kept apart, although they are undoubtedly very closely
related.
Judging from the rather meager ecological data on the tickets, this species is
largely confined to the forest edge in rather moist situations. It is worth noting
that altitudinally it ranges from near sea-level (below 200 m.) in the Paraná Valley
at the southern end of its range at about 30° S. latitude, to well above 1000 meters
in the Sierra Mérida of Venezuela, at about 8? N. latitude. What its altitudinal
location in Peru may be is not indicated, but it is perhaps somewhat higher there,
nearer the equator. It is very unlikely that the distribution of this species is as
disjunct as these scattered records indicate. A search for it along the eastern foot-
hills of the Andes on the edges of the forest openings and along trail-sides at about
the 1000-meter contour from Venezuela to Peru and gradually descending south-
ward to Bolivia and Argentina might establish many additional stations.
This species, aside from the original description, and the erroneous assignment
to it of var. lanceolata by Chodat and Hassler (see below, under E. goyazensis), I
have seen alluded to only once, by Cabrera’. This was merely a statement that it
was one of four well-known species of Erechtites in Brazil.
Chodat and Hassler undoubtedly must have assigned Hassler 8362 to E. mis-
sionum as var. lanceolata because of the resemblance of the capitula. But this
resemblance does not extend to the florets nor to the leaves, whereas in both of
these features there is detailed agreement between the several duplicates of Hassler
8362 and the type of E. goyazensis. The variety is to be excluded from E. mis-
sionum.
$ GOYAZENSES
4. ERECHTITES GOYAZENSIS anon ge Een 7: 54. 1950.
Senecio Goyazensis Gardn. Lond. Jou
preces missionum Malme var. SC SC e KE Bull. Herb. Bois. II, 5:732.
cobi suffruticose, 1 to 1.5 m. tall, branching from near the base, branches
erect. Stem glabrous, leafy, with internodes only approximately 1 cm. long. Leaves
sometimes 20 cm. long and 2 cm. wide, but usually less, with sharply and minutely
callose-serrate margins and acuminate apices; midrib inflated beneath; lateral veins
numerous, fine, prominent, diverging from midrib at an angle of 30? or less, glab-
rous; lower leaves narrowly oblanceolate or rarely obovate, subpetiolate, upper
leaves lanceolate to linear-lanceolate with semi-amplexicaul bases. Inflorescence of
several to many capitula in a paniculate corymb from terminal and axillary branch-
es; capitula at anthesis solitary on elongated peduncles 2 to 3 cm. long, which bear
27 Brittonia 7:54. 1950.
1956]
BELCHER—REVISION OF GENUS ERECHTITES 35
several linear-subulate bracteoles, calyculate with short linear bracteoles; phyllaries
of the involucre 12 to 14, glabrous, linear, 9 (to 13) mm. long, acute. Florets
rather variable, with corollas coarsely filiform to infundibuliform; the outermost
one or two florets in each spiral series pistillate or with more or less well-developed
staminodia, their style-arm apices rounded to shortly appendaged; inner florets all
hermaphroditic with corolla campanulate, deeply 5-lobed; the lobes linear-lanceo-
late, 1.0 to 1.25 (to 1.5) mm. long, 0.25 mm. wide, conspicuously papillose-thick-
ened apically; style-arm apices appendaged above a fringe of divergent papillose
hairs. Achene 2.5 to 3 mm. long, strongly ribbed, glabrate. Pappus white, slight-
ly exceeding the phyllaries and subequaling the florets.
Upper Paraná River Valley in Brazil, with an outlier in northeastern Paraguay;
co-extensive with E. ignobilis Bak.
BRaziL. coras: near Natividade, in shady woods, Dec. 1839 (or bushy places near
Ville Natividade, Jan. 1840), Gardner 3300 (BM, holotype; K, 2; F ex G, fragment;
; raph e
1865, Reenell DI 795 (S, 4); 1844, Weddell 1603 (P ex Hb. Sch. Bip., det. as S. ignobilis).
s&o PAULO: "in paludosis ad Mugi", Nov. 1833, Lund 844 (G. Prodr., det. as F. bieraci-
folia). wrrHour specific DATA: Pohl 633 (K, 2); Pohl 2056 (W). PARAGUAY: Bellayis ta
(Apa), Hassler 8362 (S); El
BM: K, 2; Sp : Y) ): Spec e d pes 16174 (BM, K; det. as S Gore uis
H H . uded: ll, > H . D
but is Ee Bak. var. leptotus Cabr.).
Cabrera gave the following reason for transferring Senecio Goyazensis Gardn.
to Erechtites: ‘The marginal florets of this species are tubulous without anthers.
The involucre is also typical of Erechthites.” He did not cite specimens, nor is it
clear that he examined the type. The sheets at Kew and the British Museum
(Natural History) had not been loaned, but Gardner 3300 appears to have been a
very widely distributed number, and he probably saw an isotype.
Dissection of representative capitula on the type and on other specimens reveals
that the trend toward unisexuality of the outer florets is only partially and quite
variably developed. Some capitula have all the series-terminating florets with
rudimentary stamens or an occasional floret with one or two stamens polliniferous.
Other capitula show complete abortion of anthers in the terminating florets, with
some reduction in the next one or two florets inward in each spiral. Furthermore,
the terminal appendage of the style arm is variable. Occasionally it is merely short-
conic with only a few very short papillae at its tip, but with a corona of low pollen-
presenting papillae below, while sometimes the apical papillae are more elongated
and fused, with a much closer approach to the tuft of fused hairs which charac-
terizes E. hieracifolia, Never have I observed a truncated or merely low-domed
apex such as is common in Senecio and is seen in S. leptanthus Phil., which has been
also (I believe, wrongly) reduced to Erechtites by Cabrera.
I retain Gardner's species in Erecbtifes where Cabrera has placed it because of
this conic and sometimes appendaged style-arm apex, and because of the size of the
capitulum and the number of florets. In these latter features it is much closer to
E. hieracifolia and E. missionum than it is, for example, to the erechthitoid species
[Vol. 43
36 ANNALS OF THE MISSOURI BOTANICAL GARDEN
of Senecio in Australia, which are characterized by usually much smaller heads and
by quite abruptly truncated style arms. Finally, the large and strongly ribbed
glabrate achene indicates closer affinity with Erechtites than with Senecio, it being
in fact scarcely distinguishable from that of E. bieracifolia.
The type of E. missionum var. lanceolata, Hassler 8326, has leaves rather nar-
rower than those of Gardner 3300, but is identical with it in floral features, includ-
ing the very deep lobing of the corolla. The difference in leaf width is bridged
by Pohl 633, which is intermediate between these two. The several specimens of
Gardner 3300 are not all alike, the holotype (BM) having leaves both shorter and
wider than on the other sheets, whence the measurements in Gardner's description.
Pohl 2056 (W) is another broad-leaved specimen which is otherwise in good agree-
ment.
Both Gardner 3300 and Hassler 8362 have corolla lobes linear-lanceolate, 1.25
mm. long, 0.25 mm. wide, with the apices glandulose-thickened; whereas Malme
744 (holotype of E. missionum Malme) has corolla lobes deltoid, 0.5 mm. long and
0.33 mm. wide, with apices only slightly thickened. Malme 744 has leaves petio-
late, broadly lanceolate to ovate, and is irregularly toothed or subincised; whereas
both Gardner 3300 and Hassler 8362 have leaves sessile, narrowly lanceolate, and
regularly serrate. I am certain that the proper affiliation of Hassler’s specimen is
with E. goyazensis rather than with E. missionum, as Chodat at first supposed.
The sheets in the Delessert Herbarium reveal that Chodat later determined Hassler
8362 as a small-headed form of E. ignobilis, but neither Chodat nor Schultz Bipon-
tinus ever published this name. Actually, there is good reason to suppose that the
name Senecio ignobilis may have originally been given by Schultz Bipontinus to a
specimen of this taxon of Gardner's rather than to the large-headed taxon to which
Baker later applied it, judging by specimens in his herbarium (see below, under E.
ignobilis) .
The affinites of E. goyazensis are much closer to E. ignobilis than to any of the
other species of Erechtites, but the two species differ not only in the sizes of the
several parts of the capitulum but also in corolla lobing and in leaf margin and
venation. Were it not for these features, E. ignobilis might be simply a polyploid
of E. goyazensis, and these differences do not exclude the possibility. The relation-
ship of these two co-extensive perennial species might well be the subject of a cyto-
genetic study.
5. ERECHTITES IGNOBILIS Baker, in Mart. Fl. Bras. 63: 299. 1884.
Senecio ignobilis Sch. Bip. in sched. ex Baker, l. c., in syn. nom. nud.
Perennial from a woody base; shoots subherbaceous, glabrous, 40 to 80 cm.
high (or more?), sparsely branched in the inflorescence only. Leaves rather
crowded toward the base, margins remotely and sharply dentate, apices acute;
lowest leaves obovate to oblanceolate, subpetiolate, 7 to 10 cm. long, 12 to 18 mm.
wide, upper leaves lanceolate to linear-lanceolate, sessile, smaller. Inflorescence
corymbose, consisting of a few capitula borne singly on long peduncles which
terminate the stem and the few axillary branches; capitula 18 to 25 mm. long,
1956]
BELCHER
ERECHTHITOID SPECIES OF SENECIO 37
9 to 12 mm. wide when pressed. Involucre oblong, of about 15 linear, glabrous
phyllaries 16 to 20 mm. long, with 4- or 5-nerved keel and scarious margins,
exceeded by both pappus and florets at full maturity; outermost one or two florets
of each spiral-series filiform, functionally pistillate but frequently containing rudi-
mentary stamens; remainder of florets hermaphroditic; style-arm apices shortly ap-
pendaged; corollas 16 to 18 mm. long. Achene 4 mm. long, subcylindric, strongly
ribbed, glabrate, dark brown; pappus abundant, white, exceeding phyllaries, sub-
equalling florets.
Easily distinguished from E. goyazensis by the much larger and fewer capitula.
Lectotype: "Brasilia, ex herb. hort. Petropolitani, rec'd. 11/66, Senecio igno-
bilis Sch. Bip. (teste Sch. Bip.) ", Riedel (K!).
Apparently confined to the Paraná River Valley.
AZIL. "Brasilia, etc.", Riedel Ph d ex hb. hort. Petrop., S. ignobilis Sch. Bip
in Sch. Bip.), Brasilia", Riedel (W); “ in paludosis Yttu, Febr. 1836”, Riede
(P, ex Hb. Sch. Bip., not det. as S. Mallet Ge GERAIS: Lagoa Santa a,
(K, F; photograph as no. 106 ex Hb. Haun., F); Lagoa Sant Lund (S). s&o ee
Canna [?] verde, Feb. 1849, Regnell III 796 (S). PARANÁ: Turma, Jan. 1910, Dusén
9061 (S); Jaguariahyva, April 1910, Dusén 9698 (S, 22 cm. high, only 2 capitula) Dusén
9727 (S), Oct. 1910 Dusén 10528 (S), Dec. 1910, Dusén 11006, Nov. 1914, Dusén 15994
(S), Dusén 15998 (F, S). PARAGUAY: Caaguazú, Hassler 9255 (K, S, W); in regione
fluminis Corrientes, LE E 5864 (K, G Deless., S).
Baker described this species as an annual herb, but several specimens, including
Riedel 379 (W), Hassler 9255 (K, W), and indeed two of the sheets at Kew deter-
mined by Baker, including the lectotype, have stems with distinctly woody bases
and portions of perennial rootstocks attached.
Five collections are cited by Baker in the original description. Since no one
specimen is designated as the nomenclatural type, all must be considered as syn-
types. The name-bringing specimen being Riedel's, I have designated it as the
lectotype.
The inclusion of Senecio ignobilis in synonymy by Baker does not constitute
valid publication, and its association with Riedel's specimen appears to be the result
of a curatorial error, probably at the St. Petersburg herbarium, which Baker, un-
suspectingly perpetuated. The specimens in Schultz’s herbarium which bear this
epithet all belong to other taxa.
ERECHTHITOID SPECIES OF SENECIO IN AUSTRALASIA
Senecio L. Gen. Pl., ed. 5, 375. 1754, et auct., in sensu extenso.
Trees, shrubs, or herbs; the erechthitoid species mostly semi-woody perennials.
Capitula with all florets perfect and infundibuliform, or with the marginal florets
more slender, pistillate, and either ligulate, subligulate, or irregularly or regularly
2- to 5-fid and sometimes with rudimentary stamens that very rarely may be pol-
liniferous; style-arm apices truncated or low-domed, with crown of divergent papil-
lose hairs at least in the perfect florets, and not prolonged into an appendage of
fused papillose hairs.
[Vol. 43
38 ANNALS OF THE MISSOURI BOTANICAL GARDEN
The erechthitoid species are distinguished from true Erechtites by the style-arm
apices truncated or low-domed rather than prolonged in an appendage of fuse
papillose hairs, as well as by smaller and less heavily ribbed achenes and usually
pgs capitula. Represented by numerous species in Australasia, with outlying
pecies in New Guinea and one in Java; occasionally met with elsewhere, as S.
imita Phil. in Chile and forms of S. flavus Sch. Bip. (S. Decaisnei DC.) from
Arabia. Two Australasian species, S. minimus and S. glomeratus, are adventive on
the Pacific Coast of the United States.
De Candolle divided the Australasian species of Erechtites into three sections,
MICRODERIS, TULODISCUS, and PLAGIOTOME. Subsequent authors have only rarely
attempted to classify their new taxa as to section. This probably reflects the fact
that the separations made by de Candolle are highly artificial and do not stand
examination. In returning these species to Senecio I have not retained these sections.
Neither is it advisable to create a new section in Senecio for these erechthitoid spe-
cies, Although they can, for the most part, readily be separated from the other
Australasian species of Senecio, there are within the group some species which clear-
ly intergrade into the discoid group and others which intergrade into the radiate
group. Therefore, pending a badly needed revision of Australasian species of Senecio
proper, I shall refer to the heterogamous material only as erechthitoid Senecio.
Our understanding of the relationships between the non-erechthitoid species of
Senecio in Australasia is poorly developed, since it has not passed beyond the crude
separation into discoid versus radiate species. The soundness of this separation has
been questioned by virtually every competent syntherologist from the time of Lin-
naeus to the present. For example, Bory de St. Vincent?5, in criticizing Thunberg's
confused efforts to restore Jacobaea Tourn., claimed that Linnaeus joined (dis-
coid) Senecio and (radiate) Jacobaea as Senecio on the premise that the presence
or absence of ligulate florets is not a constant character even in the same individuals
of a single species. In support of this, Bory cited his own experience of finding
that the late autumnal flowers of several radiate species, notably Senecio Doria L.
(Jacobaea pratensis, altissima, Limonii folio Tourn.) lacked the ligulate florets
ordinarily so conspicuous in earlier flowerings, and so could have been classified
into a different genus from the one to which they would have been referred some
months earlier, if the radiate and the discoid species were separated as Thunberg
proposed.
A related situation is the existence of radiate varieties of discoid species, such
as the not uncommon var. radiatus of S. vulgaris, the standard species of the genus,
or the ligulate var. fallax (Greenm.) Fern. of S. pauciflorus Pursh. This latter
species was regarded by W. J. Hooker?? as a "rayless state of S. aureus".
Examination of a large number of specimens of S. glomeratus (E. arguta DC.)
has revealed that in this species in particular the boundary between radiate and non-
radiate conditions is very imperfect. A single specimen often possesses some pistil-
28 Ann. Gén. Sci. Phys. 1:304-305. 1819.
?9 Flora Boreali-Americana 1:332-333. 1834.
1956]
BELCHER—ERECHTHITOID SPECIES OF SENECIO 39
late marginal florets so obliquely one-toothed as to be actually briefly ligulate,
some that are irregularly bidentate, others that are irregularly 3-fid with one sinus
twice or thrice as deep as the others, and still others that are perfectly regularly
3-fid with shallow sinuses. The same situation has been observed, but much less
frequently, in other erechthitoid species.
In the folder of Senecio aureus var. subnudus at Kew there is a specimen (Great
Slave Lake, Gates & Mellenby) which indicates that the outermost whorl of florets,
those terminating each phyllotaxic spiral in the capitulum, is unstable in another
direction. This is a discoid specimen which has rare marginal florets with stamens
non-polliniferous, although the style-arm apex is clearly truncated. This is a close
approach to the condition in Erechtites, especially in section Goyazenses, where an
occasional marginal floret may bear a set of rudimentary stamens. It is a situation
found over and over again in the erechthitoid species of Australasia, and is the basis
for the statement made above concerning the intergrading of the discoid and the
erechthitoid groups.
Another instance of an erechthitoid Senecio is the Chilean S. leptanthus Phil.,
recently transferred by Cabrera?? to Erechtites because of its rather coarsely fili-
form pistillate marginal florets. This has a velutinous, weakly ribbed achene and
low-domed, non-appendaged style-arm apex and is much better left in cio.
The significant fact is that erechthitoid states of Senecio are not confined to Aus-
tralasia, but are also found elsewhere. This adds weight to the point made by
genus Senecio could have slender pistillate florets. The limits of Senecio are not
being unduly expanded or strained by the inclusion of heterogamous non-ligulate
material from Australasia.
KEY TO ERECHTHITOID SPECIES OF SENECIO
A. pee 5 to 9, rarely 1 ome but not all capitula B
ginal florets riis silia disc florets androgynomorphic but with
Pauls abortive and style ar unded and de nsely hairy on oute Verona "e itg ae
o
BB. Marginal florets pistillate or occasionally with incomplete set «eel stam
dentate, disc florets fertile, style arms truncate, not densely hairy on gie ba KC
hairs
1. S. bipinnatisectus
D
rets not or scarcely ex bell in nu
. Leaves scarcely or not at all dps gens oarsely and irregularly toothed,
teeth varying in size and spacing and often denticulate; florets 5-fid, margi-
nal florets devoid of rudi + stamens, disc florets infundib pay
. S. biserratus
EE. Cauline leaves auriculate, coarsely but regularly toothed, teeth almost Sen
in size and spacing; all I ets 4- e subfiliform, and perfect, but es num
r of stamens varying from 1 t caf esc iia
DD. Achenes with fine white sg in hairs on sharp narrow LM esaet in
the grooves; florets twice or more as numerous as the phyllari
30 Not. Mus. La Plata 14 (Bot. No. 69):76. 1949.
40
[Vol. 43
ANNALS OF THE MISSOURI BOTANICAL GARDEN
F. Bo ria and regularly toothed, 8 per cm., variably arachnoid
beneath, glabrate with age, sparsely dd or glabrete shave, ot lobe
28 or blong Til: stem glabrate or beset wi cen Pes ute unicellular
nimus var. minimus
FF. Lori less regularly det ees with pre hispid geris ce in
addition to arachnoi irs, more or less lobate, especially ied lower ones,
Dirt to oblong; stem grise with hispid — hai
. mimimus var. Mero
AA. Phyllaries 11 to 13 or nips or deed a minority of capitula ih n or 10
G. Ph
yllaries 16 o
c cH: iniret es Si Zeie variably
ees irregularly “dentate or lobate, teeth or lobes use, pubescence
ixed, hispid and arachnoid; phyllaries MEAT on inner fa
5. S. squarrosus
HH. Achenes long and attenuate; leaves linear-lanceolate, margin sharply
coarse loboid teeth; pubescence
h oe on stem, leaf, and inflorescence; phyllaries GE
both i and outer fac 6 Ps
GG. Phyllarie rad to 15, usually 12 or 13 I
1
es only about 3 times as dius as wide; plants glabrous or nearly
» Ace enes 1.5 to 2.0 mm. long, short-cylindric, densely and uni-
formly covered with very short appressed Bed — d
p to 8 cm. lon
wide, ovate-lanceolate or broadly Ai LaS irregularly
coarse-toothed, with sharply €— teeth, auriculate and semi-
amplexicaul at base, entirely glabro 7. S. laceratus
JE et d to A m m. long, hert subc ylindric, not densely and
Se leaves oblong,
eiert: sinuate- dentate with 3 to 5 teeth
rous above, minutely
4 to 5 Eu E and 1 to 1.5
cm. wide; plant of New Guinea
II. Leaves more than 3 times longer than wide; pubescence various ................
K. Se
va
and stems glabrous or sparsely Bess REY when
L "beret eege Zeite the geg long, — -
retrorse, apical segments on upper lea orm-acum
ae 1 cm. ia ma cu us al oasis E 40, disc flor ets
abou m. long, subrostrate,
m hairs on uide ribs..9. S. See SC HS
es subentire to iie bens: with lobes broad,
Sege pinnatifid
henes 4 mm. long, faintly gege E or ie
white-haired i in ~ rooves; leaves oblong-line
ong, obtuse, s uate-den RE Fuse San or e.
ifid; marginal florets 10 to 15, disc florets about twice as
numerous; phyllaries 6 mm. "Bas with short-acute spices
E
3
E
8
4
. Wairauensis
MM. Achenes 3 mm. long, gege ribbed, hairs suberect on
or beside the ribs but not in the bottom of the grooves;
phyllaries 5 to 5.5 mm. dud ith apices abruptly nar-
rowed and then bluntly pig be radical leaves oblan-
ceolate, long-attenuate to subpetiolate, 3.5 cm. long an
6 mm. wide; cauline leaves linear-lanceolate to vam
oblong, 6 cm. long, 5 mm. wide, remotely a
to subentire S. d SE
rid us may z ge ci here;
y leaves linear-lance
(Glabrate states of S. quadridentat
may p» zed b , with mar-
gins minut deleite and strongly aos and phyl-
laries reset acuminate.)
1956]
BELCHER—ERECHTHITOID SPECIES OF SENECIO 41
KK. Leaves and stems conspicuous sly pubescent, hairs arachnoid or
hispid or both combined; sometimes nearly icit at maturity
do entire or denticulate; achenes more or less fusi-
form, 2.5 . long or lon
O. Leaves Do to emen to as oe, as 9
ide; phyllaries 6.5 to 8 long, i Gr
had then glaheatá, except ses in 2- jaa
lower third only, apices acuminate ........ Wi E
OO. Lower leaves oblanceolate or obovate lancet te,
cm., 0.3
st
to 12 cm. long, 1.5 to 1. cm. wide; p es 6 mm.
long, glabrous or Kee arachnoid at jeep: -nerved,
ratte n apices acum 13. S. gunnii
N. Leaves toothed, D Gd Firepulcy lobed, achenes
ver? cylindric, 1.5 to on, P
. Rec e an As ase oe involucre lanate; leaves
amie quu beneath, sparsely arachnoid to
glabrate abov Q
Q. Achene with 10 low broad ridges; apex of phyl-
lary acute with scarious margin ............ . S. glomeratus
QQ. Achene iioi £ extrem z narr igh thi
ridges; ape
phyllar
acuminate eu alios locke ioci marg
S. laticostatus
PP. Receptacle and involucre glabrous; E "m
strongly 2-ridged; leaves with crisped mulcicellulr
mr beneath and his spid y fece Mort: rarely gl
ec renge auricu
callos di to coarsely dentate
or echt "lob ate
S. Hairs on upper leaf surface with mta tin
late Zoch achenes + st D 75 mm. long........
6a S. bispidulus v e? EEN
SS. Hairs on upper leaf surface with
tuberculate bases; Portam es 1. Y^
Gs RS RUN ETE CE : sidan var. en "e
RR. pow once- prse isl e Ker nts denticu
16 hispidulus var. TH
PPP; Receptacle and involucre gl e us or densely lan
ate; leaf pubescence both arachnoid and hispid;
chis usually short, various ico
S. glom
ratus X S. pre
1. SENEcIo bipinnatisectus Belcher, nom. nov.
d ëmt? F. Muell. Frag. Ate do 5: 88.
iae F. Muell. ibid. ut synonym, nud.; non Senecio Atkinsonii C. B.
age Chee. Ind. 207. 1876.
Stout plant with stem strongly striate, subglabrous or sparsely short-haired,
branched above with branches ascending, densely leafy; basal portion not seen.
Cauline leaves and bracts pinnatisect or bipinnatisect, with segments irregularly
denticulate or subentire, slender, markedly revolute, sessile, with pinnatisect auricles;
lower leaves up to 10 cm. long with segments up to 5 cm. long, size gradually
reduced upward; older leaves glabrate, younger subarachnoid beneath with a few
glandular or tuberculate-hispid hairs above. Inflorescences decompound, corym-
varying in expansion from very simple corymbs with 3 or 4 capitula per
bei to repeatedly rebranched clusters 25 cm. in diameter with scores of capitula,
axes glabrous throughout or sparsely short-haired on peduncles; peduncles with
[Vol. 43
42 ANNALS OF THE MISSOURI BOTANICAL GARDEN
several simple, subulate, erect, sometimes ciliolate bracteoles. Capitula usually with
8 phyllaries, sometimes as many as 11 on some but not all capitula; phyllaries 6
to 6.5 mm. long, slender, usually 2-nerved and only moderately keeled, glabrous or
minutely glandular-pubescent, apices shortly tapered, obtuse or very slightly acute.
Florets slightly longer than the phyllaries and equaling pappus, marginal ones pistil-
late, filiform, 3- or 4-fid, twice or thrice as numerous as the hermaphroditic slen-
derly infundibuliform 5-fid disc florets, Achene 2 mm. long, short-subcylindric,
not attenuate-rostrate, dark red, ribbed, glabrous or sparsely set with very short
white hairs especially when immature.
Apparently confined to the Coastal Ranges of New South Wales and south-
eastern Queensland.?!
AUSTRALIA. NEW SOUTH WALES: Blue Mountains, Louise Atkinson (K, det. by Muel-
ler, syntype duplicate and lectotype); Monkey Creek towards Port Jackson, Woolls (MEL,
syntype); Sydney, Wilkes Exp. (US); "Senecio, Australasia, N. S. W.", Hugel (W);
rushy Mountains near Gloucester, 1881, Betche (NSW); Port Macquarie, 1898, Boorman
(NSW); Port Stacking, Feb. 1899, Camfield (NSW); Blackheath, April 1899, Maiden
(NSW, W); Bulli Pass, 1900, Hamilton (NSW); Blackhurst, swamp, Jan. 1903, Cam-
field (NSW); Jervis Bay, 1926, Rodway (K, NSW); Beaumont, 1935, Rodway 1687
(K); Braidwood, 1936, Stenfield (NSW). QUEENSLAND: Blackwall Range, April 1918,
W bite ; Candle Mountain, May 1918, White (NSW); Main Range, top of Mt.
Mitchell, 3760 ft., White 6872 (NY). wirHouT LocaLirY: “Iter Australiense 1802 to
1 n R. Brown 2279 (K); "Nova Hollandia, Erechtites sonchoides Cand.", Ferd. Bauer
( S
The new epithet refers to the characteristic deep division of the leaf and bract
blades. As stated by Mueller, this species differs from all its erechthitoid congeners
by the form of itsleaves. But in Senecio it resembles S. anetbifolius in its pinnat-
isect leaves with narrow divisions. Comparison of specimens of S. bipinnatisectus
with the type of S. anethifolius (G!) shows them to be readily distinguishable
even in the sterile state. The leaf segments of the former are strongly revolute, ir-
regularly denticulate, minutely tuberculate-hispid above and subarachnoid beneath,
whereas those of the latter are slightly revolute, non-denticulate, and glabrous.
The floral features are, of course, much more distinct, the former having mod-
erately keeled and minutely glandular-pubescent phyllaries 6 to 6.5 mm. long, and
marginal florets pistillate, filiform, and scarcely exceeding the phyllaries; whereas
the latter has glabrous phyllaries 5.0 to 5.5 mm. long, and marginal florets her-
maphroditic, infundibuliform, and much exceeding the phyllaries. I do not know
any other Australasian species with which either of these might be confused, unless
it might be S. bispidulus var. dissectus (Benth.) Belcher. In this variety, however,
the leaves are only once or very imperfectly twice dissected, the segments are
sharply angulate-lobate rather than denticulate or entire, and the pubescence is
coarsely tuberculate-hispid. The inflorescence is, of course, unmistakably distinct
from both the above species.
31 The description of this and each subsequent erechthitoid species is followed by a list of speci-
mens examined, with the herbaria in which they are found indicated by the symbols proposed by
Lanjouw. In the case of the more commonly collected species, the list is a selected one to show
the range of distribution and variation. Of less common species, all material examined has been
cited. In each case the typifving specimen, if seen, is listed first.
1956]
BELCHER—ERECHTHITOID SPECIES OF SENECIO 43
Mueller gave the number of phyllaries as "11 to 13”. I have not found any
capitulum with more than eleven, and the majority had only eight. The lectotype
at Kew does have a few heads with up to eleven. A certain amount of instability
in the number of phyllaries seems to be common in those genera of Compositae with
essentially uniseriate involucres, and indeed seems rather widespread in the family.
For individual plants, however, it is usually possible to find that the modal phyllary
number of a majority of the capitula is very close to one of the peaks of the phyl-
lotaxic series: 8, 13, 21, etc. Further, there is generally a correlation with other
characters of taxonomic significance, so that the use of the modal number of phyl-
laries as an easily seen key character is valuable. Because this modal number for
S. bipinnatisectus is 8, and seems not to reach the value of 13 assigned by Mueller,
I have placed this species in the group with eight phyllaries.
e number of florets per capitulum is quite variable, and little dependence
should be placed on the exact numbers given in descriptions, as by de Candolle.
Richard, who appears to have studied these species more carefully, omitted almost
all reference to specific numbers of florets. Approximate numbers, however, are
helpful, and the ratio between pistillate and perfect florets may be significant.
For this species a representative count is sixteen pistillate to five perfect florets in
one capitulum, a ratio of about three to one.
Immature buds of this species closely resemble those of S. minimus in size,
shape, number of phyllaries, and particularly in a spiraled appearance. This is
caused by the unexpanded phyllaries running slantingly rather than lying straight
from base to apex. The leaves also agree with those of S. minimus in having usually
quite large auricles, and these two species are undoubtedly quite closely related.
2. Senecio biserratus Belcher, nom. nov.
Senecio flaccidus A. Rich. Sert. Astrolabe, 110-112. 1834; non Less. Linnaea 5: 161. 1831.
gn sonchoides DC. Prodr. 6: 296. 1838; non Senecio sonchoides Kunth, in HBK.
v. Gen. & Sp. 4:178. 1820.
Erebi prenantboides Benth. Fl. etra 3: 658. 1866, pro parte, non DC.; Black, Fl.
ustral. 4: 609-610, pl. 50, 1929, non DC.
Ze erect, to 5 feet tall, Akan or glabrate, simple or sparingly branched
above, leafy. Leaves rather crowded and suberect, membranous, as much as 11
cm. long and 4 cm. wide, gradually and proportionately reduced upward, sessile, the
lower ones slightly or much attenuated and not clasping, the upper ones auriculate
and semiamplexicaul, all oblong to oblong-lanceolate, serrate, with irregular larger
teeth again finely toothed, glabrous or sparsely set with rather long multicellular
hairs. Inflorescence a corymbose panicle, open and lax at maturity, the branches,
peduncles, calyculi, and involucres essentially glabrous; bracteoles broad, subulate.
Involucre of 7 or 8 phyllaries; phyllaries 5 to 6 mm. long, alternately narrower and
broader, to 1 mm. wide, keel more or less conspicuously 2-ridged with a broad or
narrow shallow groove between, apices alternately moderately long-acute and quite
bluntly obtuse (ie. probably actually 2-seriate), all minutely ciliolate, with a
marked tendency to curl after achenes have been shed. Florets scarcely exceeding
phyllaries in number. Marginal florets pistillate, filiform, 5-fid, lacking rudi-
[Vol. 43
uL ANNALS OF THE MISSOURI BOTANICAL GARDEN
mentary stamens; disc florets hermaphroditic, slenderly infundibuliform, 5-fid.
Achene 2.5 to 2.75 mm. long, subcylindric, slightly tapered toward base, apex not
attenuate-rostrate but slightly contracted below the apical callus, with white hairs
in grooves between heavy low flattened ribs. Pappus and florets exceeding in-
— by 1 to 2 mm.
USTRALIA. TASMANIA: “N. Holl. détr. d'Entrecasteaux”, ex itin. Baudin (P. holotype);
“Neus Hollande cote merid*, Mus. de Paris, 1821” (G Prodr., isotype?, holotype of E
sonchoides ); Deloraine, Jan. 1902, Maiden ); unt Field East, ca
March 1906, Maiden (NSW); Eaglehawk Neck, Nov. 1924, Lucas ( ; Hobart, near
Myrtle Gully, Casca ope — burnt out previous summer, ca. 900 ft t
n u: u: D
right herb 414 ft. tall”, March 1940, Gordon (HO, leaves as much as 11 cm. long, 4 cm
wide, coarsely lobed); Teapa Valley Adamsfield track, 1500 ft., Feb. 1943, Gordon
(HO); Gordon, Dec. 1952, Curtis (HO). victoria: Port Fairy, W ban (NSW); Port
Fairy, Nov. 1900, Walker 7 (NSW). NEW SOUTH WALES: Port E R. Brown 2277
(K). New ZEALAND. SOUTH ISLAND: Canterbury, Akaroa, Raoul 30 (K); Otago, Lyall
(K); Otago, Milford Sound, Lyall (K ex Hb. Hk. as E. prenanthoides, left- hand specimen
only; right is S. wairauensis). STEWART ISLAND: "Ia arenosis maritimis", Thouroude (P).
AUCKLAND ISLANDS: “Erechtites arguta var. glaberrima”, Wilkes Exped. (K, US).
Bentham’s reduction of this perfectly good species to Erechtites Zeene
DC. Ge, S. minimus Poir.) is unjustified, despite several points of similarity. By
direct comparison of the types in the Prodromus Herbarium they may be separated
rather easily by differences in achenes and leaf margins. The achene of S. biserratus
is 2.5 to 2.75 mm. long, flat- and broad-ribbed, with hairs between the ribs; that
of S. minimus ranges from 1.5 to 2.0 (to 2.25) mm. long, is sharply and narrowly
ridged, and is hairy on the ridges rather than between them. The leaves of the
former are coarsely and irregularly doubly serrate, as the new epithet indicates,
whereas those of S. minimus are uniformly finely denticulate. Again, the former
species is virtually glabrous except for the achene and sparse appressed hairs on
both sides of the leaf, lacking the arachnoid pubescence on the lower surfaces of
the leaves and on the juvenile parts which characterize the latter.
E. prenanthoides Black, by figure and description, is S. biserratus. The figure of
the cross-section of the achene which shows hairs only in the narrow grooves is
critically diagnostic, as is the irregularly toothed leaf. Black gives the distribution
simply as "South-East". If this species actually is in southeastern South Australia,
then it should also be more generally present in Victoria, to link up with its pres-
ence in New South Wales and Tasmania. The specimen from the Auckland Islands,
incidentally, is the most southerly specimen of erechthitoid Senecio which I have
seen
De Candolle’s diagnosis of E. sonchoides, based on a specimen at Geneva, was
formed independently of Richard’s, but agrees fairly well with it. He was uncer-
tain of the identity, and queried Richard’s name in the synonymy. He need not
have been so cautious. His type and that of Richard are certainly conspecific, and
are probably a part of the same gathering, by the expedition of Baudin. From de
andolle’s account of the small number of phyllaries and florets, it seemed E. son-
choides might be an Arrhenechthites. Accordingly, the florets, and particularly
the style arms of the central florets, on the type were carefully examined with the
aid of detergent solution. The central florets are hermaphroditic, developing nor-
1956]
BELCHER—ERECHTHITOID SPECIES OF SENECIO 45
mal rather than abortive achenes; the style arms are long, recurved, and stigmatic;
the style-arm apices are squarely truncated and not papillose on either apex or
dorsum. It is a good Senecio despite the reduced number of parts
3. SeNEcIO kermadecensis Belcher, sp. nov.
Perennis (?), caule herbaceo, solum versus apicem ramoso, dense foliaceo; foliis
sessilibus late auriculatis, amplexicaulibus, vix constrictis super auriculas, ovato-
lanceolatis, inaequaliter dentatis, acutis vel subobtusis, nonnihil scabris, subglabris,
majoribus (infimis haud visis) 13 cm. longis, 4 cm. latis; inflorescentiis corymbosis;
capitulis cylindricis, calyculatis; involucri squamis plerumque 6 (5 — 7), linearibus,
glabris, 5 mm. longis; floribus paucis, numero squamis similibus, omnibus tenuiter
filiformibus, apice vix dilatatis, 4-dentatis; staminibus 1-4 numero variante in flo-
ribus capituli singuli, omnibus polliniferis, ramis styli truncatis, papillis marginal-
ibus divergentibus praeditis; achaeniis subcylindricis, 2 mm. longis, 10-costatis, sub-
appresse puberulis inter costas; pappo niveo, subsetaceo, pluriseriato.
Specimen typicum legit W. R. B. Oliver in insula prope Novam Zeelandicam
"Sunday Island" dicta in Archipelagine Kermadecensi. Endemica species. Speci-
men in herbario Kewensi.
Cheeseman, in 1925, confused this species with E. prenanthoides DC.,
and attributed to W. R. B. Oliver the statement that it was "not uncommon on
Sunday Island" of the Kermadec Islands. Oliver's specimen at Kew, the holotype,
bears that statement on its label. Erroneously determined as E. prenanthoides DC.,
this specimen superficially resembles that species, but is actually distinct and appar- |
ently undescribed.
It is remarkable in the genus for its unusual capitulum, with phyllaries 5 to 7,
usually six, and the florets about equal in number. All florets appear to have styles
and functional stamens, the number of stamens varying within the same capitulum
from 4 to 1. The florets are nearly filiform, very little dilated apically, and 4-fid;
the style-arm apices are truncate with a fringe of diverging papillae. Some achenes
in the capitula examined were less developed than others in the same capitulum,
suggesting partial sterility, but this appeared not to correlate with the position of
the achene in the capitulum nor with the number of stamens. The achenes, although
similar in size to those of S. minimus, resemble those of S. biserratus in being
hairy between the ridges. The leaves, though resembling those of S. minimus in
general shape, were much more broad and coarse than is typical for that species.
Senecio kermadecensis must be regarded as a discoid Senecio of unusual interest,
Suggesting a transition between the discoid and the erechthitoid kinds, as shown by
the partial sterilization and the slenderness of the florets. It also suggests a transi-
tion between Senecio and Arrhenechthites, as shown by reduction in floret number
and possible tendency to abortion of some ovaries. Its closest affinity in the erech-
thitoid group is not with S. minimus, as Oliver and Cheeseman thought, but with
S. biserratus, as shown by the drastic reduction in floret number and by the acheni-
al pubescence. I hesitated to describe it without examining more material. But
since it raises the interesting possibility that other peculiar endemics may be
masquerading as well-known species, it seemed desirable to call attention to it.
[Vol. 43
46 ANNALS OF THE MISSOURI BOTANICAL GARDEN
4. SENECIO MINIMUs Poir. in Lam. Ency. Méth. Bot. Suppl. 5: 130. 1817.
Inflorescence a corymbose panicle, usually large with very many capitula, dif-
fusely much branched, branches suberect or lax; arachnoid in bud, glabrate when
expanded; bracts subtending the branches rapidly and progressively reduced in size
upward, two or three bracteoles on each ultimate peduncle, capitulum with calycu-
lus of 5 or 6 short slender inconspicuous bracteoles. Involucre of 8 phyllaries,
rarely one or two more or less; phyllaries 6 to 7 mm. long, linear, alternately 2-
nerved and broader with broadly obtuse apex, and 1-nerved and narrower with
narrowly obtuse apex, glabrous or subglabrous but with apices ciliolate and slightly
darkened or not. Florets exceeding involucre, slightly exceeded by pappus; margin-
al florets filiform, briefly 3-, 4-fid, somewhat more numerous (ca. 12 versus 7)
than the disc florets, which are hermaphroditic, slender, slightly expanded into a
narrowly infundibuliform 5-fid limb. Achene 1.75 to 2.0 (to 2.25) mm. long,
short-cylindric, dark reddish-brown or brown, with fine white subappressed hairs
on the narrow ribs, the grooves glabrous, non-attenuate but with apex callose-an-
nulate. Pappus white, filiform.
4a. SENECIO MINIMUS var. minimus
b y.
Erechtites pumila DC. Prodr. 6: 297, 1838, said by DC. to be based on “S. pumilus Poir.”,
ich is non existent; non Armstrong, Trans. N. Z. Inst. 13: 338. 1887.
6. 1838; non A. Rich. (as Senecio, based on
Gaudicbaud 4, P!, which is S. quadridentatus) ; non Greenm. & Hieron. Engl. Bot.
:296. 1838, ut syn.; non Linn. Sp. Pl.
Erechtites prenanthoides Hook. f. Fl. N. Z. 141. 1853, pro parte, excl. var. minor Hook. f.
Senecio Miilleri Regel, Ind. Sem. Hort. Bot. Imp. Petrop. 31, 1863; non Kirk, Trans. N. Z.
Inst. 15: 359. 1883; non Erechtites Muelleri Lange, Ind. Sem. Hort. Haun. 28. 1861
[not seen]; Bot. Tidskr. II. 4: 5. 7. 3. 1874
Senecio Warscewiczii Hort. Berol. apud Vatke, App. Ind. Sem. Hort. Bot. Berol. 21. 1875;
B A
non A. Br. & Bouché, Ind. Sem. Hort. Bot. Berol. App. 13. 1851; Linnaea 25: 29
1852 (from Guat
emala).
Senecio heterophylla [sic " Colenso, Trans. N. Z. Inst. 27: 389. 1894.
Erecbtites Labillardieri Hieron. Engl. Bot. Jahrb. 29: 63. 1900, superfluous.
Herbaceous plant of somewhat glabrate aspect, robust specimens exceeding 1
m. in height, others as low as 15 cm. Stem erect, sulcate, simple or sparingly
branched below the inflorescence or with numerous short axillary branches with
much-reduced leaves, glabrous or sparsely set with minute white hairs. Leaves
rather variable in size and shape, lower cauline ones on robust specimens reaching
or somewhat exceeding a length of 10 cm., 1.5 cm. wide, size rather rapidly reduced
upward on stem, width decreasing proportionately more rapidly than length; lower
leaves broadly lanceolate, upper linear-lanceolate; lower portion of blade not or
only slightly constricted, rarely subpetiolate, the base slightly to conspicuously
expanded into clasping lobes or auricles; the reduced leaves of axillary branches
sometimes appearing non-auriculate, subpetiolate, oblanceolate; margins evenly
denticulate, with 5 to 8 teeth per cm., or slightly irregularly denticulate with some
teeth a little coarser but never pinnatifid; youngest leaves densely arachnoid below,
sparsely so above, older leaves glabrate.
1956]
BELCHER—ERECHTHITOID SPECIES OF SENECIO 47
Generally distributed in temperate Australia, Tasmania, and New Zealand;
adventive weed in California and Oregon.
ALIA. TASMANIA: "Capite Van Dieman”, Labillardiére Gr. “Senecio hiera-
Gieres Pres Bi Mardiére , N. Holl.”, with specific epithet deleted and "minima" written
beneath in De serps erch P, Te cio minima Enc. Sup., labii Nov. Holl."
fragment of holot x Hb. Poir.; BM; K; G Deles.; G Prodr., "Senecio byrracifolius e e
nova hollandia, m. A abillardiére 1808", laus of E. prenantboides DC.); G 175,
Georgetown (K, 2), Marlborough (K), St. pores (K, NSW); Maiden, Rechocché Bay
1908, Swanport to Swansea 1902, Port Arthur 1906, Mount Field East 1906 (all NSW);
Russell Falls, March 1910, Cheel NSW); Lilydale, Jan. 1943, Wardrop (HO). victoria:
` Vendu Vale, Robertson 469 (K, NSW); Dandenong Range, Mueller (K); Blacks Spur,
(NSW). NEY N
e x Po c (NSW,
W); Mt. Kembla, Nov. 1899, Fletcher (NSW); Blackhurst, à AH D. Canfield (ve in
non-contiguous pieces, the sterile one with leaves to 16 cm. long and 4 cm. ut
otherwise PA uent
LAND: "New Zealand 1769-70", Banks & Solander (US); Nelson, Graham
River, Cheeseman (US); Wairau Valley, Travers 16 (K); Colenso (K, isotypes (?) of S.
beterobbylla) ; Hooker (W ex Hb. Sch. Bip.) ; Westland, Parvu, Jan. 1937, Lothian (K).
UNITED STATES. CALIFORNIA: Ge Re 6 July 1931, Jones 59096 (BM); Humboldt
Co., Trinidad, Sept. D Parks 01058 (BM, F, MICH, NSW); Marin Co., Almonte,
edge of salt menm, July 1944, Howell 19857 (F, S). OREGON: Lane Co., Aug. 1949,
S
TED: E *809 Senecio Warscewiczii” 62 Hort. Berol. (P ex Hb. Sch. Bip.);
a, Mueller?”, Hort. Bot. Petrop., 1863 (K, isotype (?) of S. Miilleri Regel).
I designate the piece of S. minimus at Paris as a fragment of the holotype,
rather than an isotype, because the jagged base exactly matches the stump of a
branch on the holotype. Poiret evidently obtained this small branch for himself
when he described the species. All the other sheets of Labillardiére’s “Senecio
hyeracifolius” which I cited are authentic isotypes, with the sole exception of the
ligulate fragment on the sheet at Florence from Labillardiére’s own herbarium,
and there are probably still other sheets in other herbaria.
The isotype of S. Miillerii Regel was compared with the holotype fragment of
S. minimus and found conspecific. It was unquestionably distinct from presumably
authentic specimens of E. Muelleri Lange which were raised at Vienna from seed
from Copenhagen and which appear to be a form in the hybrid swarm, S. hispidulus
X S. quadridentatus. Vatke’s assumption of identity between Regel's species and
Lange's, made in a note clarifying the status of S. Warscewiczii Hort. Berol., was
an error.
This is one of the most distinctive of the erechthitoid species of Senecio, and
shows little affinity with the other species. Its closest resemblance is to S. biserratus,
as discussed above, although the two are readily separable. Cheeseman’s statement
that this species is “not uncommon on Sunday Island” of the Kermadecs appears
to be based on Oliver's specimen, the holotype of S. kermadecensis. S. minimus
is not to be included in the flora of the Kermadec Islands on this evidence.
[Vol. 43
48 ANNALS OF THE MISSOURI BOTANICAL GARDEN
4b. SENECIO MINIMUS var. picridioides (Turcz.) Belcher, comb. nov.
Erecbtites picridioides 'Turcz. Bull. Soc. Imp. Nat. Mosc. 24: 200. 1851; Black, Fl. S.
Austral. 4: 610. 1929; non Sond. & Muell. Linnaea 25: 253. 1852, which is S. runcini-
folius Willis.
Erechthites prenantboides DC. var. picridioides (Turcz.) Benth. Fl. Austral. 3: 658. 1866.
Differing from var. minimus in having the stem more robust and coarse and
beset with hispid multicellular hairs; the leaves larger, lobate, and hispid, with
acute callose teeth; and the corollas somewhat variable in number of lobes.
WESTERN AUSTRALIA: Swan River, 1845, Drummond 132 (K, 2; FI; all isotypes).
I have not seen the holotype of this taxon, but the isotypes which I saw agreed
well with the description. Dissection of florets, however, revealed much variation
in lobing of the corollas. Pistillate florets were 2-fid, 3-fid, or 4-fid; the perfect
florets were 3-, 4-, or 5-fid. I have not observed such variability in var. minimus.
From hispid pubescence and callose-dentate leaves, as well as this variation in lob-
ing, I suspect possible hybridization with S. bispidulus. But because of the great
similarity in the other floral and the fruit characters between these specimens and
the holotype of var. minimus 1 have maintained the varietal status given them by
Bentham, pending further study.
Turczaninow stated: "Species Er. senecioidi et argutae affinis." I find no
other reference to the former, and it is presumably a nomen nudum. The Kew
sheets of Drummond 132 have tickets reading: “Erechtites / E. sonchoidi DC.
Prodr. / Sw. riv. Drummond”. “E. sonchoidi” is written hastily, and likely was
miscopied by Turczaninow.
Black maintained this taxon as a species, separate from his E. prenanthoides.
This was fully justified, since the latter is actually S. biserratus! He gave for E.
picridioides a wide distribution in South Australia; namely, Encounter Bay, Kan-
garoo Island, Murray Lands, Yorke and Eyre Peninsulas, South-East. I have seen
none of these specimens.
5. SENECIO SQUARROSUS A. Rich. Sert. Astrolabe, 107, tab. 35. 1834.
Erechtites Richardiana DC. Prodr. 6: 297. :
Erechthites bispidula Benth. Fl. Austral. 3: 660. 1866, pro parte; non (A. Rich.) DC.
838; non Black, Fl. S. Austral. 4: 610. 1929.
Stem erect, simple, leafy, sparsely arachnoid. Leaves lanceolate-linear, 7 to 10
cm. long, 0.8 to 1.0 cm. wide, acute, minutely to coarsely remotely toothed to sub-
lobulate, glabrous or sparsely arachnoid beneath and somewhat scabrid above; lower
subpetiolate, upper sessile with more or less coarsely toothed auricles. Inflorescence
a terminal cyme of relatively few capitula, sparsely arachnoid on peduncles and
bracteoles, Capitula about 10 mm. long, 9 mm. wide, calyculate, bracteoles of the
calyculus linear, arachnoid, Involucre of 16 to 20 linear phyllaries 7 to 8.5 mm.
long with arachnoid bases, hyaline margins and recurved apices. Marginal florets
pistillate, corolla filiform, 7 mm. long, 5-fid, lobes long-acute, 0.4 to 0.5 mm. long,
0.1 mm. wide, apices glandular, sinuses not all always of same depth. Disc florets
hermaphroditic, corolla slightly infundibuliform, 5-fid, lobes similar to those of
the marginal florets but slightly broader, style arms exserted. Achene 2 mm.
long, blackish, thick-cylindric, rather thickly set with short white or tawny hairs.
1956]
BELCHER—ERECHTHITOID SPECIES OF SENECIO 49
Pappus white, slightly exceeding phyllaries and florets.
AUSTRALIA. NEW sovin WALES: Port Jackson, erreurs d A, agus frag. G
rodr.). vicrORIA: Wendu River, in forest, Sept. 1842, Robertson 289 (K); Wendu
Vale, Sept. 1843, Robertson 690 (K, on same sheet Wa 289 and another large-headed
Robertson speci th indet.); in meadow between Melbourne and D t's
Creek, O 52, Mueller (K e k., immature pens appears to be this species); ?
Cataract Hills, 1863, Mueller (NSW). TASMA a, 4 Nov. 1844, Gunn
(K, on same sheet specimen of Launceston, Gunn 508, which is indet. but
definitely not this species); Launceston, "1 1844, Gunn 508 (K, a separate ber
from the preceding) ; Van Dieman's Land, Pis 5 08 (W, rather more pubescent than the
Kew sheets); Hobart, Nov. 1923, Lucas (NSW); Blackman’s Bay near Kingston, Nov.
1935, Rodway 2039 (K
The identity of S. squarrosus was completely lost, due to Bentham’s inclusion of
it in his E. bispidula. Yet it is easily one of the most distinctive of the erechthitoid
species, readily recognized by its large capitulum, numerous phyllaries, short
blackish achene, and coarsely toothed leaves. The holotype agrees well with
Richard's plate and description, including the serration of the leaves. The abscis-
sion line (?) depicted at the base of each leaf is misleading, however. No such
feature can now be seen on the type, and it is likely only an artist's device for
marking the leaf base. One of the distinctive features, the number of phyllaries,
is not described by Richard, but can be recognized in the drawing of the capitulum.
Erecbtites bispidula Benth. is a mixture of at least two separate elements, as
shown both by the description and by the specimens at Kew determined by Bent-
am. Bentham mistook certain specimens of Gunn's distribution number, 508,
to be true E. hispidula, and to these added other specimens with large capitula
which superficially resembled them but which have long attenuate-rostrate achenes,
hence his statement, ““Achenes slender and striate as in E. quadridentata or rather
shorter.” A comparison of the Launceston specimen of Gunn 508, which Bentham
determined as E. hispidula, with the types of S. hispidulus and S. squarrosus showed
that is was identical with the latter, not the former, and has the short achene as
shown in the figure of S. squarrosus.
The large-headed specimens with long, attenuate-rostrate achenes are also char-
acterized by somewhat fewer phyllaries, nearly linear subentire leaves, and cottony
pubescence with sometimes a few hispid hairs added. It is to this group, rather
than to S. squarrosus, that Erechtites bispidula Black applies, judging from his
description of it as "near the preceding, E. quadridentata", and “achenes as in E,
quadridentata”. Certainly S. hispidulus is excluded by his description, particularly
by the dimensions given for the capitulum. The status of this group is still unset-
tled, but a plausible suggestion is that it may represent a polyploid state of S.
quadridentatus. It has not yet been given a name of its own by any one, and I
have refrained from doing so until its status can be clarified.
6. SENECIO PYROPHILUS Zoll. & Mor. ec Zoll. Nat. - en Geneeskundig Arch. v.
Neérland’s - Indié 2: 266. 1845; Syst. Verzeichn. 125. 1854. MER occ aei
i at. 2: 97. 1856; Koorders, Nat. Ty
"e : Tai 60: gt bei ra ob et Java 3: 342. 1912 (as E. byrobbila
(Zoll) Sch. Bip.).
Erecbtbites me ERE O. EE Rev. Gen. Pl. 1: 325. 1891; »on DC.
[Vol. 43
50 ANNALS OF THE MISSOURI BOTANICAL GARDEN
Perennial, with several to many spreading, then ascending, subherbaceous shoots
from a short woody base. Stems sulcate, arachnoid to lanate especially on younger
portions, with erect branches, leaves crowded on the basal portion and more distant
above. Lower leaves 6 to 9 cm. long, 4 to 6 mm. wide, linear-oblanceolate, long-
attenuate, not auriculate, sharply callose-denticulate with occasionally a few larger
coarse lobate teeth, subglabrous to arachnoid above, arachnoid to lanate beneath;
upper leaves only slightly shorter, not attenuate, linear-lanceolate, briefly and nar-
rowly auriculate just above the attachment. Inflorescences corymbose, at first
compact, then rather lax, with peduncles elongate and distant at maturity; lanate
when young, later glabrate; bracteoles few, scattered, linear-subulate. Capitula
few, with calyculus and receptacle arachnoid. Phyllaries 16 to 18, long-linear, 7
to 8 mm. long, 0.75 to 1 mm. wide; keel broad, flat, scurfy-arachnoid on entire
length, obscurely 2-nerved with nerves narrow and only slightly darkened, scarious
margins quite narrow, apices long-acuminate, minutely ciliolate. Marginal florets
approximately 40, pistillate, corolla filiform, 3- to 4-fid, about 6 mm. long; disc
florets about 20, hermaphroditic, corolla very slenderly infundibuliform, 5 to 6
mm. long, ca. 0.3 mm. in diameter, 5-fid. Achene 4 mm. long, light olive-brown,
narrowly subcylindric, very markedly attenuate-rostrate, nerves 5, rounded, low,
hairs short, scabrid, subappressed in shallow grooves between the nerves. Pappus
copious, slender, exceeding the phyllaries and subequalling the florets, niveous to
faintly tawny.
With somewhat the aspect of S. quadridentatus Labill. and doubtless related to
it, but distinguished from it by denser pubescence, more numerous phyllaries
scurfy-arachnoid all over the keel, and Ee achenes,
JAVA: Tengger, "Senecio ,Pyrop hilus flos lutei. In arenosis epe. M.
Tenanja-an (Tengger) 6-8000', XI.” D wie vn (P, 4 sheets, including one ex Hb.
Sch. Bip. and annotated b kim, isotypes); Bromo, 7000’, 16 Sept. 1875, i ries 6026
(NY, det. by him "Erecbtbites CECR DC. e E E E, byrobbila Sch, Bip. f;
*Ost- Java: auf dem Tengger bei und oberhalb Ngadisari von 2000-2400 m. ü. m an
den trockensten Stellen", Koorders 37403, 37404, 37780 (K, det by Koorders as E. his
dula [sensu Benthami]); Res. Besoeki, Yang Plateau, 2100 m., 11 Aug. 1916, Koorders
& Koorders-Schumacher 43646 (K).
The packet on Schultz's isotype sheet bears a notation: “. . . . anth. ecaud.,
flores ex. foem., cent. hermaph.” The reduction of this species to Erechtites, attrib-
uted by Miquel to “Schultz Bip. mss.”, doubtless was made as a result of this dissec-
tion. Schultz appears not to have examined the style-arm apices, which in this
material are definitely senecionoid. I do not find where Schultz himself published
the combination, and conclude that both citations by Koorders are technically in-
correct. In his “Exkursionsflora” he wrote of E. pyrophila: *. . . comp. Erechtites
hispidula DC. Prodr. VI. (1837) 296; Benth., Fl Austral. III. 660. Ich halte
diese australische Art vermutlich fur identisch mit der japanischen [sic !] E. pyro-
phila.” In this connection he cites Koorders 37402 bis 37404 and 37780, which,
however, he publishes as E. pyrophila. The sheets at Kew are determined as E.
hispidula DC., a misidentification which undoubtedly arose as a direct result of
1956]
BELCHER
ERECHTHITOID SPECIES OF SENECIO 51
his working at Kew with those Australasian sheets so thoroughly confused by
Hooker and Bentham. Some components of E. hispidula Benth. do resemble S.
byropbilus, but these are misidentified and are not S. bispidulus A. Rich. It is to
Koorders’ credit that he never actually published the reduction implied by his
comment.
But Koorders' provisional comment is reflected in the determinations of Com-
positae in Diel's “Beitrage zür Flora des Sarawaket-Gebirges". In commenting on
the determination of Keysser 40, Mattfeld?? wrote to the effect: that only a single
Indonesian species, E. pyrophila from East Java, belonged in the New Zealand and
Australian section of Erechtites, concerning which Koorders supposed that it was
identical with the Australian E. hispidula, to which then Keysser's plant might also
long, but that he, alas, lacked authentic material of E. hispidula, and so had to
determine it as E. arguta because of its pubescence and small capitula.
This confusion has prevented accurate identification of Keysser 40. could
not locate duplicates, and the specimens at Berlin and Breslau presumably are
destroyed. The characters given by Mattfeld could apply to either S. hispidulus or
S. glomeratus (E. arguta auct.). At least one can be certain from the small capi-
tula that Keysser's plant was not S. pyropbilus. It could, of course, be an endemic
novelty.
Although this species has somewhat the aspect of S. quadridentatus in the nar-
row elongate leaves, long-attenuate achene, and arachnoid pubescence, it is easily
distinguished by its coarser leaves with an occasional exserted tooth on the lower
ones, by its 16 to 18 phyllaries, by the very short pubescence which extends over
the entire length of the phyllary on both its faces, and by the slightly longer and
more slender achene. It differs from S. squarrosus by the phyllaries being pubes-
cent and lacking the reflexed apices which give the latter its name; it also has much
longer achenes and quite different leaves.
This species is known to me only from eastern Java, and is apparently the only
species from Sundaland, west of ““Wallace's Line”. A careful search eastward along
the summits of the Sunda Islands and particularly on Timor might conceivably turn
up additional stations or, possibly, even related species. At present S. pyrophilus
remains an interesting phytogeographical anomaly in the distribution of erech-
thitoid Senecio. It appears to be confined to the higher mountains at and above
2000 meters. From the notes of Zollinger and of Koorders it seems to prefer dry
sandy habitats or volcanic ash.
7. SENEcIo laceratus (F. Muell.) Belcher, comb. nov.
Erecbtites lacerata F. Muell. Linnaea 25: 417. 1852.
Erechthites eate Benth. Fl. Austral. 3: 659. 1866, pro parte; non (A. Rich.) DC. Prodr.
An erect annual with well-developed taproot, virtually glabrous. Stem un-
branched below the inflorescence or with a few small branches from the medial
32 Engl. Bot. Jahrb. 62:500. 1929.
[Vol. 43
52 ANNALS OF THE MISSOURI BOTANICAL GARDEN
axils, to 30 cm. high and disproportionately thick, up to 5 mm. in diameter at the
base, striate, densely clothed above with numerous appressed and overlapping
leaves. Leaves up to 8 cm. long, 3.5 cm. wide, usually not over three times as long -
as wide, ovate-lanceolate, the lower sometimes broadly oblanceolate, irregularly
coarse-toothed, each tooth usually with several sharply acute denticulations, the
upper leaves less irregular, auriculate and semi-amplexicual at the base, entirely gla-
brous. Inflorescence corymbose, rather crowded and compact especially before an-
thesis, glabrous except for occasional minute simple hairs on peduncles and brac-
teoles; bracts subtending main axes broadly denticulate-auriculate, rapidly tapering
to acuminately triangular; bracteoles on peduncles broadly subulate, appressed; caly-
culus of a few short slender subulate bracteoles. Involucre of 10 to 13 phyllaries,
much exceeded by florets and pappus; phyllaries glabrous, to 4 mm. long, 0.5 to
0.75 mm. wide at base, apices abruptly narrowed to acute or narrowly obtuse points
often reflexed at maturity, lower half of phyllary strongly keeled and usually with
a very narrow median nerve more or less prominently raised above the keel surface
but sometimes submerged in it. Capitulum of 25 to 30 florets, of which about 10
are hermaphroditic. Marginal florets pistillate, corolla 4-, 5-fid, 3 mm. long, fili-
form, styles slightly exserted, style-arm apices bluntly truncated. Disc florets
hermaphroditic, corolla 5-fid, slenderly infundibuliform, 3.5 mm. long, anthers
minutely sagittate, style-arm apices truncate. Achenes 1.5 to 2 mm. long, short-
cylindric, light brown and angular when immature, becoming plump, dark reddish
brown and very indistinctly ribbed when mature, whole achene except the straw-
colored annulus covered uniformly and rather densely with very short appressed
papillose hairs, glabrate at maturity. Pappus white, capillary, slightly exceeding
the florets.
Apparently confined to the interior of Australia.
AUSTRALIA. SOUTH AUSTRALIA: Cudnaka River, Mueller (MEL, paratype, teste Willis
[isotype?]; Basedow Range, 20 July 1889, Tietkins (MEL, det. by Muell). NORTHERN
TERRITORY: Bagot’s Creek, Horn Expedition, 1894, Tate (K).
Bentham included Mueller's type of E. lacerata in his E. arguía complex, pre-
sumably because of its rather small capitula and compact inflorescence, plus the
resemblance of its phyllaries to the larger glabrous ones of S. bispidulus, to which
the larger part of his description of E. arguta (S. glomeratus) applies (see below).
Mr. J. H. Willis has kindly informed me by letter that: “Bentham treated this as
an inland, coarse and glabrous variety of E. arguía; in which he has been followed
ever since at the Melbourne Herbarium". Mueller, at least as late as 1889, regarded
it as a separate species, as shown by his determination of Tietkin's collection.
is distinctive species is especially noteworthy for the proportionately greater
width of the leaf blade, approached among erechthitoid species only by S. papuanus,
from which it is otherwise distinct. It is also almost entirely glabrous, a feature
readily separating it from S. glomeratus, from which it can also be distinguished
by the entirely regular corolla of the pistillate florets and by the uniformly and
densely hairy (immature) achene.
1956]
BELCHER——ERECHTHITOID SPECIES OF SENECIO 53
The distribution can scarcely be indicated by the three specimens seen, except
as "inland". The Tietkins and the Tate specimens are both of terminal fragments
only; they have leaves somewhat smaller than those of the type, and are more
mature, The description is based on all three specimens.
8. SENECIO papuanus (Lauterb.) Belcher, comb. nov.
nura papuana Lauterb. Fedde's Repert. Spec. Nov. 13: 242. 1
Perennial; stem erect, leafy toward apex but lower leaves abscissed, very nearly
glabrous, terete, striate. Leaves 4 to 5 cm. long, 1 to 1.5 cm. wide (or larger,
below?), gradually reduced in size upwards, cuneate, oblong, sinuate-dentate with
3 to 5 teeth on each side, acute, glabrous above, minutely short-haired on nerves
beneath, sometimes purplish beneath; lower subpetiolate and slenderly auriculate,
upper sessile with coarser auricles, Inflorescences axillary and terminal, corymbose,
of two to several capitula borne singly on peduncles 1 to 5 cm. long and bearing a
few linear acute bracteoles 3 mm. long. Capitulum 11 mm. long, 4 to 5 mm.
wide, with calyculus of 2 to 5 linear bracteoles subapical on the peduncle. Phyl-
laries 12 to 14, 7 to 8 mm. long, obtuse. Marginal florets pistillate, in two rows;
corolla filiform, apex slightly expanded and 5-fid; style-arm apices slightly domed,
without corona. Disc florets hermaphroditic; corolla slenderly infundibuliform,
regularly 5-fid; style arms short, apices truncate and flat with corona of very short
diverging hairs. Achene (immature) 3 to 3.5 mm. long, slenderly subcylindric,
not attenuate-rostrate but with expanded apical annulus; pappus multiseriate,
white, slightly exceeding phyllaries and equaling florets.
NEA: Kaiser-Wilhelmsland, Bolan, 2400 to 3000 m., 1913, Keysser 317
(cited by Mattfeld as in Hb. Breslau, fragment and photograph in Hb. Berlin, both
presumably destroyed. Isotype, BM!) ; Papua, Albert Edward Mts., central part, 3680 m.,
"common weed on burnt areas", Brass 4225 (NY); Mt. Wilhelm, on open places above
the tree-line, 11000 to 15000 ft., 5 Aug. 1953, Semple & Rayner (MEL). Other speci-
mens reported, but not examined this author: Southeastern New Guinea, Crest o
Owen Stanley Range, 1889, MacGregor (MEL, holotype of Senecio erechthitoides F.
Muell., too fragmentary now to loan) ; Northeastern New Guinea, Sarawaket Mts., 2 March
1937, Clemens 5682 (cited by Mattfeld).
As far as is known from these five widely scattered collections, this species is
endemic in the mountains of eastern New Guinea at elevations above 2500 meters.
Further explorations should clarify the nature and distribution of this distinctive
outlier, which bears so little superficial resemblance to most other erechthitoid
species of Senecio. It is similar to S. laceratus in the proportionately very broad
leaf, but is readily distinguished from it by the size of the capitulum, almost twice
as large as that of S. laceratus.
[Vol. 43
54 ` ANNALS OF THE MISSOURI BOTANICAL GARDEN
9. SENECIO RUNCINIFOLIUS Willis, Proc. Roy. Soc. Queensl. 62: 106, pl. 7, figs.
34-37- 1952.
Erecbtites picridioides Sond. & Muell. Linnaea 25: 523. 1852; non Turcz. Bull. Soc. Nat.
Mosc. 24: 200. 1851.
Erechthites mixta Benth. Fl. Austral. 3: 659. 1866, pro majore parte; Black, Fl. S. Austral.
10. 1929; non (A. Rich.) DC. Prodr. 6: 297, 1858.
Herbaceous, pale green, glabrescent, 15 to 80 cm. high, young parts and leaf
axils slightly arachnoid-pubescent. Leaves sessile, lanceolate, acuminate, largest
12 cm. long and 4 cm. wide at base, with up to 8 more or less retrorse sinuate-lobed
segments on each side, uppermost leaves less toothed, with very long subfiliform
apices. Inflorescence corymbose, lax, up to 100 capitula or more, peduncles slender,
as much as twice as long as the calyculate capitulum. Involucre minutely arach-
noid, glabrous in fruit; phyllaries 12-14, about 10 mm. long, 0.5 to 0.7 mm. wide,
narrowly deltoid, with obtuse or subacute non-sphaceolate apex. Marginal florets
pistillate, about 40, corolla slenderly filiform, usually 3-fid, often with one or two
much deeper sinuses, giving a subligulate aspect, or sometimes unequally 4-fid,
lobes papillose, thickened. Disc florets about 13, hermaphroditic; corolla infundi-
buliform, tube about 0.5 mm. in diameter at base of the 5 papillose-thickened lobes;
anthers 5, about 1 mm. long, ecaudate; style arms about 0.33 mm. long, apices
enlarged and recurved, without fused terminal papillae. Pappus silky, lustrous, at
length exceeding florets by 2 mm. and phyllaries by as much as 5 mm. Achene 2.5
to 3 mm. long, about 0.3 mm. in diameter, subrostrate, 9- or 10-ribbed, with short
suberect papilliform hairs on the ribs.
AUSTRALIA, SOUTH AUSTRALIA: Moorundee near Blanchetown, Murray River, Feb.
1851, Mueller (MEL ex Hb. Sonder, holotype); “towards Spencer’s Gulf”, Warburton
(ME VICTORIA: “10 mi. west of Cohuna", Aug. 1946, Vickery (NSW); Berribee
Tank, Murray River flood plain, 31 Aug. 1948, Willis (MEL); Cohuna, 8-9-1952 (NSW
ex , 39” tall, lower leaves to 10" long). NEW SOUTH WALES: “ ; i
Hill", Fraser (K, syntype of E. mixta Bede ves of E
Fraser (BM, same gathering as preceding?); Warrego River, Western Plain: , Sept. 1885,
Betche 15 (MEL); Warrego River, 12-8-1885 (Betche ?) (NSW); junction of Murray
and Darling rivers, 1889, Mrs. Holding (NSW ex MEL); Zara, Wanganella, Dec. 1905,
Officer (NSW, US); Brindingabba, Arrara, 1912, Boorman (NSW, “eaten by stock
voraciously") ; Nelia Yari (Menindee Dist.), 20 Nov. 1947, Constable (NSW de
LJ
This is a comparatively rare plant, apparently confined to riparian habitats
mostly in the Murray River system, and to be expected in, but not yet reported
from its extensions into extreme southern Queensland. Its scarcity may be explained
by the note on Boorman's specimen.
As pointed out by Willis, this species was originally described from very depau-
perate specimens, and under a name already preoccupied. Bentham evidently
recognized the relationship between the type and the more robust specimen from
Spencer's Gulf. To these he united collections by Robert Brown and by Fraser,
and identified the whole group with E. mixta DC. His description was entirely
original and applies well to the Fraser specimens. Neither the description nor the -
specimens agree with Richard's type of S. mixtus, which belongs to the genus
Arrhenechthites. ‘The “Memory Cove, R. Brown" specimen cited by Bentham as
1956]
BELCHER—ERECHTHITOID SPECIES OF SENECIO — 55
E. mixta appears to be "Senecio plebejus 8, Memory Cove, R. Brown 2282” (BM!).
Too over-mature for accurate identification, it belongs in the vicinity of S. bispi-
dulus, with glabrous achenes. It is neither S. mix£us nor S. runcinifolius, definitely.
There should be no confusion between this species and the similarly named S.
runcinatus Less., a true discoid Senecio from Mexico, which has a large leaf up to
30 cm. long and 12 cm. wide with its margin closely set with callus-tipped teeth
and the runcinate lobes at essentially right angles, instead of reflexed. Lessing's
species was erroneously placed in Erechtites by de Candolle, but must be excluded
from that genus.
The material examined includes five of the six collections cited by Willis,
through whose kindness I was able to examine them. The 1952 specimen from
Cohuna appears to be part of a collection received by Willis after the publication
of his paper. In a personal communication he stated that it has: "robust stems
twice as tall as the limit fixed in my diagnosis", which was forty centimeters, I
have emended the diagnosis accordingly.
10. SENECIO wairauensis Belcher, nom. nov.
HE glabrescens 'T. Kirk, Trans. N. Z. Inst. 9: 550. 1877; Student's Fl. N. Z.
s e spiun, Man. N. Z. Fl. ed. 1. Se? Lond ed. 2. 1008. 1925; Illus. N, z H
- 1914; non DC. Prodr. 6: 295.
E Bes a Ts DC. var. B minor Hook. ` Fl. N. Z. 141. 1853.
Stem herbaceous, erect, sulcate, 30 to 80 cm. tall, simple or sparingly branched
above, glabrous throughout or with a few minute soft hairs in the grooves of the
peduncles; leafy. Leaves suberect, crowded and overlapping, up to 15 cm. long, 5
cm. wide, gradually reduced in size upward usually with little change in propor-
tions, sessile, often auriculate-amplexicaul or even sagittate below an attenuated
narrowly winged petiole, or rarely merely sessile without auricles, varying from
oblong-linear and obtuse to broadly oblong with a large ovate terminal lobe, sinu-
ate-dentate, sinuate-lobate, or pinnatifid with denticulate lobes, glabrous or with
soft hairs scattered on lower surface, especially along nerves, and more sparsely on
upper surface, wings of petiole often ciliolate, blade very thin and membranous
when dried, often purplish beneath. Inflorescences terminal and axillary, race-
mose in bud, becoming paniculate or subcorymbose. Capitula borne singly on
slender peduncles of irregular length, often 1 to 2 cm. long, 3 to 8 (to 14) per
branch, never solitary. Peduncles glabrous or with a few minute hairs in grooves;
two or three linear-subulate bracteoles 3 to 5 mm. long on the peduncle proper,
and one slightly longer bract subtending each peduncle. Phyllaries of involucre
(10 to) 12 to 14, 6 mm. long, 0.5 mm. wide, glabrous, shorter than florets and
Pappus, keel bearing two broad low nerves with a narrow median fissure between
them, margins scarious, apices shortly acute. Pistillate florets 10 to 15, hermaphro-
ditic florets 20 to 25. Marginal florets pistillate or occasionally with some rudi-
mentary stamens also, with style-arm apices truncate and without hairs; corolla
4-fid, filiform, 4 mm. long. Disc florets hermaphroditic, style-arm apices truncate
with a marginal fringe of a few short hairs; corolla 5-fid, slenderly infundibu-
[Vol. 43
56 ANNALS OF THE MISSOURI BOTANICAL GARDEN
liform. Achenes long-tapered, subcylindric, non-rostrate but with slightly expand-
ed annulus, faintly ribbed, glabrous or sparsely white-haired in the grooves, 4 mm.
long. Pappus very fine, capillary, white, slightly exceeding florets.
New ZEALAND: SOUTH ISLAND: Roto Iti, Kirk 824 (K, syntype and lectotype) ;
Wairau Gorge, 3000 ft., Cheeseman (K, extreme marginal florets 3-fid, pistillate, next
k Di
wh a
S. dunedinensis) ; Canterbury, Southern Alps, 2000-4000 ft., 1862, Haast 527 (K) ; Otago,
Lyall (K, very immature); Lake District, Hector % Buchanan 2 (K, very immature);
i . Benth., det. as E. prenanthoides var. B, leaves very narrow
. Hook., right specimen only, left is S. biserratus). STEWART
ISLAND: "In arenosis maritimis", Gode , immature, peduncles lanate, det. E. arguta).
Also reported by Cheeseman from a few localities on North Island.
Kirk's sketchy original description of this endemic New Zealand species was
only slightly amplified by him later. Cheeseman gave an improved description
and a good figure. This is a well-defined, valid species which, because of its trun-
cated and unappendaged style arm, is certainly a Senecio, not an Erechtites. The
new epithet is based on Kirk’s identification of the original locality of Travers.
Hooker’s diagnosis of E. prenanthoides var. minor is simply, “var. B, minor;
foliis sinuato-lobatis subpinnatifidisve". In the description of the species, further,
the leaves are, “all sharply toothed, lobed and pinnatifid throughout their length in
var. B." But the two specimens cited by Hooker for this variety, "Milford Sound
and Otago, Lyall", unmistakably are referable to this species and not to S. minimus
— E. prenanthoides DC.). Furthermore, in Hooker's annotated copy of the
"Flora of New Zealand’ in the library at Kew he has written, "Also Haast 132, 25”,
in the margin beside var. minor. This sheet, determined as var. B in Hooker’s
script, is S. wairauensis except for the right-hand specimen.
11. SENEcIo dunedinensis Belcher, nom. nov.
Erecbtites diversifolia D. Petrie, Trans. & Proc. N. Z. Inst. 19: 323-324. 1887; non Senecio
diversifolius Du Mort., 1827
slender erect herb from a perennial rootstock, unbranched or only sparingly
branched below the inflorescence, and the inflorescence more or less branched. Stem
shallowly sulcate, glabrescent or sparsely cottony, leafy for its entire length, the
lower leaves slightly crowded. Radical leaves to 3.5 cm. long, 6 mm. wide,
oblanceolate, long-attenuate to subpetiolate; cauline leaves as large as 6 cm. long,
mm. wide, linear-lanceolate to linear-oblong, shortly attenuate to sessile but not
clasping; margins remotely sinuate-denticulate on lower leaves, becoming sub-
entire on upper leaves; blades glabrescent to sparsely arachnoid beneath, nowhere
hispid. Inflorescences terminal and axillary, corymbose, with relatively few capi-
tula, at first moderately congested, becoming very lax at full maturity, peduncles
glabrous or sparsely arachnoid, with 2 to 6 subulate purple-tipped bracteoles 2 mm.
long or less. Capitula calyculate; involucre of 12 to 14 phyllaries; phyllaries 5 to
5.5 mm. long, shorter than florets and pappus, sparsely and very shortly glandular-
1956]
BELCHER—ERECHTHITOID SPECIES OF SENECIO 57
pubescent, apices rather suddenly narrowed and then prolonged in a bluntly
acuminate point, shortly ciliolate and darkened, keels basally two-ridged with a
median nerve emerging midway and becoming more prominent as the ridges are
reduced, margins scarious. Pistillate florets marginal in two rows, corolla 4-fid,
lobes thick-tipped; hermaphroditic florets central, 5-fid, lobes thick-tipped.
Achenes somewhat fusiform, slightly constricted below the expanded rim, 3 mm
long, nerves prominent, rounded, rather narrow, grooves very narrow, hairs short,
white, suberect on or beside the nerves but not in the bottom of the grooves. Pappus
very fine, slender, equaling the florets and much exceeding the phyllaries.
NEw ZEALAND. SOUTH ISLAND: Otago, Naseby, 1800 ft., Dec. 1892, W. Petrie 858
(K, det. as "Erechtites diversifolia D. Petrie, ex Herb. W. Pere. Dunedin.") ; interior of
Otago, D. Petrie (NSW, undated); Prov. Canterbury, 1860-61, Sinclair & Haast 132, 25
(K, Mo hand specimen weg others being S. wairauensis) ; Canterbury, Godley River bed,
Haast 617 (K).
SCH did not designate a type nor cite any specimens for E. diversifolia. 1 do
not find that any other author has designated a type, although several gatherings
have been listed. I did not, in limited correspondence, locate a collection by Petrie
dating from 1887 that might have served as his type. The Petrie specimen at Kew,
collected some years later, agrees with the description and could serve as neotype if
further search for the holotype should be unsuccessful.
Cheeseman? reported this species on North, South, and Stewart Islands, and
common in some places. Judging from its scanty representation in the herbaria,
it must be rather rarely collected. I have named it S. dunedinensis in allusion to
the type locality. Although apparently not very close to any other erechthitoid
species, its leaf and achene suggest S. wairauensis, from which it can be distin-
guished by its smaller capitula and shorter phyllaries.
12. SENECIO QUADRIDENTATUS Labill. Nov. Holl. Pl. Spec. 2: 48, tab 194. 1806.
Neoceis tomentosa Cass. Dict. Sci. Nat. 48: 458. 1827, nom. prov
ae ee A. Rich. Sert. oe 96. 1834, non $ prenanthoides DC.
r. 6, 1838, which is S. min
pad ras es os M pa (Labill.) DC. ode: 6: 295. 1838, et auct.
SE glabrescens DC. Prodr. 6: 295, 1838; non T. Kirk, Trans. N. Z. Inst. 9: 550.
Erechtites glandulosa DC. Prodr. 6: 295.
S landulosus Cunningh. apud Doe x Se E? IE 28:498. 1845.
Starts glabrescens (DC.) Sch. Bip. Flora 28: 498.
Erecbtites incana 'Turcz. Bull. Soc. Nat. Mosc. 24: Se
e quadridentata var. glabrescens (DC.) ub "TL Austral. 3: 660. 1866, pro
Reid erecta F. Muell. ex Lange, Bot. Tids. 4: 6. 1874.
Perennial; stems erect, striate, densely incanous-arachnoid to lanate when
young, more or less glabrate later, nowhere hispid or scabrid, sparingly or much
branched below the inflorescence, with branches suberect to erect. Leaves linear
to lanceolate and as much as 9 cm. long and 0.3 cm. wide or sometimes the lower
————
53 Man. N. Z. Fl. p. 1007. 1925.
[Vol. 43
58 ANNALS OF THE MISSOURI BOTANICAL GARDEN
ones oblong-lanceolate, sessile, attenuated toward the base, sometimes with minute
linear and simple auricles, more or less minutely and distantly callose-denticulate,
usually revolute, typically densely arachnoid to lanate especially when young, some-
times becoming subglabrous especially on upper side, acute to subacuminate. Inflo-
rescences terminal and axillary, forming corymbose cymes, slightly congested be-
coming lax, usually densely arachnoid, sometimes glabrate; capitula calyculate with
linear bracteoles about 2 mm. long; phyllaries of the involucre 11 to 13, 6.5 to 8
mm. long, 0.4 to 0.5 mm. wide, at first arachnoid later glabrate towards the apices,
2-nerved with nerves prominent on the lower third only, acuminate, ciliolate.
Marginal florets pistillate with filiform (3-) 4-fid corolla 5.5 to 6 mm. long. Disc
florets hermaphroditic with slenderly infundibuliform 4- (5-) fid corolla with blunt
lobes papillose-thickened apically. Achenes 2.5 to 5 (to 4) mm. long, straight or
slightly arcuate, more or less attenuate-rostrate, narrow grooves between broadly
flattened ribs beset with short white subappressed hairs, reddish or olive-brown at
maturity. Pappus multiseriate, niveous, exceeding the phyllaries by 2 mm., equal-
ling the florets.
AUSTRALIA. TASMANIA: "Nova Hollandia in capite Van-Dieman", Labillardiére (FI,
een", 13 1878, Beccari (Fl); Blackman's Bay, Feb. 1929, Rodway H506 )
"Between Nat’l. Park and Westerway in railway cutting", 30 N 7
TH WALES: ckson, Gaudicbaud 4 (P, holotype of S. prenantboides
A. Rich.; G. Prodr.) ; Barham, 13 Oct. 1949, Vickery : :
Vale, 17 ‘Nov: 1843, Roberiton 691 (K); Werribee, 24 RS 2, Morrison (K); 14 Jan.
89
1924, Williamson (F, center and right; left is S. glom d Melbourne, Tooroorong, 8
Nov. 1936, Mauritzon (S). SOUTH AUSTRALIA; “Austr. felix, Exp. Novara", Mueller
(W); js cin meridional.", Mueller (Fl, W); Mt. Lofty ranges, Sept. 1903, Koch
$37 (K). TERN AUSTRA LIA: Swan River, Drummond 379 (FI; K, 4 sheets; W; iso-
types vd En incana EL urcz.); dw idis Hugel (W); Toodyay, Preiss 73, 126 (S).
NORTH ISLAND Sait? Ps Woodhill, Kapara, Oct. 1882
Geer US). OUTH ISLAND: Can EPA Waiau Marble Quarry, limestone cit
Dec. 1936, Lotbian (K); Prov. Ca ee 860-61, Sinclair & Haast 321 (W); Awatere,
Kirk 17 (US); Central Otago, Alexandra, 15 Nov. 1929, Sledge 391 (K).
IMOR: Decaisne (P).
This was the first of the erecthitoid species of Senecio to be published. The type
material, including the holotype sheet which has pinned to it the manuscript draft
of the original description plus four specimens, reveals much of the range of varia-
tion in habit, leaf shape, and pubescence of this species. Subsequent collections
have added to the known variation, and several extremes have received specific
rank. There has been apparent introgression with allied species, especially S.
hispidulus, and some such states have also been described as species, It is doubtful
if a really satisfactory disposal of all the names in this complex can be made without
intensive experimental investigation. |
Richard redescribed S. quadridentatus Labill., making the corolla lobes non-
glandular. He then described S. prenanthoides to contrast sharply with his S.
bispidulus. But both his specimens of the former two have corolla lobes glandu-
1956]
BELCHER——ERECHTHITOID SPECIES OF SENECIO 59
lose-papillose, and are in full agreement except for pubescence. The inflorescence
of Gaudichaud 4, the holotype of S. prenanthoides, is virtually glabrous, although
what pubescence there is, particularly on the phyllaries, is arachnoid, as on Labil-
lardiére’s types. These extremes of pubescence are unite by specimens showing
all intermediate degrees, and by some which show one extreme on one branch and
the other on another. I can not justify retaining S. prenanthoides even as a form.
The type of E. glabrescens DC., Cunningham 134 (G!), is quite comparable to
Gaudichaud 4 in pubescence but has less revolute leaves. Broader subpetiolate
basal leaves also give this and similar specimens a somewhat different aspect,
although such leaves probably develop normally in this species but are usually lost
before maturity. They certainly were present on seedlings raised from Tasmanian
achenes furnished me by Dr. Curtis. This specimen is identical in floral details
with the type of S. quadridentatus.
The variety glabrescens Benth., defined by specimens determined by him at
Kew, included not only the specimens of Cunningham 61 which accurately repre-
sent E. glabrescens DC., but also such diverse elements as Adamson 343 and Gunn
508 (Circular Head), which appear to be hybrids between S. quadridentatus and
S. hispidulus A. Rich. Bentham regarded E. glandulosa DC. as a luxuriant state of
E. quadridentata. The type, Cunningham 141 (G!), is definitely related to S.
quadridentatus by achene, leaf shape, and lanate-arachnoid pubescence, but differs
in being less pubescent and in having larger and auriculate leaves. I agree with
Bentham, although the auricles may be evidence of hybridity with S. hispidulus.
Erechtites erecta Mueller ex Lange has never been indexed as published, prob-
ably because the name occurred in a discussion of the affinities of E. Muelleri, writ-
ten in Danish. I am indebted to Mr. W. C. Worsdell at Kew for a translation of
Lange’s article, from which I quote the passages pertaining to E. erecta:
"This second name of Müller's (E. erecta) is ascribed to a plant which differs from
that kee? — Ge Mueller E denser and longer, mei loose f over oi entire € (also
iff and earl eaves,
on the upper s f the leaves), by st erect, rly adpressed, narrow linea
with sug EN eed d margins and almost dese of a, E. more rigid ye sd more
congested on and a distinct ee - the achene. This E. erecta F. Mill., sched. pl.
exsic aer in ]l.) agrees fairly closely with the original S. quadri-
(no Bth. d :
dogs Labill. fl. nov. po ae 194, and with the ‘description of the same in DC, per-
haps answers to the nii Bth. & Müll., mpa pS vini Se ok very v": white,
leaves mostly fuh, Pon entire SE a few r eeth’. must be n i
most leaves on the Müller specimens are not stalked, ut San bad: iker. they are not ‘oblong’
linear .
ut just narrowly
I consider this valid publication.
I have not seen Lange’s type, but have seen a number of sheets determined as
E. erecta, some by Mueller himself. These specimens bear out Lange’s description,
except that the achenes vary some in the attenuation of the neck. The differences
between these specimens and typical S. quadridentatus are in the direction of greater
xeromorphism, and probably are an ecological adptation to the more arid conditions
of South Australia, from which most of the authentic material has come. Lange’s
tentative identification, with var. Gunnii Benth. (= E. Gunnii Hk. f.) is of course
[Vol. 43
60 ANNALS OF THE MISSOURI BOTANICAL GARDEN
not supported by comparison with Hooker: type. I regard this simply as an
extreme of S. quadridentatus, lacking sufficient discontinuity from the type to be
recognized.
13. Senecio gunnii (Hook. f.) Belcher, comb. nov.
Erechtites Gunnii Hook. f. Lond. Jour. Bot. 6: 122. 1847; Fl. Tasm. 1: 220, tab 63. 1860.
Erecbtbites quadridentata var. Gunnii (Hk. £.) Benth. Fl. Austral. 3: 660. 1866.
Perennial from rhizome, annual shoots herbaceous or slightly woody at base.
Stems erect, arachnoid to lanate but eventually glabrate. Leaves sparsely arachnoid
and hispidulous above, moderately lanate and hispid beneath; lower ones obtusely
oblanceolate or obovate-lanceolate, 8 (to 12) cm. long, 1.5 (to 1.8) cm. wide,
subentire but minutely denticulate, base long-attenuate and subpetiolate, slightly
broadened at attachment and sometimes with small auricles 1 to 2 mm. long;
upper ones smaller, sessile, not attenuate. Inflorescences a corymbose-panicle,
varying from rather compact to lax, branches lanate to arachnoid or eventually
glabrate, capitula several to numerous on rather short peduncles, calyculi lanate;
phyllaries 11 to 13, 6 mm. long, glabrous, minutely glandular-papillose, or basally
sparsely arachnoid, more or less prominently keeled, 2-nerved, often purplish, apices
acuminate, ciliolate; marginal florets pistillate, corolla filiform, apices of the 4
lobes thickened-glandulose. Disc florets hermaphroditic, corolla subfiliform,
slightly and abruptly dilated above, apices of the 5 lobes thickened-glandulose;
anther bases obtusely lobulate. Achenes 3 mm. long, slightly arcuate, light brown,
strongly ribbed, with short white hairs in deep grooves between the ribs or lacking,
apex definitely attenuate-rostrate. Pappus slightly exceeding phyllaries and flo-
rets, white.
AUSTRALIA. TASMANIA: Marlboro, Jan. 1841, Gunn 700/1842 (K, syntype; NSW);
Arthur’s Lakes, 17 Jan. 1843, Gunn 700 (K, syntype; NSW); 1844, Gunn 700 (K ex Hb.
Benth.) ; Western Mountains, 18 Feb. 1843, Gunn 700
Lake on
Bronte Road, 27 Jan. 1949, Burbidge 3426 (HO). victoria: Yarra Yarra, Mueller (K); (?)
Mt. St. Bernard, Jan. 1899, Walker (NSW); Mt. Hotham, Jan. 1900, Maiden (NSW);
Buffalo Mt., 4300 ft., 19 Jan. 1913, Cambage 3753 (NSW). NEW souTH WALES: Watcha
Road, Dec. 1893, Kretschmann (NSW); Mt. Kosciusko, Jan. 1898, Maiden (NSW);
Thredbo River, Jan. 1899, Maiden es Forsyth (NSW); Bemberi Peak, 6100 ft., 15 Jan.
1912, Cambage 3441 (NSW); Kosciusko, 9 March 1949 Skottsberg (NSW, S); near
Charlotte Pass, 14 March 1949, Skottsberg 174 (S).
In the original publication of Erechtites Gunnii, Hooker merely stated: “Hab.
Alpine situations, Gunn.” In the ‘Flora Tasmaniae’ he cited Gunn YOO; and" ...
Arthur's Lakes, Marlborough, etc. Lawrence, Gunn.” One sheet of Gunn 700 at
Kew has two gatherings, "Gunn 700, Arthur’s Lakes, 17/1/45, V. D. Land", and
"Gunn 700/1842, Marlboro, 5/1/41." Beside the former are the diagnostic figures
drawn for plate 63 in the ‘Flora Tasmaniae”. Below the latter specimen is “Erech-
lites Gunnii" in Hooker's script. I interpret these gatherings as the syntypes. This
same sheet has been annotated by Bentham as the variety.
1956]
BELCHER—ERECHTHITOID SPECIES OF SENECIO 61
The systematic status of S. gunnii is still somewhat uncertain. It is surely
closely allied to S. quadridentatus. That it may be a hybrid between, say, S.
minimus and S. quadridentatus, with perhaps some introgression into S. bispidulus,
can not be ruled out entirely without cytogenetic study. The principal argument
against its hybridity is its lack of variability in those features which intergrade in
the group of putative hybrids between S. bispidulus and S. quadridentatus. On the
other hand, Bentham may have been right in considering it a variety of S. quad-
ridentatus. Pending further clarification, I side with Rodway?* in considering it
a species.
From the elevations given by the collectors, S. gunnii appears to be alpine and
subalpine in distribution. The only other ecological datum available to me is that
of Gunn, preserved in the type cover at Kew: “Very common on the summits of
the Western Mountains, growing in moist places in the pasturage.”
14. SENECIO GLOMERATUS Desf. ex Poiret in Lam. Encyc. Suppl. 5: 130. 1817.
Senecio glomeratus Desf. Cat. Hort. Paris. 124. 1815, nomen nudum; ex Link, Enum. Hort.
erol. 2 :
Erechtites glomerata and vars. subincisa and polycephala DC. Prodr. 6: 297, 1838.
Neoceis microcephala Cass. Dict. Sci. Nat. 34: 388. 1825
Senecio argutus A. Rich. Fl. Nouv. Zél. 258, 1832; Sert. Astrolabe, 104. 1834; Endl.
Prod. Fl. Norf. 51. 1833; non Kunth in HBK. Nov. Gen. & Spec. 4: 183, 1820.
Erechtites arguta (A. Rich.) DC. Prodr. 6: 296. 1838; Cheesm. Man. N. Z. Fl. 364. 1906;
ibid. ed. 2, 1007. 1925; Hook. f. Fl. N. Zeal. 1: 142. 1853, pro parte; Fl. Tasm. 1:
219. 1860, pro parte; Benth. Fl. Austral. 3: 659. 1866, pro parte; Rodway, Tasm. Fl.
95. 1903, pro parte; Black, Fl. S. Austral. 4: 610. 1929, pro parte.
Senecio Lessonianus Sch. Bip. Flora 28: 498. 1845; non Steud. Nom. ed. 2, 2: 562. 1841.
Senecio plebeius Banks & Soland. ex Hook. f. Fl. N. Z. 1: 142. 1853, ut syn., nom. nud.
Senecio Lessoni F. Muell. Ann. Rept. Gov't. Bot. 26. 1858, mom. nud. (publ, as nom. nov.
for “E. an , error for E. arguta?).
Annual (?) ; stem erect, herbaceous or slightly woody at base, simple or branch-
ing above, more or less densely arachnoid-pubescent, sometimes glabrate below,
often sublanate on juvenile parts and inflorescences. Leaves ovate-lanceolate to
linear-lanceolate, variable in length and width and in length-width ratio but usual-
ly more than three times as long as wide, denticulate and revolute to irregularly and
coarsely sharp-toothed to sinuate-lobate with obtuse sinuses and callose-denticulate
deltoid lobes to more or less profoundly pinnatifid with long lanceolate denticulate
or subincised acute or obtuse lobes, sessile, usually auriculate; lower leaves some-
times subpetiolate above the auricles; sparsely arachnoid to glabrate above, more
densely arachnoid beneath. Inflorescences terminal and axillary, forming a con-
gested corymbose panicle, arachnoid to lanate, rarely glabrate after anthesis, capitula
small, 5 to 7 mm. long, 2 to 3 mm. wide, borne singly or in small clusters, calycu-
late; phyllaries 11 to 13, 3.5 to 5 mm. long, usually near 4 mm., with basal one-
third to one-half usually densely short-arachnoid to lanate, or rarely glabrate after
fructescence, obscurely 2- to 3-nerved, acute to acuminate, minutely ciliolate at
the apex. Marginal florets pistillate, occasionally with rudimentary stamens, with
corolla filiform, usually 3-fid, sometimes irregularly 2-fid and subligulate. Disc
—
9^ Tasmanian Flora. pp. 94-95. 1905.
[Vol. 43
62 ANNALS OF THE MISSOURI BOTANICAL GARDEN
florets hermaphroditic, with slenderly infundibuliform corolla 4-, 5-fid, and with
style-arm apices truncated. Achene short-subcylindric, 1.5 mm. long or less, not
attenuate-rostrate, ribbed, with short whitish or tawny subappressed hairs between
the ribs. Pappus white, capillary, exceeding involucre, subequalling florets.
CULTIVATED: "Senecio quadridentatus Labill. Senecio glomeratus h. par." (FI, Hb.
Webb. ex Hb. Desf., 2 pieces, right-hand one designated as neotype); "Senecio aggregatus
hort. Paris., 1813" (P); “Jardin des Plantes . . . October 1816” (K ex Hb. Gay, det. as S.
glomeratus Poir.) ; "Senecio glomeratus h. pl. 1815” (G Prodr., syntype of E. glomerata
var. subincisa DC.).
Bridgeport, April 1946, Wilson (HO); Somerset, Feb. 1948, Curtis (HO). WESTERN
AUSTRALIA: "Riv. des Cygnes", iter Baudin (P); Swan River, 1839, Drummond
Drummond 257 (W). SOUTH AUSTRALIA: “Aust i
K:
( ra.
, 2, marginal florets frequently more or less ligulate, next row sometimes perfect
ue
ne, Nov. 1936, Mauritzon (S); Sand-
ringham, Nov. 1936, Meebold 21799 (NSW). NEW souTH waLEs: Mt. Koscuisko,
(K).
NEw ZEALAND. NORTH ISLAND: “Ad riv. Punakitere, Nov. 1874", Berggren (S);
Auckland, Panmore, Kirk (F); Auckland, Cheeseman (F); Wellington, "
sea to 2000 ft., common," March 1909, Travers (G Deless., W)
de l'Astrolabe, détroit de Cook" (P, holotype of S. argutus A. Rich.); Lake Dist. in the
bush, March, Hector & Buchanan 3 (K). WITHOUT SPECIFIC LOCAL : "Seneci bius"
cino County, July 1931, Jones 29097 (BM, F); San Mateo Co., open rocky hills, Sharp
cum es Rose 13980 (W); Humboldt County, Spruce Cove, Trinidad, Parks 24033
» 9):
Poiret’s description is rather difficult to apply literally to the herbarium mate-
rial to which his epithet has been applied, the disagreement being mainly with leaf
shape and pubescence. The name was based on a living plant, and there might
be no holotype. I have, however, obtained from Florence a sheet with two speci-
mens from Desfontaines’ herbarium, which is ticketed “Senecio quadridentatus
Labillard. Compos. h. p., S. glomeratus h. Par." The latter determination is in
the same script and ink used on the holotype of S. minimus Poir., and is, I believe,
in Poiret’s writing. I therefore select the right-hand specimen on this sheet as
neotype of Senecio glomeratus Desf. ex Poir.
The two specimens have somewhat different leaf forms. The neotype approxi-
mates Poiret’s description of the leaf as linear-lanceolate, incised or toothed, but
clearly shows a few pinnatifid leaves near the base. The specimen on the left
accords well with the description of S. glomeratus Desf. ex Link as petiolate and
pinnatifid. It also agrees with a number of specimens bearing determinations as _
1956]
BELCHER
ERECHTHITOID SPECIES OF SENECIO 63
S. glomeratus Desf., most of which, like the one in the Prodromus Herbarium, were
taken from cultivation. These two specimens are in excellent agreement in floral
characters; both were included in the determination, and I see no basis for separat-
ing them. Nor could I justify forming subspecific categories of this variable
species on such an unstable basis as these foliar differences, when every degree of
intergradation exists in the material.
Richard based S. argutus on a scrappy specimen, little more than an inflores-
cence, from New Zealand. The floral characters of this specimen, and of a some-
what larger pair of branches from Richard’s herbarium, ticketed simply as “Senecio
argutus nob., Nie Zélande", are unmistakably those of S. glomeratus. The same is
true for the sheet of six fragmentary specimens collected by the naturalists of the
Baudin expedition from “Nouv. Holl. détr. d'Entrecasteaux” (Tasmania), which
Richard determined as Senecio argutus. He rightly cited these as extending the
range of this species to Australia.
The d'Urville specimen cited by de Candolle in the ‘Prodromus’, in contrast to
«the one retained by Richard, has quite pinnatifid leaves but agrees with Richard's
type and description in pubescence and floral features. This New Zealand speci-
men forms the basis for “Erechtites arguta . . . Senecio argutus A. Rich. et
Lesson. . . . non Kunth” in the ‘Prodromus. De Candolle also maintained E.
glomerata as a distinct species, and even placed it in a different section of the
genus! The holotype of his typical variety is a specimen "ex itin. Baudin", which
is almost an exact replica of the left-hand specimen on the Baudin sheet cited by
Richard as S. argutus. These two specimens likely are of the same gathering,
possibly even from the same plant!
With this merely sinuate-dentate specimen de Candolle associated two coarsely
lobate to pinnatifid specimens as E. glomerata var. subincisa DC. This was based
on "Senecio glomeratus Desf.! hort. Paris 1824 [sic!]. Poir. Suppl. 5. p. 130.
Link enum, 2. p. 325. Neoceis microcephala Cass. dict. 34. p. 388. (v. v. et KI
These specimens approach the left-hand specimen from Desfontaines' herbarium,
but do not quite equal it in depth of lobing and slenderness of lobes. One is
ticketed as "Senecio glomeratus h. pl. 1815", and is probably the basis for the
exclamation mark after the basonym, since with de Candolle that meant that he
had seen the "type". The other specimen establishes the earliest date, 1807, which
I find for this species in cultivation.
The type of E. glomerata var. polycephala, from Port Western on Bass Strait,
has a somewhat larger capitulum with larger and wider bracts than usual, and three
large leaves which are merely shallowly dentate. Although I have seen no other
specimen which exactly duplicates this, it is definitely a part of the S. glomeratus
complex. It is linked to the more typical state by several collections, notably the
Travers specimen from Wellington, on which the phyllaries were 5 mm. long and
the leaves similarly large. I therefore regard var. polycepbala as only a luxuriant
state of S. glomeratus.
[Vol. 43
64 ANNALS OF THE MISSOURI BOTANICAL GARDEN
J. D. Hooker disregarded E. glomerata and took up E. arguta in his floristic
papers on Australasia. In attempting to deal with the extreme polymorphism
shown by the (hybrid) material in his herbarium,® he created four varieties of E.
arguta for Tasmanian specimens, and later formed two varieties for New Zealand
specimens. These specimens are at Kew, duly annotated.
Hooker: var. a of New Zealand is typified by Akaroa, Raoul 31 (K), which
I regard as a hybrid between S. glomeratus and S. hispidulus (probably var. scaber-
ulus). His var. 8 of New Zealand agrees well enough with typical S. glomeratus. His
var. B glabrata of Tasmania is typified by two mixed sheets at Kew, of which that
part which most closely matches the brief description again appears to be another
hybrid of the above parentage, while the other specimens are more like pure S.
glomeratus. Hooker attributed his var. y asper to S. asper Cunn. I have failed to
find this name published, and if it were it would be a later homonym of S. asper
Ait. This variety is typified by Hooker 1125 and by “near Woolworth, 25 Nov.
1836, Gunn 843”. These were compared with Richard's types, and agreed most
closely with S. pusillus A. Rich., which I consider a depauperate state of another
hybrid between S. glomeratus and S. bispidulus. Var 8 obovata is good S. glomera-
643/1842", is much the same type of plant as the hybrid var. a of New Zealand.
tus. Var. e, without an epithet, typified by “Circular Head, 11 Dec. 1837, Gunn
Thus E. arguta Hk. f. includes both S. glomeratus and some hybrids of it with S.
bispidulus.
Bentham consolidated these errors by accepting as F. bispidula 3 mixed group
of large-headed specimens including S. squarrosus, and by placing the then nameless
specimens of S. bispidulus, along with hybrids of it with S. glomeratus, in with the
true S. glomeratus material as E. arguta (sensu Bentbami). A large part of Ben-
tham's description of E. arguta applies much more to S. bispidulus than to S.
glomeratus, and some parts apply only to the former and its hybrids.
Of modern authors, only Cheeseman seems to have had an accurate conception
of this taxon, under the name of E. arguta DC. Neither his key to species nor
his description reveals any confusion with S. bispidulus. Rodway followed Bent-
ham exactly. Black similarly followed Bentham, even to the inclusion of his var.
microcepbala, which is better treated as a distinct species (see S. laticostatus).
15. SENECIO laticostatus Belcher, nom. nov.
Erechthites arguta var. microcephala Benth. Fl. Austral. 3: 659.
microcephalus Phil. 1894.
Stem striate, arachnoid, apparently simple below the inflorescence, basal part
unknown. Leaves to 6 cm. long, 2.5 cm. wide, pinnately lobed, lobes triangular
to oblong, sharply toothed, base auriculate, semiamplexicaul especially upward,
sparsely hispid and arachnoid and rough above, copiously arachnoid beneath. Inflo-
rescence a compact corymbose panicle of numerous capitula, few or solitary on
1866; non Senecio
35 Lond. Jour. Bot. 6:122. 1847.
36 Bot, Antarct. Voy. II. Fl. N. Z. 1:142. 1853.
1956]
peduncles from 0.5 to 5 cm. long, branches striate, arachnoid, peduncles strongly
striate, moderately arachnoid, the linear-lanceolate entire bracts virtually glabrous,
receptacles lanate. Capitula short, comparatively broad; phyllaries of involucre
about 12 to 13, slightly lanate at the base, glabrous at the apex, 3.5 to 4 mm. long,
0.5 to 0.75 mm. wide at base, the center portion about 0.1 to 0.2 mm. wide between
the scarious margins which rapidly taper toward the apex so that the apical 0.75
to 1.0 mm. is essentially without scarious margins and is long and slenderly acumin-
ate, even in early bud, when the phyllaries may equal or exceed 3 mm. while the
florets are still less than 1 mm. long. Marginal florets pistillate, 20 to 25, corolla
slender, 3- or 4-fid, 2.5 to 2.75 mm. long. Disc florets hermaphroditic, 3 or 4 in
number, corolla 3.0 to 3.25 mm. long, thick, apex 1.5 mm. in circumference, with
4 obtusely deltoid lobes, each 0.4 mm. wide and 0.3 mm. long, anthers 1 mm. long;
florets at anthesis exceeding phyllaries by about 1 mm. Achene (somewhat im-
mature) olive-green, 1.5 mm. long, cylindric, non-attenuate, glabrous, having 5
extremely narrow thin high ridges extending its full length. Pappus hairs white,
unequal in length but mostly equaling or slightly exceeding the phyllaries, shorter
than the florets.
Known to me only from the holotype collection.
AUSTRALIA: Flats beyond the Brodribb River, Jan. 1855, Mueller (MEL holotype).
Mueller gave to the type specimen a manuscript name in Erechtites; Bentham
adopted his trivial as the varietal epithet. The new epithet, Jaticostatus, alludes
to the unique fin-like ribs of the achene. The type specimen has a somewhat
anomalous growth pattern, as though it had been broken over prior to flowering.
The leaves are highly suggestive of the more coarsely lobate forms of S. glomeratus,
differing unreliably in having somewhat coarser denticulations on the median lobes.
The floral features, however, are unique, particularly the short broad 4-fid corolla
of the few disc florets, with bluntly deltoid lobes, the short phyllaries with acumin-
ate non-scarious apices, and the five-finned short achene. Furthermore, the ped-
uncles of S. glomeratus are shorter and more lanate than those of S. laticostatus, and
the involucre of the former is much more pubescent than that of the latter.
It is possible that the type specimen is a hybrid between S. glomeratus and some
discoid species of Senecio, in some of which there is an approach to this type of
corolla, But I have yet to encounter any species which might have contributed
the unique features of the phyllaries and the achene. Furthermore, the pollen
grains are well formed, lacking the abortive grains one would expect from a wide
cross. I have seen no other specimen of this taxon, which in itself is suspicious,
but I see no alternative to treating it as a good species, perhaps related to but
certainly distinct from S. glomeratus.
16. SENECIO HisPIDULUS A. Rich. Sert. Astrolabe, 92, fab 34. 1832.
ual herb; stem erect, simple or sparingly branched below the inflorescence,
sometimes subscapose, glabrous or minutely puberulous or sometimes densely clo
with crisped multicellular hairs. Leaves 3 to 7 (to 11) cm. long, 0.5 to 1 cm.
[Vol. 43
66 ANNALS OF THE MISSOURI BOTANICAL GARDEN
wide (to 2.5 cm. or more in vars. dissectus and scaberulus), linear-lanceolate to
ovate-lanceolate, sessile, auriculate with coarsely bidentate auricles, or the lowest
leaves sometimes subpetiolate with minute linear auricles, more or less coarsely and
irregularly sharp-toothed, sometimes deeply pinnatisect and the segments sharply
denticulate, densely hispid beneath with multicellular hairs, less densely to sparsely
hispid above or sometimes glabrate, upper surface frequently roughly wrinkled or
scabrid after drying, with the hairs subtuberculate. Inflorescences terminal and
axillary, corymbose, usually rather congested, sometimes diffusely paniculate, glab-
rous, minutely puberulous, or rarely hispid; bracts greatly reduced, sublinear or
long-triangular, subentire, erect. Capitula slender, calyculate with triangular
bracteoles about 1.5 mm. long and 0.4 mm. wide; phyllaries of involucre 11 to 13,
4.5 to 6 mm. long, glabrous or sparsely puberulous, not arachnoid, strongly keeled
and prominently 2-nerved, scarious on the margins, acute or acuminate; corollas
of the filiform pistillate marginal florets usually 4-fid, a few 3-fid or 5 fid, of the
less numerous slenderly infundibuliform perfect disc florets 5-fid, the lobes all
obtusely papillose-thickened on the inner aspect of the apex, the style-arm apices
truncated, Achene 1.5 to 2 mm. long, plump, cylindrical, without attenuate neck
but with callose-annulate apex, blackish-brown with whitish subappressed hairs in
rows of variable width between the low rounded ribs. Pappus white, exceeding
phyllaries, subequalling the florets.
In temperate Australia and New Zealand.
16a. SENECIO HISPIDULUs var. hispidulus
Senecio bispidulus A. Rich.
Erecbtites bispidula (A. Rich.) DC. Prodr. 6: 296. 1838; non Hook. f. Fl. Tasm. 1: 220.
1860; non Benth. Fl. Austral. 3: 660. 1866; non Rodway, Tasm. Fl. 94-95. 1903; non
Black, Fl. S. Austral. 610. 1929.
Erechthites arguta Benth. Fl. Austral. 3: 659. 1866, pro majore parte.
Leaves linear-lanceolate to lanceolate, denticulate to coarsely toothed, neither
pinnatifid nor lobed, upper leaf surfaces scabrid with subtuberculate-based hairs.
Achenes 1.5 to 1.75 mm. long, beset with hairs in narrow rows in the grooves.
AUSTRALIA. TASMANIA: “Senecio hispidulus No. 18. Van Diemen, 1828, Voy. Astro-
labe" (P, holotype) ; Port Dalrymple, R. Brown 2278 (K); between National Park
Westerway, in railway cutting, Nov. 1929, Comber 1756 (HO, 2; K); Knocklofty, Ho-
t ee
one
pinnatifid, approaching var. dissectus) ; Boyamp, Oct. 1896, Morrison (K); Midland Junc-
tion, Dec. 1898, Morrison (US). NEW SOUTH WALES: Sydney, U. S. Expl. Exped. (US);
Dec. 1902, Camfield (US, left-hand specimen only, right a hybrid with S. quadriden-
fatus) ; Kiama, Harvey (K).
NEW ZEALAND. NORTH ISLAND: Coromandel, Petrie (K, 2). SOUTH ISLAND: without
specific data, ex Hb. T. Kirk (F). Bay of Islands, U. S. Expl. Exped. (US, aspect of var.
bispidulus, but only 8 phyllaries, depauperate, leaf bases non-auriculate, leaves oblong or
ovate-lanceolate).
1956]
BELCHER—ERECHTHITOID SPECIES OF SENECIO 67
Senecio bispidulus is a species whose identity in Australian floras has been
thoroughly submerged in "E. arguta”, while other taxa have masqueraded under
its name as “E. hispidula”.
Richard's description and figure of S. hispidulus, on the whole, agree well with
the holotype, but erred slightly. The marginal florets are pistillate, not staminate,
as pointed out by de Candolle, and are more usually 4-fid than 3-fid. The artist
portrayed the leaves as entire above the bidentate auricles; actually they are rather
regularly callose-dentate, as I found by use of detergent solution, but are so strongly
rolled as to appear entire. De Candolle had no specimen of this taxon, but drew
the diagnosis for the ‘Prodromus’ directly from Richard, merely correcting the
description of the pistillate floret.
Hooker, however, had the misfortune to receive from Gunn, and later from
other collectors on Tasmania, a curious assortment including hybrids of S. bispidulus
with both S. glomeratus and S. quadridentatus. The former group he referred to “E.
arguta”, and attempted to treat them as varieties, as discussed above under S.
glomeratus. The latter group he first distributed, in the "Florae Tasmaniae Spici-
legium’, as unnamed varieties of E. hispidula and E. glabrescens (i. e., a glabrate
state of S. quadridentatus). Both varieties were queried as to whether they might
belong to the other species!
In properly assigning Cunningham's, d'Uberville's, and his own specimens from
New Zealand to E. bispidula in the ‘Flora Novae Zelandiae', Hooker compared them
to this mixed Tasmanian group: "This is also a New Holland and Tasmanian plant,
but the specimens from the latter country are wooly as well as hispid. Those figured
by Richard have the leaves nearly entire." This concept is epitomized by the Kew
specimen, “Circular Head, 23/11/37, Gunn 508”, which is annotated “E. glabres-
cens DC.”, and beneath that, “E. hispidula DC.”, both in Hooker's script. (Ben-
tham rejected this determination, and cited this specimen in the "Flora Austral-
iensis’ as E. quadridentata var. glabrescens, which it is not, either.) This speci-
men has the combined pubescence of S. bispidulus and of S. quadridentatus (hence,
"Wooly as well as hispid") ; the longer phyllaries and long and attenuate-rostrate
achene of the latter; and the bidentate auricles and callose-dentate leaf margin of
the former.
In the ‘Flora Tasmaniae’ Hooker merged his varieties of E. glabrescens and E.
hispidula into what must be called E. bispidula Hk. f., non DC., and rewrote the
description to fit the Tasmanian plants in his herbarium. To the Circular Head
specimen he added others, which, as far as I could find at Kew, did not include
a single specimen of typical E. bispidulus but did include the Launcestown gather-
ing of Gunn 508, which definitely is S. squarrosus. Hooker accurately compared
his New Zealand material with this Tasmanian mixture in the ‘Handbook of the
Flora of New Zealand’: “This differs from the Tasmanian E. hispidula in the
achene only 1/12 in. long and not attenuate at the top, also in the smaller glabrous
ds and short involucral scales.” He thus correctly concluded that the two
groups were not conspecific, and set the New Zealand material apart as a new
[Vol. 43:
68 ANNALS OF THE MISSOURI BOTANICAL GARDEN
species, E. scaberula! Actually, by direct comparison of types, E. scaberula and S.
bispidulus A. Rich. are very similar and can only with some difficulty be separated
into varieties.
A careful examination of the specimens determined as E. bispidula by Bentham
for the ‘Flora Australiensis’ reveals that the misinterpretations initiated by Hooker
were only further entrenched. I believe that all of the specimens now remaining
at Kew which were determined by Bentham as E. hispidula are in fact not that
species at all. “Gunn 508 Circular Head,” he returned to E. quadridentata as var.
glabrescens. To “Gunn 508 Launcestown” and other material of S. squarrosus he
added other large-headed specimens collected by Backhouse and by Robertson, and
called the lot E. hispidula (Benth., non DC.,). His description largely applies to
S. squarrosus, which he included in the synonymy. He also included S. bispidulus,
but wrote: “from the description, but scarcely the figure, t. 34.” As noted above,
the plate actually is in good agreement with both specimen and description. `
To find S. bispidulus of Richard in the ‘Flora Australiensis’ one must look
under E. arguta, which is described as: ‘more or less scabrous-hirsute with crisped
hairs and occasionally with white cottony wool on the underside of the leaves and
about the inflorescences, rarely nearly or even quite glabrous," and "involucre in
the normal form about 3 lines long". Most of this applies to S. bispidulus rather
than to S. glomeratus, and the second quotation quite excludes S. glomeratus! In
effect, Bentham treated the S. bispidulus and the S. argutus of Richard as syn-
onyms, although he did not realize it, having already mistakenly identified the
former with S. squarrosus. j
This confusion in E. arguta Benth. becomes even more obvious when Bentham’s
synonymy and comment are considered. E. Bathurstiana DC., treated as a variety of
E. arguta, is definitely related to S. bispidulus instead. S. multicaulis A. Rich.
is, by examination of the type, unquestionably part of the S. bispidulus complex, `
with phyllaries 6 mm. long and entirely glabrous, but with the achenes also glab-.
rous instead of hairy. S. apargiaefolius Walp. is most likely a hybrid of S. bispidu-
lus with S. quadridentatus (see below). S. pusillus A. Rich, is, from the type,
definitely a hybrid between S. glomeratus and S. hispidulus, with the characters of
the latter predominating, and is essentially equivalent to E. arguta var. asper Hk. f.
The non-existent S. pumilus Poir. (E. pumila DC.) is an error on de Candolle’s
part for S. minimus Poir.
Rodway followed Bentham closely in his treatment of Erechtites. He char-
acterized E. hispidula as having 16 to 24 phyllaries about 4 lines long, the capitula
few and stout. This appears to be S. squarrosus. His E. arguta, with phyllaries
3 lines long or less, is certainly meant to include S. his pidulus as well as S. glomera-
fus. Black also included S. bispidulus in his E. arguta, but he appears to have fol-
lowed Hooker in part, rather than Bentham, in applying the name E. bispidula
to the group of specimens more resembling S. quadridentatus but with larger
slender capitula and more numerous phyllaries. E!
1956]
BELCHER—ERECHTHITOID SPECIES OF SENECIO 69
16b. SENECIO HISPIDULUS var. dissectus (Benth.) Belcher, comb. nov.
Erechthites arguta var. dissecta Benth. Fl. Austral. 3: 659. 1866.
Erecbtites Bathurstiana DC. Prodr. 6: 297. 1838.
Senecio Batburstianus (DC.) Sch. Bip. Flora 28: 498. 1845.
Differing from the typical variety in having deeply pinnatisect leaves with
obtuse sinuses, lobes 5 to 8 (to 10) on each side and broadly linear to oblanceolate,
more or less acute-denticulate; marginal florets irregularly 3-fid, with one cleft or
two much deeper, inner florets 4-fid.
AUSTRALIA. NEW SOUTH WALES: “Rocky Hills in the neighborhood of Bathurst",
ec
A. Cunningham 135 (G Prodr., holotype), A. Cunningham 102 (K, isotype?) ; "Rocky
(NSW); Burrinjuck, Feb. 1911, Boorman (NSW); Coonor's Creek, Barroba, Aug. 1913,
Rupp 6 (NSW); Bell's Paddock, Black Mountain, June 1932, McKie 454 (NSW).
VICTORIA: Wimmera, Dallachy (K, immature); Lake Albacutya, 1901, D”Alton (NSW);
3oorman, Euroa, Dec. 1901 Williamson (NSW). SOUTH AUSTRALIA: “Austr. felix",
Mueller (K).
Bentham's placement of E. Bathurstiana DC. as a variety of E. arguta Benth.
was a logical consequence of his concept of that species as virtually identical with
S. bispidulus. It has a much closer affinity with S. bispidulus proper than with S.
glomeratus, when the pubescence and general facies are considered. It is not impos-
sible that genetic analysis will reveal this variety as another hybrid, with S.
glomeratus the other parent, as is suggested by the irregular corolla of the marginal
floret and the addition of arachnoid pubescence on the leaf over the multicellular
crisped hairs. But the general characters are closer to S. hispidulus, and I leave it
in that species pending further study.
Croizat37 cited this variety (as E. arguta var. dissecta) as occurring on Lombok
in the Lesser Sunda Islands, without giving the authority for this record. In view
of the confusion over the identity of S. glomeratus and of S. hispidulus, 1 hesitate
to accept this extreme extension of range until I have examined the specimen.
16c. SENECIO HISPIDULUs var. scaberulus (Hook. f.) Belcher, comb. nov.
Erecbtites scaberula Hook. f. Handb. Fl. N. Z. 157. 1864.
Senecio bispidulus Cunn. Ann. Nat. Hist. 2: 121. 1838; non A. Rich.
Erecbtites bispidula Hook. f. Fl. N. Zeal. 142. 1853; non DC. l i
? Erechtites pumila Armst. Trans. N. Z. Inst. 13: 338. 1881, fide Kirk, Student's Fl. N.
Z. 334. 1899; non DC.
Leaves lanceolate, the lower oblanceolate and coarsely toothed or irregularly
lobed, the upper surfaces beset with coarse but non-tuberculate hairs. Achenes
2.0 mm. long, hairy in broad bands in the grooves.
I did not see Armstrong’s type, cited as “McKenzie Country, Mr. J. F. Arm-
strong, December 1877”.
37 Manual of Phytogeography. p. 538. 1952.
[Vol. 43
70 ANNALS OF THE MISSOURI BOTANICAL GARDEN
New ZEALAND. SOUTH ISLAND: Akaroa (K ex Hb. Hk., lectotype); Bay of Islands,
1853, Hooker (K, 2 sheets, syntypes) ; Dunedin, Kirk 333 (US). NORTH ISLAND: “Amon
ferns at Wangeroo", 1833, R Cunningham (K); Wellington, open places, 1500 ft., 1909,
Travers (W). Also reported by Cheeseman (l.c. 1007. 1925) from Stewart Island and the
Chatham Islands.
Neither Cunningham’s nor Hooker’s original descriptions of this taxon gave any
clue to the distinctions between it and the typical Australian variety. Hooker
described it as a distinct species only on the basis of differences between it and
what he mistakenly supposed E. hispidula to be, as described above. Since in the
description of E. scaberula Hooker cited no specimens, but referred to his treat-
ment of 1853, the specimens cited in the earlier work must be the syntypes. The
Akaroa specimen, presumably supplied by d’Urville, is selected as lectotype because
it is determined in Hooker’s script as E. scaberula. The Cunningham specimen of
this variety was not annotated by Hooker. The two sheets of Hooker’s own
gathering are included in the syntypes because they bear his determination as E.
scaberula. All are in good agreement.
Dissection of a capitulum from Hooker’s specimen revealed that all 18 florets
contained stamens, either vestigial or functional. The lobing of the corollas also
varied considerably. Most of the florets were 4-fid, some with 4 and some with 5
stamens, functional or not; two were 3-fid, one with 3 vestigial stamens, the other,
non-marginal, one with 4 functional ones; only three 5-fid florets had 5 functional
stamens each. No tendency toward ligules was discernible, the picture being
entirely one of partial emasculation accompanied by reduction in size of corolla
limb and number of lobes. This specimen thus offers an intermediate stage between
the truly discoid Senecio with all florets perfect and infundibuliform, and the
extreme erechthitoid state with all outer florets strictly pistillate and filiform. (In
some other specimens, however, which also had staminodes in 4-fid marginal flo-
rets, there were some corollas with some sinuses much more shallow than others,
thus approaching the ligulate condition.)
This variety appears to be confined to New Zealand, where it largely replaces
the typical variety. The latter is adventive in New Zealand to some extent. The
specimens from Coromandel, for example, were sent to Kew by Petrie with a note
saying that they were new to the island. The determination at Kew was: "E.
scaberula with slightly different achenes"!
SENECIO GLOMERATUS X S. HISPIDULUS
sillus A. Rich. Sert. Astrolabe, 99. 1834
recti arguta (A. Rich.) DC. var. a Hk. f. Fl. N. Z. 142, 1853 (prob. X S. bispidulus
var. d ; vars. B slebróta; y aspera, and ¢., Hk. f. Fl. Tasm. Spicil. 122. 1847;
asm. 1: 219. 1860
Senecio glomeratus and S. bispidulus appear to hybridize so freely over much
or all of their largely overlapping range that determination of material in this
complex is exceedingly difficult. This is a situation which urgently requires
investigation by as many varied techniques as can be brought to bear.
1956]
BELCHER——ERECHTHITOID SPECIES OF SENECIO 71
In my own determinations the following criteria have prevailed. For S.
glomeratus Y associate arachnoid pubescence, simple auricles, lanate receptacles,
bracts, pedicels, and phyllary bases, small capitula, short (3.5 to 4 to 5 mm.),
obscurely nerved phyllaries, usually 3-fid pistillate florets, with slightly irregular
corollas, and short (1.5 mm.) brownish achenes. For S. bispidulus 1 associate
hispid multicellular pubescence, with the bases of the hairs usually subtuberculate,
bi-dentate auricles, glabrous or minutely puberulous inflorescence, larger capitula,
longer (4.5 to 6 mm.), strongly 2-nerved, glabrous phyllaries, usually 4-fid
regular pistillate corollas, and slightly longer (1.5 to 2 mm.) blackish-brown
achenes.
Many specimens are intermediate in some degree in one or another or several
of these characteristics, and I have felt obliged to call such specimens hybrids. If
this entire complex is considered as one vast polymorphic species, which is what
Bentham in effect did, one is left with the problem of appropriate subspecific
categories for the many intermediate forms and the cause for such extreme
variability is ignored. Hooker's reaction to contact with limited numbers of
examples of these hybrid swarms was to form varieties for each; Bentham's response
to a somewhat larger suite was to merge them. Later authors have largely followed
Bentham.
Determination of specimens of S. glomeratus is further complicated by the
rather extreme range of foliar variation of the species and by certain trends in the
floral features as well. I strongly suspect this species of crossing rather frequently
with S. quadridentatus, although this is much more difficult to detect because of
their similarities in pubescence. I also suspect introgression, less commonly, with
one or more radiate species of Senecio, but I am not sufficiently acquainted with
these to name the probable offenders. PASE cursory examination of the material
laid in under the several names in the discoid and the radiate groups of
Australasian Senecio leads me to fear diis there is as much confusion of identity in
those taxa as in the erechthitoid material!
SENECIO HISPIDULUS X S. QUADRIDENTATUS
Erechtites tenuiflora DC. Prodr. 6: 296. 183
(? ecio epale Walp. Linnaea 14: 309. 1840.
Senecio tenuiflorus (DC.) Sieb. ex Sch. Bip. Flora 28: id 1845.
Pis ed apargiaefolia (Walp.) Sond. Linnaea 25: 524.
Erecbtites Muelleri Lange, Ind. Sem. Hort. Hafn. 28. er jour. Bot. 1874: 5, £. 3. 1874,
The evidence of hybridization between these two species is presumptive, based
on the existence of specimens intermediate in some particulars. Three related
collections from Tasmania, "between National Park and Westerway, in railway
cutting, 500 ft. alt., 30 Nov. 1929" (K!) are of especial interest: (1) Comber
1756, the first of these, is annotated: “Erechthites arguta DC. Tall erect peren-
nial, 2-4 ft. high with shrubby base; whole plant green; flowers yellow”.
specimen matched precisely the holotype of S. bispidulus. (2) Comber 1757 is
annotated: ‘‘Erechthites quadridentata DC. Tall erect perennial 2-6 ft. high
[Vol. 43
72 ANNALS OF THE MISSOURI BOTANICAL GARDEN
with a shrubby base; leaves and stems white wooly; flowers yellow". This speci-
men is S. quadridentatus, quite typical except for slightly more glabrate phyllaries
than usual. (3) Comber 1758 is annotated: “Erechtites tenuiflora DC. Inter-
mediate between above numbers; 3 — 5 ft. high; involucre purple". This specimen
is not quite identical with the type of E. tenuiflora, although the resemblance is
quite close. The leaf is shaped like that of S. quadridentatus but is broader and
auriculate at the base, although the auricle is not bidentate. The leaf is hispid
above, arachnoid and hispid beneath. The stem is cottony below, glabrate above
and on the inflorescence. The phyllaries are glabrate as in S. hispidulus but 7 to
8 mm. long, slender, and not strongly keeled. The achenes, however, are short and
non-attenuate. The general aspect is that of S. quadridentatus, but it has several
of the features of S. bispidulus. It is undoubtedly a hybrid, and quite possibly
the offspring of numbers 1756 and 1757 themselves!
With this clearly intermediate specimen as a guide, re-examination of a large
suite of specimens revealed numerous other individuals which were intermediate
in various degrees in the several features in which the two species differ, and which
I interpret as also being of hybrid origin.
One such hybrid, which has been dignified as a species, is E. tenuiflora. The
type material, Sieber “Fl. Novae Holl. no. 435” (BM!, G!, K!, P!), shows appreci-
able variation among the several specimens. They agree, however, in having rather
short phyllaries (6 rather than 7 to 8 mm. long), achenes of intermediate length
without necks, small but definite auricles, very little arachnoid pubescence and
more or less abundant hispid multicellular pubescence, features indicative of an
admixture of S. quadridentatus with S. bispidulus, with the former predominating.
Another such hybrid is E. apargiaefolia Sonder, the type of which (Fiedler's
Section, Bebr, MEL ex Hb. Sond. !) was located for me by Mr. J. H. Willis. Its
connection with S. bispidulus is shown in the auriculate leaf bases, the crisped
multicellular hairs on the lower leaf surfaces beneath the arachnoid pubescence and
the roughened and scabrous-haired upper leaf surface, the glabrous 2-nerved phyl-
laries, and the short cylindric achene. Its relationship to S. quadridentatus is seen
in the moderately long slender phyllaries, arachnoid pubescence, narrowly lanceo-
late leaves, and 4-fid florets with thickened glandulose apices. It resembles E.
tenuiflora but has a somewhat shorter achene and broader and more scabrid lower
leaves.
Sonder published E. apargiaefolia as a new combination for Senecio apargiae-
folius Walp., although based on a different collection. Walpers’ type, cited as
"Nova Hollandia Lhotsky legit", is supposed to be in Herb. Lucaeni at the Botan-
ische Institut, of Kiel University. Although the greater part of the herbarium was
saved in the destruction of the Institut during the war, it (as of 1952) was in
temporary storage and inaccessible. Although Sonder stated that he compared
Behr's specimen with Walper's type and found them conspecific, I have reduced
S. apargiaefolius with a query, pending examination of the type, if it still exists.
1956]
BELCHER——ERECHTHITOID SPECIES OF SENECIO 73
Walper's description actually applies very well to the Behr specimen except
for four points: (1) the widest leaf is only 31 lines instead of 5 to 6, and is not
repand-dentate; (2) the capitula are heterogamous, not homogamous; (3) the
leaf base has small semi-clasping auricles and is not simply attenuate; and (4) the
achenes are not all glabrous. The difference in leaf width may mean only that
Walper’s specimen is larger, or, more likely, that its leaf configuration trends more
toward that of S. bispidulus. Walpers stated: “Flores homogami tubulosi hermaph-
roditi”. Sonder, after examining the type, wrote: ‘Flores marginales feminei
tubulosi!”, and so transferred Walper's name to Erechtites. This is certainly true
for the Behr specimen, and I accept it for the Lhotsky specimen until it is available
for further study. Walpers possibly overlooked the auricles, which is easily done,
especially if there is much arachnoid pubescence around the node. It is also possible
that the Lhotsky specimen has auricles lacking or very slightly developed, for this
is a very variable character. The sheets of Sieber 435, for example, vary among
themselves from no auricles to auricles as long as 2 or 3 mm. is suggests a
multiple-allelic character appearing in varying degrees, and is one of the lines of
evidence in support of the hybridity of this complex.
All the achenes in the packet on the Behr specimen are definitely hairy on their
ridges, after the manner of S. hispidulus, and are approximately 2 mm. long. Most
of the achenes visible in the capitula, and all of the obviously immature ones, are
also hairy. A few achenes, however, apparently the most mature ones retained,
are certainly glabrous. This coincides with other observed instances, both in this
genus and others, of achenial pubescence diminishing with increasing maturity.
Accordingly, I accept Walper's characterization as based upon the most mature and
accessible achenes and not in actual disagreement with Sonder's type.
My inquiry to Copenhagen concerning the type of E. Muelleri Lange went
unanswered, but two sheets so determined and said to have been raised from seed
from Copenhagen were included in the loan from Vienna. These agreed well with
Lange's emended description and plate (1874), and may be presumed authentic.
(I was unable to locate the 1861 seed list in which Lange originally published.)
Their achenes ranged from 2.4 to 3 mm. long, and were heavily ribbed and only
slightly attenuated. The leaves were long and rather wide, regularly denticulate
to sinuate-dentate or with an occasional long-exserted linear-lanceolate tooth.
phyllaries were glabrous or glabrate, the leaves variably arachnoid, mostly glabrous
above. From these features I conclude that this represents another hybrid between
S. bispidulus and S. quadridentatus, essentially stabilized near the latter parental
line. The relative uniformity of these plants and their immediate ancestors is not
necessarily an effective argument against their hybrid nature; as pointed out by
Anderson??: “variation between [hybrid] individuals will lessen as parental comb-
inations are approac
I have not cited specimens of hybrid nature, except those indicated above, for
lack of space. They are, however, quite common in all the herbaria examined.
38 Anderson, E. Introgressive hybridization. New York. 1949.
[Vol. 43
74 ANNALS OF THE MISSOURI BOTANICAL GARDEN
ARRHENECHTHITES
ARRHENECHTHITES Mattf. Engl. Bot. Jahrb. 69: 288. 1938.
Perennials with shoots herbaceous or suffruticose. Capitula heterogamous with
few florets. Involucre of 5 to 8 sub-biseriate phyllaries, the narrow scarious
margins of the exterior ones inserted into grooves beneath the lateral nerves of
the interior phyllaries. Marginal florets pistillate, as many as or slightly more than
the phyllaries and opposite them; corolla filiform with base inflated and more or
less indurated, oblique and truncate at the apex, minutely and irregularly toothed,
or subligulate; style exserted, arms short and glabrous; achene subcylindric, with
a callose rim base at the apex and also sometimes callose-annular at the base. Disc
florets fewer than or as many as the marginal florets, structurally perfect but
functionally staminate, corolla with limb infundibuliform, deeply 5-lobed, lobes
recurved; style arms erect, short and astigmatic, or style apex subentire, papillose-
hairy over the outer faces; achene narrowly cylindric or stipiform, slightly longer
than that of the pistillate floret, usually with abortive embryo. Pappus filiform,
white, exceeded by disc florets.
enus characterized by functionally staminate disc florets with = arms
reduced, astigmatic, and papillose over the outer faces.
rechthitoi cio differs in having all florets fertile, perfect lts with
short erect corolla lobes and style arms glabrous except for a terminal margina
fringe of papillae. Brachyglottis Forst. of New Zealand approaches most closely
in composition of the capitulum, having marginal florets 8, opposite the phyllaries,
functionally pistillate with sterile stamens; disc florets 2, apparently perfect
Endemic in the higher mountains of New Guinea, with an extended outlier in
the Australian Alps and Blue Mountains. Six species are known.
Type of the genus: A. tomentella Mattf.
KEY TO yya OF ARRHENECHTHITES*
A. Phyllaries 5, pistillate florets 5; disc florets solitary
A. tomentella
AA. than one 7 or 8, pistillate ds s as many or slightly more numerous; disc Sec more
2. A. mixta
- long m. long; corolla of marginal florets
ne 4- "Zeen or ei diéca d obliquely truncated; leaves entire
or mer nice oothe
C. Style di vei been? scarcely bifid, clavate with shallow notch and a ec:
papillose hairs; es 2.5 to 5 cm. long vo oe
CC. Style of ped tins pac Uh arms erect, uniformly papillose on the outer
Ap leaves 10 cm. lo
D. Leaves broadly elliptical, ‘we to 15 cm. long, 4.5 to 7 mm. wide ........4. A. —
DD. Leaves narrowly lanc Hs te, 15 cm. long or longer, Gees 3 cm. > wide
E. Leaves acute, 15 to 16 m ea ng; disc florets 2 5 A. posa aru
EE. Leaves ene cm. long or longer; disc florets 4 to 6 ........
6
6. A. dolichocephala
*Modified from Mattfeld, Engl. Bot. Jahrb. 69:288-289. 1938.
1956]
BELCHER—ARRHENECHTHITES 75
1. ARRHENECHTHITES TOMENTELLA Mattf. Engl. Bot. Jahrb. 69:290. 1938.
Known to me only from Mattfeld's description. Apparently collected only
once.
Holotype: Northeast New Guinea, "Bezirk Morobe: Sarawaket, Busu Tamunac,
an offenen Grasland, 24-2700 M. ü. M.", 31 Jan. 1937, Clemens 5251 (B,
destroyed).
2. ARRHENECHTHITES mixta (A. Rich.) Foi comb. nov.
Senecio mixtus A. Rich. Sert. Astrolabe, ier? ES 18
Wir e mixta ~g Rich.) DC. Prodr. 6:2 gree non Benth. Fl. Austral. 3:659. 1866;
Black, Fl. S. Austral. 4:610. 1929.
Sinn herbaceous, erect, simple or sparingly branched, purpurascent, sparsely
arachnoid. Leaves glabrous to sparsely arachnoid above, with sparse arachnoid hairs
overlying numerous short strigose multicellular hairs beneath, sometimes purpuras-
cent; lowest leaves alate-petiolate, abruptly broad-lanceolate to ovate, with large
coarse teeth; medial léaves with attenuated portion progressively shorter and sinuses
between teeth progressively deeper so that within 5 or 6 nodes the blade has become
lobate to pinnatisect, the segments notably variable in size and shape, borne es-
sentially at right angles to the midrib, more or less denticulate, shortly acuminate,
acute, or rarely obtuse; sinuses equalling or exceeding the segments in width; leaf
abruptly reduced below first node of inflorescence to a linear to ensiform bract;
bracteoles of peduncle about 4 mm. long and 0.2 mm. wide. Inflorescence cymose-
paniculate, much branched, lax; capitula solitary on elongate peduncles to 5 cm.
long, ecalyculate but surrounded by 3 to 5 linear bracteoles 5 to 7 mm. long and 0.5
mm. wide, which arise from the terminal 2 or 3 mm. of the peduncle; end of pe-
duncle above bracteoles abruptly doubled in diameter from 1.0 mm. to form a very
short receptacle. Involucre (12 to) 20 to 24 mm. long, sub-biseriate; phyllaries
long, linear-lanceolate, tightly appressed in bud, becoming widely separated in fruit
or sometimes adhering tightly over the undeveloped achenes to simulate an elongat-
ed receptacle, flat-keeled, usually indistinctly 4-nerved, with mixed pubescence
similar to that of leaf, often purplish, acute to acuminate, ciliolate; outer phyl-
laries narrower (0.71 mm.), non-scarious, frequently edged with purple; inner
phyllaries wider (1.25 mm.) with light scarious margin 0.3 mm. wide on either
side; intermediate phyllaries scarious on one margin, non-scarious and sometimes
purple-edged on the other. Marginal florets pistillate, or having corolla with the
base about 1 mm. long and 0.4 mm. in diameter, the tube 12 to 14 mm. long, 0.2
mm. in diameter, the oblique limb 1.0 to 1.5 mm. long, 0.4 to 0.5 mm. wide, and,
imperfectly 2- or 3-dentate; style ege? vum exceeding the tube; style arms
divergent at an angle between 45^ and 9 t full anthesis, about 0.5 mm. long,
glabrous or minutely pubescent, Bac on the adaxial faces. Central florets
2 or 3, structurally perfect but not ripening achenes, corolla about 17 mm. long,
the tube above the incrassate base slender, 14 mm. long and 0.2 to 0.5 mm. in
diameter, the limb 3 mm. long and 1 mm. in diameter, regularly 5-fid with lobes
deltoid, 1 to 1.25 mm. long, 0.5 mm. wide at base; anthers about 2 mm. long,
1/2 to 2/3 exserted, minutely sagittate, the terminal portion non-polliniferous and
[Vol. 43
76 ANNALS OF THE MISSOURI BOTANICAL GARDEN
broadly flattened, 0.5 mm. long; style scarcely exserted; style arms brief, appressed,
clavate, non-stigmatic, long-papillose over all the abaxial surfaces. Achene o
pistillate floret fusiform to subcylindric, callose-annulate at base and apex, 6-7
mm. long, 1 mm. in diameter in the middle, 0.5 mm. in diameter at either end,
about 12-ribbed, brown, glabrous or with a single row of short slender hairs down
the middle of each groove. Achene of central floret flattened, 6 mm. long, 0.2
mm. in diameter in the middle, 0.5 mm. in diameter at either end. Pappus white,
scarcely barbed, subequalling florets and phyllaries.
AUSTRALIA. NEW SOUTH WALES: Port Jackson, Gaudichaud 6 e berati elh lee
56
Cheel (NSW); Eskbank, Jan, 1915, Hamilton Sid ); Mt. Tomb, Gregson (NSW).
VICTORIA: East Gippsland, spurs north of Mt. Drummer, Dec. 1937, Hunter (MEL);
Bidwell, Feb. 1943, Hunter 9 (MEL); Genoa Peak, Ta 1947, Willis (MEL). “Australia
orient.”, Hugel (W, very meagre specimen).
Richard interpreted the subligulate marginal floret of this species as indicating a
transitional form between the discoid and radiate sections of Senecio. He accur-
ately described the unusual style arms of the central florets but did not recognize
their uniqueness. The achenes of his specimen unfortunately were quite immature
and he did not note the abortive development of those of the central florets. Mature
specimens, however, show the imperfect central florets, reduced number of florets,
and erect style arms with pubescent outer faces which together characterize Ar-
rbenecbtbites. Y therefore transfer Richard's species to this genus, although this
requires emending Mattfeld's diagnosis to admit pistillate florets subligulate and
more numerous than the phyllaries, These are minor changes when the excellent
agreement in other respects is noted.
Mr. J. H. Willis has very kindly written as follows on the ecology of A. mixta:
“It is a perennial with fleshy rooting system and bright reddish purple coloration on
the stems and undersides of the leaves, and it favors well-watered stony hillsides
in montane forests, from about 1000 to 4000 feet altitude." Inclusion of this
species in Arrbenecbtbites extends the range of this genus, formerly known only
from the higher mountains of New Guinea, by some 27 degrees of latitude.
The holotype at Paris apparently has not always been recognized as such. An
original label as S. sonchoides A. Rich. was only corrected by an anotation by Joret
in 1949! 1 find no explanation for the earlier epithet. The specimen, although
undoubtedly in much poorer condition now than when illustrated, is certainly the
one from which the plate was prepared. The plate, however, fails to show that
the phyllary margins are free to the base, even though in firm contact with one
another in the lower half of the involucre. Also, the corolla lobe in the detail of
the pistillate florets is shown as too short. Most of the capitula shown, and some
of the leaves, are now gone, apparently clipped off and distributed to other her-
baria.
1956]
BELCHER—ARRHENECHTHITES Fi:
Very probably the leaves and capitula of this species which were admixed with
the fragments of S. squarrosus in the packet labeled E. Richardiana in the Prodr-
omus Herbarium came from this specimen at Paris. De Candolle left no specimen
identified as E. mixta in the Prodromus Herbarium, no. 15 under Erechtites being
omitted. The diagnosis in the ‘Prodromus’ is taken entirely from Richard, except
for the data on number of phyllaries and length of capitulum. These he must have
derived from a study of Richard's plate, for it would seem that if he had recognizec
the fragments mixed in with E. Richardiana he would have separated them, as I did,
and identified them in the proper sequence.
Bentham’s description of E. mixta resembles that of de Candolle but is original,
based on Fraser, Piper’s Hill, New South Wales (K!). This specimen is S, runcini-
folius, as discussed above in that connection. Black’s description, drawn from
Bentham's, is of the same.
3. ARRHENECHTHITES HAPLOGYNA (F. Muell.) Mattf. Engl. Bot. Jahrb. 69: 292.
1938.
Senecio baplogynus F. Muell. Trans. Roy. Soc. Victoria 1:14. 1889.
Erechthites haplogyna (F. Muell.) Mattf. Engl. Bot. Jahrb. a 422. 1929.
Peduncle glabrate, narrowly and sharply ridged. Bracteole linear-lanceolate, 2
mm. long, subtending capitulum immediately below receptacle. Capitulum 9 mm.
long, technically ecalyculate, with sub-biseriate involucre of 8 phyllaries 6 mm.
long, 0.7 to 1.2 mm. wide, with about 10 narrow and sharply ridged nerves, acute,
minutely ciliolate. Marginal florets 8, almost 6 mm. long; corolla 0.7 mm. in
diameter at inflated base, with tube 4 mm. long and gradually tapered and indur-
ated in lower 3 mm., apically slightly expanded, irregularly and briefly 3- or 4-fid
with one sinus much more deeply cleft than the others; style exserted, coarse;
style arms about 0.3 mm. long, recurved, glabrous, blunt; achene (immature) 2.25
mm. long, about 0.4 mm. in diameter. Disc florets 8; corolla 6 mm. long, in-
fündibuliform, with inflated base indurated and 0.7 mm. in diameter, tube 3.5
mm. long and abruptly reduced in diameter to 0.3 mm. and then gradually enlarged
to throat diamecer of 0.5 mm., regularly 5-lobed with lobes acute and multinerved,
2 mm. long and 0.5 mm. wide; anthers 5, polliniferous, 1.75 mm. long, with bases
obtuse and apices long-appendaged and narrower; style subequalling corolla lobes,
almost undivided, abruptly dilated at apex to about 0.5 mm. in diameter and
marginally papillose, probably non-stigmatic; achene very slender, about 2.5 mm.
long and 0.2 mm. wide, immature and probably abortive. Pappus slightly tawny,
barbellate, equalling phyllaries, exceeded by central florets but not by marginal
ones.
Known only from a single collection in the Owen Stanley Range of New
Guinea, Summit of Mt. Knutsford, 1889, McGregor, holotype (MEL). Material
examined: A single capitulum of the holotype.
The above description is based on an examination of this capitulum after moist-
ening it with detergent solution, and is intended only to supplement Mueller's
ription.
[Vol. 43
78 ANNALS OF THE MISSOURI BOTANICAL GARDEN
This species is quite similar to A. novoguineensis (S. Moore) Mattf. Indeed,
in transferring Mueller's species to Erechtites in 1929, Mattfeld stated that they
might be identical. Later, in transferring them both to Arrhenechthites, he kept
them separate on the basis of a difference in leaf size. I at first supposed that the
species named by Moore might simply be based on a more luxuriant shade form of
Mueller's plant, a supposition gaining some support from ecological data in the
two reports. After comparing the florets from the two types, however, I believe
that the following differences in floral characteristics reinforce the imperfectly
known vegetative differences:
A. baplogyna
A. novoguineensis
Central florets
Corolla tube o
marginal florets
Achene
8, v corolla lobes
m. long, 0.4 mm. wide
Rigidly incrassate, gradually
tapering for about 34
length
its lengt
Apparently about 2.5 mm.
ong when mature
2, with corolla lobes 1.5 mm.
long, 0.35 mm. wide
Rigidly incrassate for about V4
to L4 its length and abruptly
and 3 in gege eter
About 2 mm. long
Style-arm apex
Extremely short, almost undivided, |0.42 to 0.45 mm. rne little or
and abruptly dilated not at all dilate
4. ARRHENECHTHITES MASTIGOTHRIX Mattf. Engl. Bot. Jahrb. 7: 477. 1940.
Known to me only from Mattfeld's description. Apparently collected only
once so far. This is the only one of the species of this genus for which Mattfeld
explicitly stated: "Typus in Herb. Berol.".
Holotype: New Guinea, Saruwaged Mts., Masak river, am Wasser, 1800-2400
meters, 10 Nov. 1937, Clemens 7509 (B, destroyed).
5. ARRENECHTHITES NOVOGUINEENSIS (S. Moore) Mattf. Engl. Bot. Jahrb. 69:
292. 1938.
Erechthites novoguineensis S. Moore, Trans. Linn. Soc. 9: 86. 1916.
Marginal florets pistillate, 8 to 10; corolla base inflated and indurated, abruptly
softened and reduced in diameter, tube slender, obliquely truncate or irregularly
briefly toothed. Central florets 2, with 5 corolla lobes 1.5 mm. long, 0.35 mm.
wide; anthers polliniferous; style scarcely exserted, arms only 0.42 to 0.45 mm.
long (rather than 1 mm., as given by Moore) and slenderly clavate, nonstigmatic,
and papillose-hairy on the abaxial sides only.
oe known only from the type collection from dortiwestern New
Holotype: New Guinea, Nassau Mountains, Carstensz Peak, 6551 to 7956 ft.,
27 Jan. 1913, Boden-Kloss (BM!).
The holotype, and only, specimen is a small fragment, consisting of a short
apical length of stem with two main leaves, their axillary (and flowering) bran-
1956]
BELCHER—EXCLUDED SPECIES 79
ches, the large terminal corymb, and an additional large leaf (detached). It agrees
well with the generic characterization as given by Mattfeld for Arrhenechthites,
especially the structure of the capitulum, except for one minor detail. Whereas
Mattfeld specified that the marginal florets should be of the same number as the
phyllaries and opposite them, the marginal florets in this specimen number from
8 to 10 per capitulum, whereas the number of phyllaries is 8. As pointed out in the
discussion of A. mixta, there is also in that species an excess number of marginal
pistillate florets, for the accomodation of which I have slightly expanded the gen-
eric limits. It appears that Mattfeld did the same by implication when he included
Moore's species in the new genus. The supplementary description given above deals
only with the finer structure of the capitulum, which was inadequately treated by
Moore.
6. ARRHENECHTHITES DOLICHOCEPHALA Mattf. Engl. Bot. Jahrb. 69: 289. 1958.
Known to me only by Mattfeld's description. Apparently not again collected.
Holotype: Northeast New Guinea, "Bezirk Morobe, Sarawaket, Busu Tamunac,
an offenen Stellen,” 2100 to 2400 m., 30 Jan. 1937, Clemens 5287 (B, destroyed).
SPECIES TO BE EXCLUDED FROM ERECHTITES
In addition to the Australasian species of Erechtites which I have placed in an
erechthitoid group of Senecio, and the three species referred to Arrhenechthites, six
other taxa have been described as Erechtites which do not belong to that genus. One
is a Blumea. The other five belong in Senecio; one is erechthitoid, one is radiate,
and three are discoid. These six are discussed here. In addition, a seventh name,
a nomen nudum, is shown to be based most probably on still another discoid Senecio.
BLUMEA TENERA Merr. Philip. Jour. Sci. Bot. 7: 250. 1912.
Erechthites Bukaensis Rech. & Muschl. in Rech. Denkschr. Akad. Wiss. Wien 89: 620.
1914.
Rechinger 4043 (W!), syntype of E. Bukaensis, has a pluriseriate involucre,
caudate anthers, and other features of the tribe Inuleae, genus Blumea. Among the
Indonesian sheets of Blumea at Kew was an undetermined sheet from the Solomons
and two sheets from the Philippines which agreed with the Rechinger specimen in
fullest detail. The Philippine sheets included Merrill 7363, isotype of Blumea
tenera, and agreed fully with the description. This species is now seen to have a
wide, if spotty, distribution from Luzon to the Solomons. It appears to be a low-
growing member of the rank-grass or "cogonal" association. Further search in
such habitats should reveal additional stations.
Bram. LUzON: Prov. Rizal between La Loma and Maypajo, near
Manila, Dec. 31, 1910, "in open grasslands a few meters above sea-level, not common",
Merrill 7363 (K, isotype of Blumea tenera Merr.) ; Palawan, Taytay, May 1913, Merrill
9397 (K). New GUINEA: “am Ramm Fluss", Jan. 1902, R. Sch K
SOLOMON Is.: Buka, "in Alang-Alang (Imperata)”, Sept. 1905, Rechinger 4043 (W, syn-
type of E. Bukaensis) ; New Pomerania, Simpsonhafen, Sept. 1905, Rechinger 4205, 4267
(W, both det. as E. prenanthoides DC.).
Merrill also cited “Merrill 679 from Culion, and For. Bur. 5874 Curran, from
Zambales Province, Luzon, both small forms", but I have not seen them. In addi-
[Vol. 43
80 ANNALS OF THE MISSOURI BOTANICAL GARDEN
tion, Rechinger 4044, from the same station as Rechinger 4043, the other syntype
of E. Bukaensis, is most probably B. tenera also, but I have not seen the specimen,
which is at Vienna.
SENECIO LEPTANTHUS Phil. Anales Univ. Chile 88:15. 1894; emend. Reiche, Fl. Chile
4: 225-226. 1905.
Erechthites leptantha (Phil.) Cabr. Not. Mus. La Plata 14 (Bot. No. 69): 75-78. fig. 1.
1949.
CHILE: Prov. Antofagasta, Taltal, Oct. 1925, Werdermann 825 (K, 2); Desert of
Atacama, 1890, Morong 1292 (K).
I have not seen Philippi’s type (Geisse, near Caldera). The specimens cited
above, however, agree with his description except for the achenes, which are very
densely clothed with short blunt white hairs, rather than glabrous. This is recon-
ciled by Reiche: “Aquenios cortamente blanco-peludos (no peludos, como se dice en
la diagnosis original.)" Cabrera also describes the achenes as “densa y cortamente
papiloso-pubescentes”.
In Werdermann 825 the first row of marginal florets was filiform and pistil-
late, those of the second row functionally pistillate but with rudimentary stamens,
and those of the third and subsequent rows perfect with functional stamens. Morong
1292 had some florets in the outermost marginal row with rudimentary stamens
and others without, while those in the second and subsequent rows were all func-
tionally perfect. Of more importance, all of the specimens agreed as to style-
arm apices. The tips of the style arms in the marginal florets were blunt, without
dorsal protruding papillae. In the perfect florets the style-arm apices were sharply
truncate with a half-crown of moderately enlarged but non-fused papillae fringing
the apex (cf. Cabrera, fig. 1-H).
Cabrera transferred this species to Erechtites because of the pistillate filiform
marginal florets. In view of the lack of the appendage of fused papillose hairs
which characterizes Erechtites, I believe it better to return this species to Senecio.
In structure and size of capitula it strikingly resembles erechthitoid species of
Senecio in Australasia. This resemblance is reinforced by the unusual velutinous
achene, quite like that of certain Australian species, which is so very different in
shape, size, ribbing, color, and pubescence from the distinctive achene of the genus
Erecbtites.
SENECIO glossanthus (Sond.) Belcher, comb. nov.
Erecbtites glossantba Sond. Linnaea 25:524. 1852.
Senecio brachyglossus F. Muell. ex Benth. and var. major Benth. Fl. Austral. 3:670. 1866;
d es elatior Benth. 1. c.; non S. brachyglossus Turcz. Bull. Soc. Nat. Mosc. 2411:
Annual; stem erect, 20 cm. tall, sparingly branched above, glabrate, faintly
striate, purplish below. Leaves 2 mm. broad at the base, alate, broader toward
apex, bearing on either side one or more narrow lobes at an angle of from 45°
to 60° and approximately 5 times as long as wide, their width being about one-half
that of the rachis, the margins all minutely and irregularly callose-denticulate,
glabrate or with occasional long coarse isolated hairs, especially along the nerves
1956]
BELCHER——EXCLUDED SPECIES 81
beneath, lower leaves sessile and neither auriculate nor clasping. Upper leaves
similar to the lower but more lobate and smaller, with base auriculate and semi-
amplexicaul. Inflorescence compactly cymose, capitula rather few, peduncles
short, primary floral bracts equaling or exceeding the branches. Involucre of 8
to 10 (to 12) phyllaries 4 to 6 mm. long, inconspicuously 2- to 4-nerved, with
margins scarious and apex acute, sphacelate and minutely papillose; calyculus of
6 to 8 broadly lanceolate short bracteoles, 1.5 to 2 mm. long and 0.5 to 0.6 mm.
wide, margins laciniate, apex sphacelate; peduncles long-haired in the axils. Margin-
al florets few, ligulate, pistillate, without rudimentary stamens; corolla tube slender,
2.5 to 2.75 mm. long, 0.25 mm. in diameter, 4-nerved; ligule up to 2 mm. long
and 0.5 mm. wide, 2-nerved, lanceolate, apex acute, revolute; style arms ca. 1
mm. long, bluntly truncate or slightly rounded on apex, without encircling papil-
lae, exserted. Disc florets perfect, about 25; corolla very slenderly infundibuli-
form, 4 mm. long, 0.4 mm. in diameter, minutely and regularly 5-fid, teeth about
0.3 mm. long; style arms about 1 mm. long, not exserted, apices slightly rounded,
marginally papillose. Achene 2 to 2.5 mm. long, very densely clothed with ap-
pressed tawny hairs about 0.5 mm. long which completely conceal the slender
achene and extend beyond its apex. Pappus white, capillary slightly exceeding
phyllaries.
AUSTRALIA. S. AUSTRALIA: "ad litus sinus St. Vincent", Mueller (MEL, syntype) ; Mt.
Lyndhurst, Sept. 1898, Koch 220 (K). victoria: Aug. 1843, Robertson 468 (K);
Melbourne, 1854, Adamson 342 (K); Point Nepean, Mueller (K, syntype of S. bracbyglos-
sus Var. major). WESTERN AUSTRALIA: Swan River, Drummond 44 (K); Drummond 377
(K); Klondinin, Sept. 1932, Young (K).
This briefly ligulate and rather variable species bridges the gap between erech-
thitoid species with irregular corollas in the marginal florets and definitely ligu-
late eusenecionoid species. It was assigned to Erechtites section PLAGIOTOME by
Sonder, despite his description, “ligulis stylum aequantibus vel superantibus." i
might indicate that he also felt the vagueness of the boundary between Senecio and
the Australian species assigned to Erechtites. Sonder described the capitulum as 3
lines long with about 12 phyllaries and 20 to 30 florets. Bentham described S.
bracbyglossus as having capitula 2 lines long with about 8 phyllaries and 16 to 18
florets, and then wrote: "var. (?) major. Flower-heads larger. Involucres about
5 lines long with about 12 bracts. Florets also more numerous." Thus var. major
agrees exactly with Sonder's description of E. glossantba.
The syntype from St. Vincent's Gulf has phyllaries 6 mm. (3 lines) long, as
stated by Sonder, but the number ranges from 8 to 10, usually 9, and never reaches
12. The other syntype collection by Mueller, cited by Sonder as "ad agros prope
urbem Adelaide", may be the specimen cited by Bentham for his typical variety as
"near Adelaide, F. Mueller". If so, it probably has 8 phyllaries 4 to 5 mm. long,
as do the other specimens at Kew which Bentham determined as typical S. brachy-
glossus. The Pt. Nepean specimen, cited by Bentham for var. major, agrees exactly
with the one from St. Vincent's Gulf except that it does have 12 phyllaries. Drum-
mond 377, cited by Benthan for var. major, differs from both the above in having
[Vol. 43
82 ANNALS OF THE MISSOURI BOTANICAL GARDEN
only 8 phyllaries, much broader leaves, and the corolla merely obliquely trun-
cated, but since the achene is characteristically pubescent I consider it a variant of
S. glossanthus. The Wilson Promontory specimen is to be excluded from the species
altogether, as it had a bifid ligule 2.5 mm. long, a glabrate achene with an oblique
annulus, and leaves with broadly amplexicaul bases. Thus Bentham's var. major
is based on a mixture of unrelated elements and should not be maintained.
I conclude that Senecio glossantbus is best interpreted as a variable species dis-
tinctively characterized by it unusually hairy achene. The variation of the phyl-
laries from 8 to 12 in number, and from 4 to 6 or 7 mm. in length, may be an
edaphic response, which would make it unwise to make varietal distinctions without
further information
Bentham also published a var. elatior, based on a specimen from the Blue Moun-
tains. The type specimen at Kew and isotypes sent from Melbourne and Sydney
agree in having the ray florets with a trifid ligule only 1 mm. long, 10 to 12 phyl-
laries, and a leaf much larger and more dissected than that of S. glossanthus. In
addition, their achenes are distinctively different in size, shape, and pubescence
from those of S. glossantbus. Var. elatior is to be excluded from this species.
SENECIO GEORGIANUs DC. Prodr. 6: 371. 1838; non Greenm. Ann. Mo. Bot. Gard.
25: 803. 1938.
Erecbtites candicans Hook. f. Lond. Jour. Bot. 6: 122. 1847.
Hooker did not specify a type in his description of E. candicans, stating merely:
"Hab.? (Gunn in Herb. Hook.) ". There is a sheet at Kew from Hooker's herb-
arium on which the right-hand specimen bears the following annotation: "Van
D[ieman's] L[and], Gunn", düneilun candicans nob." (in Hooker's script),
and "701. Senecio Georgianus DC.". I take this to be the holotype, and it agrees
with the description given by Hooker. Dissection of a capitulum at the bud
stage showed it to have marginal florets 5-fid with polliniferous stamens; that is,
it is a discoid rather than an erecthitoid Sezecio. Comparison of this specimen with
both the isotype (K!) and the holotype (G!) of S. georgianus DC. shows them
to be conspecific, as later concluded by Hooker and as given in the 'Index Kew-
ensis’.
Senecio georgianus Greenm. was published as a new name for S. Hallii Hieron.,
1895, non Britton,,1889. Apparently Greenman overlooked the anterior Candol-
lean name. Britton’s plant was from Wyoming, Hieronymus’ from South America.
As far as I am aware, the latter still requires re-naming, but that I leave to students
of Senecio of South America, if indeed they have not already done so.
SENECIO POLYPODIOIDES (Greene) Greene ex Greenm. Monogr. Senecio, T. 1: 25.
ics od ; Engl. Bot. Jahrb. 32: 21. 1902.
io poly podioides Greene, Pittonia 3: 90. 1896, nom . prov
Erebies ol poly podioides Greene, Pittonia 3: 90. 1896, nom. nov. for S. gracilipes Robins.
Senecio gracilipes port & Greenm. Amer. Jour. Sci. 50: 156. 1895; non A. Gray, Proc.
. 5: 14
e poly podioides Kate & Greenm. ex Jacks. Ind. Kew. Suppl. 1:393. 1906, error.
1956]
BELCHER—EXCLUDED SPECIES 83
The isotype at Kew has functional stamens in all florets and belongs in Senecio,
Sg MULGEDIIFOLII, where Greenman finally placed it.
Type: Mexico, State of Oaxaca, Sierra de Clavellinas, 9000 ft., 24 Oct. 1894,
Pringle 6010 (GH; K!).
SENECIO RUNCINATUS Less. Linnaea 6: 410. 1831.
hig ? runcinata (Less.) DC. Prodr. 6: 295. 1838.
Thi cies e uniformly homogamous capitula. Not one filiform pistillate
floret was rei among several hundred florets which I examined from scores of
heads from many different specimens (K, BM, P, G, W, MICH). The style-arm
apices are truncated, with a lateral whorl of stubby pollen-presentation papillae
but with no terminal tuft of fused hairs such as characterizes the genus Erechtites
in sensu strictiore. This species is, in fact, a discoid Senecio, section MULGEDIIFOLIL.
Its delimitation and synonymy will be treated in a separate publicat
Type: Mexico, Vera Cruz, “Malpays de la Joya, Novbr.”, Sobiede & Deppe
(B, destroyed).
ERECHTITES ELONGATA A, Gray ex Jacks. Ind. Kew. 2: 860. 1893, nomen nudum;
DC. Prodr. 6: 294, 1838, nomen nudum.
As pointed out in the discussion of Erechtites, this "E. elongata", in the light
of Rafinesque's letter to de Candolle, refers most probably to Senecio elongatus
Pursh, Fl. Am. Sept. 2: 529. 1814. This was treated by Torrey and Gray as a
discoid form of S. aureus var. obovatus Torr. & Gray, and now appears in ‘Gray’s
manual’ (8th ed.) as S. obovatus forma elongatus (Pursh) Fern. It is as unrelated
as possible to E. bieracifolia.
At least four other names have been published in Erechtites as nomina nuda.
These are: E. senecioides Turcz. (Bull. Soc. Nat. Mosc. 24: 201. 1851), probably
an error for E. sonchoides, as explained on page 48; E. macroglossa Muell. (Rept.
Gov’t. Bot. 1853, p. 15. 1853), entirely unidentified; and E. pauciflora and E.
uniflora Raf. (Herb. Raf. 68. 1833), both of which most probably were applied
to depauperate forms of typical E. hieracifolia.
84
[Vol.
ANNALS OF THE MISSOURI BOTANICAL GARDEN
INDEX TO ScIENTIFIC NAMES
43
Previously published valid names are in ordinary type; new names and the final mem-
bers of new combinations are in bold face type; synonyms are in italics.
Anonyme Robin
Arrhenechthites
dolichocephala
haplogyna
mastigothrix
mixta
novoguineensis
nthoides
Ge socrpbelum valerianaefolium `
rechtites
agrestis
ambigua
apargiaef olia
arguta
WI
-—
var. cacalioides
var. cacalioides
. pubescens
var. carduifolia
var. glabrescens 14
var. hieracifolia 13
var. intermedia 14
var. megalcarpa .—— . ———— M
minor 14
var. praealta 14
r. typica 14
Gees 19
Hieraciifoliae 1, 13
pispid 48, 66, 69
ignobilis E
ana GE
Labillardieri 46
lacerata 51
leptantba 80
macroglossa 83
megalocar pa cH
minima 46
missionum 32
mission 34
var. lanceolata 32, 34
mixta 54, 75
Muelleri 71
novoguinensis 78
organensis 29
pauciflora - 83
apuana 53
petiolata 31
picridioides 48, 54
poly podioides 82
praealta 2
realta 10, 14
prenantboides 31, 43 46
var. mino 55
var. picridioides 48
pumila 46, 69
pyrophi 49
quadridentata 49, 57
var. glabrescens 57
. Gunnii 60
Richardiana 48
runcinata 83
scaberula 69
senecioides 83
Senecioneae 7
ecioninae 2,8
soncboides 43
sulcata 19
tenuiflora 71
uniflora 83
valerianaefolia 25
1956]
BELCHER—ARRHENECHTHITES
f. organensis 29
var. organensis 29
f. prenanthoides — . 30
f. reducta 30
f. valerianaefolia |. 26
Erecbibites 10
aspera 20
malasica 20
papuana 53
rosea 26
zeylanica var. malasica o 2 26
eis 10
carduifolia 19
hieracifolia 14
microcephala 61
idu 14
tomentosa 57
j 10
bieracifolia 14
necio 3; 37
albifloru. 31
zeg 71
argutus 61
Atkinsoniae 41
rsti 69
bipinnatisectus 41
biserratus 43
brachyglossus 80
ar. elati 80
ar. major 80
cacalioides 19
carduifolius 19
ecaisnei 3
diversifolius 56
dunedinensis i 55
elongatus 5, 83
erechthitoides SE 29
Fischeri 19
flaccidus 43
georgianus 82
glabrescens De *
gland 57
— EE,
61
glomeratus X hispidulus Gi, 70
oaa — 80
Goyazensis Se
gracilipes 82
60
beslogymaus 77
85
beteropbylla 46
hieracifolius 13, 19, 46
var. gigantea 14
hispidulus 65
var. dëses 69
var. hispidulus 66
var. scaberulus 69
bispidulus 66, 69
hispidulus X quadridentatus ............ i
ignobilis 36
kermadecensis 45
laceratu 51
lactucoi 31
laticostatus 64
leptanthus 80
Lessoni 61
Lessonian 61
microcephalus 64
minimu: 46
ar. minimus 46
var. picridioides 48
mixtus 75
ülleri 46
multicaulis 68
53
mert? $2
polypodioides
poly podioides 82
prenantboides 57
pumilus 46, 68
pusillus 70
pyrophil 49
quadridentatus .. 57
runcinat SE EE
E E EE 48, 54
$ udus a M
ides xs A4
a _~ 48
tenuiflorus 71
valerianaefolius 25, 26
Vukotinovici i M
wairauensis du 99
Warscewiczii 46
cio 10
chus
agrestis 19
iliensis 19
erythropappus 26
laevis -B
occidentalis 19
MISSOURI BOTANICAL GARDEN = >
RomarE Woonson, Jay
Berta M. TRYON, `
Volume XLIII Number 2
| Annals
of the : ae i 7 ee dex ^]
Annals
of the
Missouri Botanical Garden
Vol. 43 MAY, 1956 No. 2
THE AUSTRALIAN AND NEW ZEALAND SPECIES OF PITTOSPORUM*
ROBERT C. COOPER**
Many of the Australian and New Zealand species of Pittosporum have long
been treasured in gardens of Australia, New Zealand, France, and England, for
their flowers, foliage and sweet scent, and several are common hedge plants and
ornamentals in California and Florida. In addition to an aesthetic appeal, the
species have scientific interest and potential economic value.
The most recent inclusive treatment of Pittosporum is in the treatment of the
family Pittosporaceae by Pritzel,! who found it was impossible to make a satis-
factory arrangement or classification of the genus and relied upon the geographic
distribution of the species. He wrote as follows:
Etwa 160 Arten in den Tropen und Subtropen der östlichen Hemispháre von
P
st-Afr s zu den Inseln des Pazi e befriedigende natürliche
EAE der G di E nicht gegeben werden kann, so bleibt nichts
anderes übrig, als sie Verbreitung zu behandeln. Ein künftiges
hrer
System bs Gattung d EE vor allem auf die ceni zu gründen haben, und
diese ist bei vielen Arten noch nicht genügend bekan
He discussed and keyed the species separately i the following regions:
Malaysia ralia Other Polynesian Islands China and Japan
Philippine Islands Solomon Gë and Bismarck New Zealan Africa à
New Guinea Hawaiian Islands Macronesia
onia Fiji SI Tonga Islands India Madagascar Terr.
The species of New Caledonia were treated by Guillaumin? in 1940, and a key
to them was published in 19483. The Hawaiian species were revised by Sherff* in
» Pritzel, E. In Engler & Prantl’ e Nat. Pflanzenfam. ed. 2. 18a:273-281. 1930.
Bull. Soc. Bot. Fr. 87:333. 1
ZK Aen et Synopt. Nouv. Calédonie. Wege pp. 134-136. 1948.
* Field Mus. Nat. Hist. Bot. Ser. 22:467.
*An investigation carried out in the "eger laboratory of the Henry Shaw School o E ad
of Washington University, and submitted a thesis in partial fulfilment of the requirements for
the de imi " Doctor of Philosophy.
` t, S retine Institute and Museum, Auckland, New Zealand.
Mars ` 19,
Jul y (87)
[Vol. 43
88 ANNALS OF THE MISSOURI BOTANICAL GARDEN
1942. A preliminary account of the African species by Cufodontis? appeared in
1951, and a fuller revision? in 1952. A study of the East Asian species by Gowda’
was published in 1951, and a criticism of this work by Lif, in connection with the
Formosan species, followed in 1953. The species of the Malayan Archipelago will
be revised by one of the authors of ‘Flora Malesiana'?, and those of the Central
Pacific Islands will be dealt with in A. C. Smith's "Studies of Pacific Plants”.
The species which occur in Australia and New Zealand are revised in this study,
and an attempt is made to reconcile some of the divergent views of other workers.
History or THE GENUS
William Dampier, navigator and sometime buccaneer, was the first European
to collect a specimen of Pittosporum in the Australasian region. Between August
6 and September 2, 1699, he cruised along the northwest Australian coast in
H.M.S. Roebuck, from Shark's Bay northward to 18? 21' S. latitude. The few
specimens of plants he collected at Shark's Bay went to Dr. William Sherard and
are preserved in the Sherardian Herbarium at Oxford, England. They were described
by John Ray assisted by Sherard,'! and Plukenet.? Among them was the species
we now know as Pittosporum phillyraeoides DC. Linnaeus did not list the plant
in his ‘Species Plantarum’ (1753) and although Robert Brown examined the Shark's
Bay collection subsequent workers did not mention it.
The type specimens of the type species of the genus Pittosporum were collected
by Banks and Solander, who accompanied Cook as naturalists on his first voyage
of discovery. Between October 1769 and February of 1770 they collected four
species in New Zealand, and in May and June of 1770 they obtained another
species at Palm Island and Bustard Bay on the coast of Queensland, Australia.
During the voyage Solander had written descriptions of the plants collected which
were revised and systematically arranged prior to his death in 1782. Banks had
prepared excellent plates to accompany his Nene. but publication was delayed
and finally abandoned. In 1788 Gaertner!? published the description of Pitto-
sporum from the manuscript prepared by Solander, and described and illustrated
the fruits and seeds of two species made available to him by Banks!*.
The subsequent history of the collection of new species in Australia and New
Zealand, like the history of botanical discovery in these regions, may be divided
err. Bot. Zeitschr. 98:105. 1951.
oe —€— from Dr. C. G. G. J. Van Steenis, general editor of “Flora Malesiana”-
Y? Personal c
3 Hise. Pl. Vol. 3 5 and Apod: 1704.
12 Almalth. Bot.
13 Fruct. et Sem. A 1:286. £. 59. 1788.
14 The five species collected by Banks and Solander were: Pittosporum tenuifolium Gaertn., P.
umbellatum Gaertn., P. ralpbii Kirk, P. cornifolium A. Cunn., all from New Zealand, and P. ferru-
it. from Awink
1956]
COOPER—AUSTRALIAN AND NEW ZEALAND PITTOSPORUM 89
into two periods: 1, the period of investigations by travellers or naturalists at-
tached to voyages of discovery or survey; 2, the period of investigation by resident
botanists. Dampier, Cook, d'Urville, Wilkes, Robert Brown, and J. D. Hooker
are a few of the distinguished men who visited Australia and New Zealand during
the first period. The second period followed when systematic white settlement of
the two countries began. Bentham!5, F. M. Bailey**, Maiden?”, Lee!? and Willist?
have dealt adequately with the Australian collectors of both periods, and J. D.
Hooker?%, Cheeseman?! and 'Cockayne?? have given full accounts of the New
Zealand botanists.
The descriptions of species of Pittosporum from Australia and New Zealand
are scattered widely in the extensive literature on the floras of these regions, but
have been brought together from time to time by European and local workers. In
1839 Putterlick?? recognised thirty-six species of which twelve were recorded from
Australia and seven from New Zealand. Bentham?*, whose study was based on the
extensive Australian collections of Robert Brown, Preiss, Sieber, Allan Cunningham,
Ferdinand Mueller and others recognized nine species. J. D. Hooker? revised the
New Zealand species in 1853 and again in 1864 and recognized thirteen species.
His study was based on the extensive collections of Colenso, Sinclair, Dieffenbach,
Buchanan, Bidwill, Richard and Allan Cunningham, Hector, Lyall, Travers, and
others.
Hooker was far from satisfied regarding the status of several of the species,
and wrote?9: “I have restored the var. reflexum of Fl. N.Z, as a different species,
but with some doubt: better specimens are wanted of both”; and again”, “It
is with great reluctance that I add another Pittosporum to the series, already very
difficult of discrimination, that consists of Zenuifolium, colensoi and fasciculatum.”
Kirk?8 published a revision of the New Zealand species in 1872, based on his
own extensive collections and field observations, He did not have access to the
early literature on the genus, or to the important collections at the Royal Botanic
Gardens, Kew, but in spite of these handicaps he made a very valuable contribution
Wa Austr, 1: age o i
Proc. Roy. io ie 8:xvii, xlv.
17 Jour. & Proc. Roy. Soc. N.S.W. 42:60. rd Ist. Suppl. Rept. Austr. Assoc. Adv. Sci. 13: 224.
1912. 2nd Suppl. Jour. & Proc. perte Soc. N.S.W. 55: 150. 1921
Ea i .
19 Vict. Nat
1863.
25 Fl. Nov, Zel. 1: CH 1853; Handb. N.Z. Fl. p.18. 1864, and p. 725. 1867.
SZ loc. cit. p.21. Soe
loc. cit. p.725. 1867.
28 N.Z. Inst. "oc PC 4:260. 1872.
[Vol. 43
90 ANNALS OF THE MISSOURI BOTANICAL GARDEN
to our knowledge of the New Zealand species. Cheeseman??, who worked under
similar difficulties, recognized nineteen species in 1906 and twenty-three in 1925,
but expressed doubts regarding the status of many of these entities
The most recent study which is of importance to us is that of the small-leaved
species of Pittosporum in New Zealand, published by Laing and Gourlay?? in 1935.
This was based on extensive field observations and a considerable number of speci-
mens cultivated in the authors’ gardens, and includes complete descriptions of
seedling and juvenile stages. They observed the similarity of one of the small-
leaved New Zealand species of Pittosporum to the Australian species of Bursaria
but left the question of nomenclature changes to future workers. Pax?! treated
the genus as a whole for the ‘Die Natürlichen Pflanzenfamilien’ in 1891 and
recognised the species described by Bentham for Australia and by J. D. Hooker
and Kirk for New Zealand. Similarly, Pritzel3?, in the second edition of “Die
Natürlichen Pflanzenfamilien', adopted the Australian species from Bentham and
subsequent authors, and the New Zealand species from Cheeseman, with little
change.
Economic Uses
From the wealth of observation published on the uses of different species of
Pittosporum two generalizations can be made: (1) that certain medicinal and
poisonous properties are common throughout the range of the genus; (2) that
these properties were known to several primitive races of people who lived in com-
munities which were widely separated, if not completely isolated.
The Australian aborigines soaked the seeds and pulp of the fruit of P.
bbillyraeoides in water and used the bitter infusion internally for pains and
sprains?, The Filipinos used a decoction of the leaves of P. pentandrum as an
aromatic bath after childbirth, and the powdered bark of the same tree as a
febrifuge and for bronchitis. They also used the fruits of P. resiniferum as a
universal medicine?*, The Malayans applied the leaves and roots of P. ferrugineum
as poultices in the treatment of malaria??, and some South African natives employed
the bark of P. viridiflorum* as a febrifuge and as a bitter emetic. Again, on
Réunion Island the natives used the bark and leaves of P. senacia in medicine?”
e Malayans macerated the leaves and fruit of P. ferrugineum for fish
poison?5, and the Javanese treated P. ferrugineum and P. javanicum for the same
29 Man. ne Fl. pp. 51, a 1134. 1906; ed. 2, p. 486. 1925.
:44, 1935
30 Trans . Soc.
um Engler & ëch: Nat. ona. 3 (2a):106. 1891.
32 loc. cit. ed. 2 :265 30.
aiden, For. A prr 31:161.
34 Valenzuela, Con Santos. jour. P hilipp. ~~ a 34:34. 1947.
ha and
5 Burkill, Dict. Econ. Prod. Malay Pe cag 2:1763.
36 Githens, Drug Pl. Afr. p.100. 194
37 Niederlein, Resources Te des olen. Franc. p.55. 1902.
38 Burkill, loc. cit. 1935
1956]
COOPER—AUSTRALIAN AND NEW ZEALAND PITTOSPORUM 91
purpose??, The Fijians used the fruits of P. arborescens, P. brackenridgei, and P.
rbytidocarpum as a fish poison, the fruits of P. rhytidocarpum as a dye, the leaves of
P. arborescens as a remedy for stomach troubles, and the bark and leaves of P.
arborescens and P. brackenridgei for witchcraft. A cherished sheet of P. arborescens
in the herbarium of the Department of Agriculture, Suva, Fiji, is from a plant
which was "Exhibit A" at the trial of a poisoner charged with ritual murder.
It is possible that these medicinal and poisonous properties will be of economic
value in the future. For example, the bitter properties of P. phillyraeoides (Austra-
lia) and P. viridiflorum (South Africa) have been mentioned. Since the dawn of
history all sorts of virtues have been ascribed to bitter or "strong" medicine, but
bitters still have a place in medicine as general tonics, in the control of diarrhaea,
and in wound dressings.
The presence of saponins has been reported by a number of workers. Klein?
states that the following species contain the saponin “pittosporin” of unknown
formula. He gives no reference, but the information is probably taken from
Greshoff*!, ;
REGION SPECIES MATERIAL TESTED
Sino-Japanese Region Pittosporum tobira Leaves
Madeira-Teneriff Pittosporum coriaceum Leaves and bark
— Pittosporum densiflorum Leaves
Malayan Archipelago Pittosporum javanicum Bark
S Pittosporum floribundum Bark
South Af
i Pittosporum viridiflorum Bark
Pittosporum erioloma Leaves
: [aie phillyraeoides Fruit
A
te Pittos rbombifolium Leaves
Pittosporum undulatum Leaves and bark
Pittosporum cornifolium Leaves
New Zealand Pittosporum crassifolium Leaves
"t Pittosporum eugenioides Leaves
Pittosporum buttonianum Leaves
ED tpe
4, Hayne, Nutt. Plant. Ned. Ind. p. 688. 1927.
a 4 Han er Pflanzenanalyse. 37:1135. 1932.
Kew Bull. p. 414. 1909,
[Vol. 43
92 ANNALS OF THE MISSOURI BOTANICAL GARDEN
The presence of saponins probably accounts for most of the medicinal and
poisonous properties of the species. Githens* says:
The saponins constitute a ob large and diverse Led of teh mg have
the property nid using foaming when added t re ER e m and P
viridiflorum a dp Sen as piss "eee in Se ir pits countri e The "
find an extensive use in shampoos, toothpast <n similar cosmetic prepara-
tions. Some have no marked Sad: on the Mods, o: of them Beni
nausea or vomiting. This is accompanied by scetur secretion in the a-
tory passages and a greater c fluidi « of the mucus, leadin Wa f^ y jocis nan d EC
coughs or expectorant actio nausea is often accompanied by pan?
eadin
leading to lowering of febrile scs sa Saponins are therefore used as
emetics, expectorants and febrifuges. Many saponins act on the gills of fi Ze
interfere wit respira ration and are therefore eg as gus geng A few sapo-
Omer nk locally irritant and act as purgati X escam). cause.
deat
Saponins have been found suitable for insecticides, but neither the Department
of Insecticides at Rothamstead Experimental Station, England, nor the Section for
Insecticide Investigations at the U. S, Department of Agriculture Research Station,
Beltsville, Md., has examined any species of Pittosporum. The only information I
ve on their insecticidal properties is to be found in “A Survey of Plants for
Insecticidal Activity” by Heal and others??, These workers found that aqueous
extracts of the branches, leaves, and roots of P. semacia were toxic to American
cockroaches when injected into the bloodstream, but were inactive against German
cockroaches and milkweed bugs. Alcohol and petroleum ether extracts of P. senacia
— leaves were toxic to black carpet beetles, but non-toxic to other insects
. An extract of P. viridiflorum bark was non-toxic to all insects tested.
pedo ferrugineum of Malaya and Australia, and P. arborescens, P. bracken-
rid gei and P. rbytidocarpum of Fiji appear, from their record as fish poisons, to be
more promising than the species tested.
The essential oils of several species have been examined and described, and the
papers are listed:
kbl:
Pittosporum monticolum.................. A. 5 Faime LA Pharm. Weekblad,
Malayan Archipel,
I. Garcia-Reyes, in Rev. Filip,
M Med. Farm. 28:448. 1937.
y = Cornworth & J. C. Earl,
Pittosporum resiniferum................
Australia...................... Pittosporum aadnlatun a Jour. Roy. Soc. NSW.
72: 249. 1939.
: C. L. Carter & W. V. Heazle-
Pittosporum eugenioides... wood, in Jour. Soc. Chem. Ind.
New Zealand 68: 34. 1949.
Pittosborum tenuifolium SESS SUE A. J. Calder & C. L. Carter,
loc. cit. 68:355. 1949.
42 Githens, loc cit. 1949.
53 Lloydia 13:89. 1950.
1956]
COOPER—AUSTRALIAN AND NEW ZEALAND PITTOSPORUM 93
Essential oils** usually are the source of scent in flowers, and have many uses
as flavors and condiments in food. Frequently they have the property of checking
bacterial growth and are used for food preservatives, and even as wound dressings.
They are also valuable in medicine to regulate intestinal movements and for
respiratory, kidney and urinary infections. In 1889 Maiden recommended the
cultivation of P. undulatum on a commercial scale for its essential oil, but his
suggestion has never been followed, and the possibilities of this and other species
remain unexplored.
e gum-resins of P. bicolor, P. rbombifolium and P. undulatum of Australia
and of P. eugenioides and P. tenuifolium of New Zealand have been described by
Maiden**, who suggested that they might be found useful in medicine.
The iei of the Australian species have been described by Mueller*”, Bailey*?
and Maiden*®, and the woods of the New Zealand species have been dealt with by
Kirk5? and others. They are not available in sufficient size or quantity for com-
mercial use but are much esteemed for special purposes. The wood of P. undulatum
is preferred by some to any other timber for golf sticks?!, The wood of P. bicolor,
which was once used for clubs or “waddies” by the aborigines of Tasmania, is now
much esteemed for axe handles and billiard cues??. The replacement of ““waddies”
by billiard cues will be deplored by all lovers of the truncheon and shillelagh.
Pittosporum has made a small but valuable contribution to horticulture. The
cultivation of species of Pittosporum began when P. coriaceum was brought to
England from Madeira in 1787 and introduced into gardens by James Webster; P
undulatum was introduced into England from Australia by Sir Joseph Banks about
two years later; and P. tobira was sent to Kew from Canton, China, in 1804.
Splendid living collections are maintained today at the Royal Botanic Gardens,
Kew, the Villa Thuret, in southern France, and Golden Gate Park, San Francisco,
California. Many Australian and New Zealand species are represented in
collections and are grown in the surrounding districts as street trees, garden orna-
mentals, hedge plants, and shelter trees, The Australian P. undulatum is probably
the most widely cultivated and has been reported in gardens in Ceylon, Italy, the
Azores, France, England, the United States (Florida and California), Bermuda,
Hawaii, India, China, New Zealand and, of course, Australia. In the Azores it was
introduced as a shelter tree for the orange groves and is now naturalized. In
Bermuda and Hawaii it has also become naturalized and is an attractive addition to
the local flora,
Finnemore, H. The paid pp. 347-349. 1926.
d Maren i Austr. p.292, 1889.
ustr. Assoc. Adv. Sci. sae’. 1893.
« * Select Cat Zeen Pl. p.258. 1881.
pa oods. p.11. 1888.
so Useful Nat. Pl. Austr. p.588. 1889.
59 N.Z.
[Vol. 43
94 ANNALS OF THE MISSOURI BOTANICAL GARDEN
The widespread use of the Australian and New Zealand species in horticulture
has been accompanied by the usual nomenclatural problems. Horticulturists have
created, and persist in using, several specific epithets which have been ignored or
reduced to synonymy by taxonomists concerned only with the "wild forms". I
have attempted to solve the nomenclatural problems by citing published horti-
cultural names in the synonymy of the species concerned and in the discussion
following the description.
TAXONOMIC CRITERIA
The principal criteria for the classification of the Australian and New Zea-
land species of Pittosporum are based on comparative morphology and geographic
distribution. The chromosome number of only one species, the Chinese P. tobira
(2n = 24), is known”, and the embryology of the genus has not been investi-
gated”*, The anatomy of thirty-six species of Pittosporum, and a representative of
each of the allied genera, Hymenosporum, Bursaria, Sollya, and Citriobatus, has
been examined by Guenot, who commented:9? “La structure anatomique du genre
Pittosporum s'est montrée dans les espéces que nous avons examinées d'une con-
stance remarquable."
Gowda”* examined the pollen of twenty-six species of Pittosporum and a
representative of each of the other genera in the family, and found that they varied
in size but did not show any marked difference in form and sculpture. Lucy
Watson-Smith?" has informed me that the New Zealand species vary in size and
thickness of the exine but generally are very much alike. In the preceding chapter
on economic uses I mentioned that certain chemical properties of the genus are
remarkably constant.
A study of living and herbarium specimens has convinced me that Pittosporum
is an assemblage of very homogeneous forms and that many of its morphological
characters are constant throughout its range from Hawaii to South Africa.
the evidence suggests that it is a tropical and subtropical genus which is most
specialized and reduced at the extreme limits of its distribution, especially at the
southern boundaries in Australia and New Zealand.
MORPHOLOGY
SHOOT SYSTEM:
The Australian and New Zealand species are evergreen trees and shrubs with
verticillate and simple branching. Gowda?9 described the behaviour of the leading
shoot as follows:
53 Darlington, C. D. and Janaki Ammal, E. K. Chromosome atlas of cultivated plants. p. 112. 1945.
5 Johansen, D. A. Plant embryology. p. 174. 1950
" wen n ^i eg ibution à l'étude anatomique des Pittosporaceés. Doct. Diss. Univ. Paris. Ser.
. 523, p.5
56 SEN ës Zeg 32: 277. 1951.
Personal
nal communication.
38 loc. cit. p. 265.
1956]
COOPER—-AUSTRALIAN AND NEW ZEALAND PITTOSPORUM 95
1. Vegetative in the f
a. Producing ind Ji d in the second year and also a new
leading vegetative shoo oots the lower axils; or
b. Producing leafless peudoermini pa odia the second year from
the buds congested at the apex, and also a new leading vegetative shoot
r shoots.
2. Vegetative and terminating in an inflorescence the first year, and the second
year producing a new leading vairar shoot or shoots from its lower axils.
A plant of the Australian P. undulatum was observed in the Linnaean house
of the Missouri Botanical Garden from February 1952 until April 1953. In
February inflorescences developed in a terminal position on leading shoots. Dur-
ing April and May buds appeared in the axils of the leaves beneath the inflorescences,
and some of these developed rapidly and assumed a leading position while others
developed more slowly and formed a whorl of lateral branches. Similar behaviour
was observed in the Chinese P. tobira and on both plants two phases of growth,
reproductive and vegetative, were accomplished in the growing season.
The plants bore female flowers which were not fertilized, and the dead flowers
were forced to one side and shed when an axillary shoot assumed a leading position.
On several branches of both po one or two axillary vegetative shoots developed
at the same time as, an y below, the terminal inflorescences, In several
instances one of these vegetative shoots immediately assumed a leading position and
the inflorescence was left behind in a lateral position, On other branchlets the
vegetative shoots formed an irregular whorl and the inflorescence was left behind
in a terminal position.
An examination of herbarium material of P. undulatum showed that the
vegetative buds remain dormant in the axils of the subtending leaves when fruit
develop and the cluster of fruit persists for some time as a terminal structure.
Herbarium material of P. umbellatum, P. tenuifolium, P. crassifolium, P. corni-
folium, and P. dallii showed that where a terminal inflorescence developed fruit, the
fruit stalks were occasionally forced to one:side by the leading shoot and persisted in
a lateral position.
Agnes Arber®® has remarked that in shoot systems, “the subordination of
parent shoots to lateral shoots is, indeed, quite common; it is the essential feature of
all sympodial and cymose branching.’
The development of the leading vegetative shoot prior to the inflorescence
shoot involves a change in timing which has occurred in the allied Australian genus
Sollya. In S. heterophylla the inflorescence develops in a lateral position but is not
subtended by a leaf. There is a leaf on the side of the shoot opposite to the in-
florescence, however, indicating that the shoot above the inflorescence is really a
lateral axillary structure.
In Pittosporum divaricatum and P. crassicaule of New Zealand the branchlets
are almost spinose, but bear leaves and occasionally an inflorescence. Spines are
% The natural philosophy of plant form. p.94. 1950.
l [Vol. 43
96 ANNALS OF THE MISSOURI BOTANICAL GARDEN
well developed in the Australian P. o’reillyanum and the allied Australian genera
Bursaria and Citrobatus, and their derivation from short shoots is indicated by the
occasional presence of leaves on them.
In the New Zealand Pittosporum divaricatum, P. crassicaule, and P. anomalum
the flowers are borne on "arrested branchlets" which may attain several millimetres
in length. These are lateral structures and appear to be miniature forms of the
reproductive shoots on which the flower stalks are borne in many other species.
In the Australian P. phillyraeoides and P. bicolor, and in the New Zealand P.
obcordatum and P. rigidum, the inflorescence consists of a solitary flower, or a
fascicle of flowers, which is usually borne in the axil of a leaf. Pittosporum
phillyraeoides rarely has several flowers in a cyme, and P. obcordatum and P.
rigidum frequently have flowers on minute arrested branchlets. The New Zealand
P. tenuifolium and P. buttonianum bear both terminal and axillary inflorescences
which consist of 3 fascicle of flowers or of a solitary flower.
In all species mentioned the flowers may be considered as terminal on a shoot
which is reduced and sometimes is represented only by the scales at the base of the
flower stalks. Occasionally, in P. tenuifolium and P. buttonianum the shoot is well
developed and bears a whorl of cataphylls or 1-2 leaves beneath the terminal
fascicle of flowers, Rarely, the subtending cataphylls and leaves are suppressed
and the inflorescence appears to be compound. Other aspects of this reduction in
the reproductive shoot are discussed below.
The presence of terminal inflorescences, axillary inflorescences, and of both
terminal and axillary inflorescences is fairly constant within a species or sub-
species and is a valuable criteron for classification. However, it must be borne in
mind that the inflorescence may be pushed from a terminal to a lateral position by
the development of a leading shoot. Furthermore, in some species several reduced
forms of the fertile shoot (e-8 a cyme, fascicle, or solitary flower) may be found
together on a branchlet.
LIFE FORMS:
The life forms of the Australian and New Zealand species and subspecies,
according to the system of Du Rietz®, are summarized in the following table:
In compiling this table difficulty has been experienced in fitting the species
to the various categories defined by Du Rietz. His definition of "trees" is holo-
xyles with a distinct main trunk remaining unbranched in its lower part", and of
"shrubs," “holoxyles higher than 0.8 m., not developing a distinct main trunk, with
the stem branched from its basal part above or below the soil surface”. Some
species of Pittosporum develop both a main trunk and branches from the base.
Pittosporum undulatum, for example, when grown in an open situation usually
has branches from the base as well as a main trunk. Difficulty has also been
experienced with the size classes of Du Rietz. Again Pittosporum undulatum may
9? Life forms of terrestrial flowering plants. Acta Phytogeogr. Suec. 31. 1931.
1956]
COOPER—-AUSTRALIAN AND NEW ZEALAND PITTOSPORUM 97
| eg and Lord}
Life Form Australian we Islands New. Zealand
BIVALVAE TRIVALVAE BIVALVAE | TRIVALVAE
Tall t
(8 - 30: m. in height) 6 É 2 2
Low trees
(2-8 m.) 1 2 4 3
Dwarf trees
(0.8 - 2 m.) 4 2 1
High shrubs
(2 ~ 8 m.) 4 3 2
Ordinary shrubs
(0.8 - 2 m.) I
Dwarf shrubs
(under 0.8 m.) Z ze 2
Epiphytic shrubs i is 2
be cited as an example: in barren exposed localities it is a shrub 2-3 m. high
but in favorable situations it is a tree 10-30 m, tall. Similarly, P. bicolor, P.
rbombifolium, and P. crassifolium are very variable in habit and size.
In spite of these difficulties the summary does show: (1) that the New
Zealand species of the series TRIVALVAE are conservative in form; (2) that the
New Zealand species of the series BIVALVAE are more diverse in shrub forms; and
(3) that the Australian species of the BIVALVAE are a relatively conservative group.
The Australian species belonging to genera allied to Pittosporum have 2-valved
fruits, and if these were included in the table the number of shrub species would be
much higher, and the number of life form classes would be increased by the addi-
tion of dwarf-shrubs, dwarf half-shrubs, and twiners with a woody base.
Although the life forms of members of the Pittosporaceae have evolved along
closely parallel lines in Australia and New Zealand, the New Zealand species are
placed without difficulty in the genus Pittosporum while the Australian species are
so specialized in other characters that they are classified in separate genera. A
Possible explanation of the different rates of evolution in the two countries is given
later,
Laing and Gourlay*! considered that the small-leaved New Zealand species of
the BIVALVAE are a natural group and commen
It should first be noted that all the species dealt with . . . are highly
rphic in length and rigidity of branchlets, leaf shape and size, and in
species . . has also
size and ships, from the corresponding adult plant.
than one adult form, and the juvenile may go through many changes.
9! Trans. Roy. Soc. N.Z. 65:46. 1935.
[Vol. 43
98 ANNALS OF THE MISSOURI BOTANICAL GARDEN
There are a number of species, however, which lie between the two extremes
represented by Laing and Gourlay's polymorphic small-leaved section of BIVALVAE
and the more conservative members of the BIVALVAE and TRIVALVAE. For example,
Pittosporum turneri is a shrub or tree 2—9 m. tall, with sharply divaricating and
matted branchlets at the juvenile stage and on the lower part of the adult stem, and
ascending branches on the upper part of the stem. The leaves are relatively small
below and large above, and the plant combines the distinguishing characters of the
two sections suggested by Laing and Gourlay.
Cockayne and Allan? recorded about 200 New Zealand species as showing
more or less strongly marked dimorphy or even polymorphy as they progress from
the juvenile to the adult stage, and they eet that an explanation based on
hybridism and the "somatic segregation" in hybrids of chimaeras of various
types might be the correct one, There is, however, no evidence to support their
hypothesis.
ONTOGENETIC DEVELOPMENT OF THE SHOOT:
The vegetative shoot develops from a bud which is protected by cataphylls.
The term "cataphyll"9? is used here to include both bud scales and transitional
leaves. Usually the latest-formed cataphylls become quite leaf-like but with
the elongation of the shoot they are shed and only a whorl of leaves near the apex
of the shoot matures. The sequence of cataphylls from one whorl of mature
leaves to the next is illustrated in fig. 1. for the Australian P. undulatum, the New
Zealand P. umbellatum, and the Chinese P. tobira.
Usually the cataphylls are missing from herbarium specimens but their scars
have been found in most species of Pittosporum that 1 have examined. In the allied
Australian genus H ymenosporum the cataphylls are shed, but in Sollya, Bursaria,
Pronaya, Billardiera, Citriobatus, Cheiranthera, and Mariantbus they are retained.
Stebbins?* and others have remarked that the presence of an apical meristem
and the serial differentiation of organs result in two types of ontogeny; one is that
of the various lateral appendages, starting from the time when they become dif-
ferentiated from the apical meristem of the shoot, and the other is that of the
embryo and seedling. Agnes Arber® pointed out that recent work by Miller and
Wetmore®™ showed that the development of the shoot from the meristematic apex
is merely a modification of the original pattern in the relevant part of the embryo.
The embryo has not been investigated, but there are many observations on
development of Pittosporum seedlings and juveniles.
Generally variations in size and shape are more marked in seedling and juvenile
leaves than in the cataphylls between two whorls of mature leaves, but ithe
resemblance between the two ontogenies is marked.
*
62 Jour, Ecol. 15:239, 245. 1927
- 63 Foster, i in puc . Jour. Bot. 16:475. E
6t Variations robin in plants. 1950.
$5 The Gees en hy of plant a x 162. 1950.
66 Amer. Jour. Bot. 33:1. 1946.
1956]
COOPER—AUSTRALIAN AND NEW ZEALAND PITTOSPORUM 99
A wn
P. UMBELLATUM NEW ZEALAND
Em,
P. UNDULATUM AUSTRALIA
EM
P. TOBIRA ASIA
Fig.. 1. haa uence of cataphylls and leaves of three species of Pittosporum.
Previous mature whorl of leaves at left; new whorl at right; caducous cataphylls
indicated Greg horizontal black line. About one-third natural size. Further explana-
tion in the te
[Vol. 43
100 ANNALS OF THE MISSOURI BOTANICAL GARDEN
The following summary of seedling, juvenile, and adult foliage in representa-
tive Australian and New Zealand species of Pittosporum and in Billardiera longi-
flora is based on the descriptions of Lubbock®’, Cockayne®’, Sainsbury%, Hardy",
Laing and Gourlay™, as well as my own observations.
Species Seedling leaves Juvenile leaves Adult leaves
Australia
P. pbillyraeoides Entire, lanceolate- As for adult Entire, linear-oblong
oblong
P. undulatum Entire, lanceolate- As for adult Entire, elliptic-oblong
oblong
B. longiflora Entire, linear Trilobed, obovate Trilobed, lobes serrate,
ovate
New Zealand
P. tenuifolium ` ` Entire, ovate As for adult Entire, oblong-ovate
P. umbellatum Entire, obovate Lobed, pinnatifid, Entire, elliptic
obovate
P. divaricatum Irregularly double, Lobed, oblong or obo- |Entire or dentate, cren-
serrate, linear vate to linear ate or lobed, linear-
— to linear-
oblon
P. eugenioides Entire, — Variously notched a Entire, rd
ellipt base, POR
From the above summary and the descriptions given later, it is clear that
marked changes in seedling, juvenile, and adult foliage are due to differences in
size and general outline, and frequently to variation in the leaf margins.
Following Arber??, I am using the term “heterophylly” to describe the occur-
rence of two or more different types of leaf upon one individual. Heterophylly
has been reported in almost all the New Zealand species of Pittosporum. It is slight
in the TRIVALVAE (e.g. P. tenuifolium) but is well marked in the BIVALVAE (e.g. P.
umbellatum and P. divaricatum). Several heterophyllous species of Pittosporum
have been reported from New Caledonia'?, but none are known from Australia.
The allied Australian genera, Pronaya, Marianthus, and Billardiera have species with
$7 A contribution to our knowledge of seedlings. p. 200. 1892.
88 N.Z. Inst. Trans. & Proc. 31:362. 1899; and M :265. 1901.
$9 N.Z. Inst. Trans. & Proc. 54:572.
bis Prei- ei Soc. Vict. 28: 240. 1916.
5 Trans. . N.Z. 65: 44. 1955.
e 7 Water plants. p.143. 1920.
Guillaumin, in Bull. Mus. Hist. Nat. Paris 25:214. 1919.
1956]
COOPER—AUSTRALIAN AND NEW ZEALAND PITTOSPORUM 101
lobed or notched leaves, however, and their resemblance to juvenile forms of the
New Zealand species of Pittosporum is striking.
Distinct sun and shade forms in P. divaricatum, P. crassicaule, and P. anom-
alum have been described by Cockayne’, and Laing and Gourlay™; and Michie has
supplied me with sun and shade forms of P. pimeleoides var. major. It is clear from
the records and specimens that leaves exposed to the sun tend to be entire, and that
shaded leaves are variously lobed, toothed, and pinnatifid; but exposure and shade
are obviously not the cause of heterophylly. Stebbins' discussed heterophyllous
leaves from the ecological viewpoint but admitted that similar lobed and pinnatifid
leaves in Quercus had no apparent selective value. Ashby™ studied the changes in
size and shape from leaf to leaf on a shoot of Ipomoea caerulea and remarked:
On the basis of these fragments of evidence it is suggested that lobing is
determined very early in leaf ontogeny by a growth €— which changes
. an
in concentration or composition during development, . . which is sup-
presed at low light intensities and also in short days with uninterrupted
nights.
Elsewhere, he summarized past research as follows:
p .
some plants no correlation has been discovered between leaf shape and
external conditions. In these plants (e.g. species of Ipomoea and Hedera) leaf
shape is a function of the age of the plant and the position at which the leaf
arises. Heteroblasticl'?] development is presumably controlled by unknown
internal factors. :
Both leaf shape and rate of heteroblastic development are influenced by
genes. For Gossypium the effects of some specific genes on heteroblastic
development have already been worked out.
In the New Zealand P. umbellatum the change from lobed and pinnatifid
leaves is abrupt, but in P. virgatum it is gradual; in fact, lobed leaves are frequently
found in the adult foliage. There are not two phases, juvenile and adult, in this
species, but a series of phases in which the leaves differ markedly in size and form.
Ross Michie, of Kaitaia, N.Z., a keen amateur botanist, has a number of plants of
P. virgatum in cultivation on which two, three and four types of foliage are present
at different levels on the trunk, and he observes that some plants develop entire
leaves after the first phase of lobed and pinnatifid leaves, while others pass through
two or three more or less distinct leaf forms before reaching the adult foliage.
Leaf forms of juvenile and adult plants of P. umbellatum, P. virgatum, P.
patulum, P. turneri and P. pimeleoides are illustrated in fig. 2, and measurements of
their flowers and fruits are given in the chapter on Taxonomy. There is some
similarity between the adult leaves of P. pimeleoides, and the juvenile leaves of P.
patulum, P. turneri, and P. virgatum, and some resemblance between the adult
"7 Rept. Austr. Ass. Adv. Sci. 13:219. 1912.
75 Trans. Roy. Soc. N.Z. 65:44. 1935.
5, Variation and evolution in plants. p.490. 1950.
xd pe —— 49:386. 1950.
H D 21 . .
79 The sah heveroblastic” is used by Ashby in the sense given it by Goebel (Organographie
der Pflanzen, ed.1. p.123. 1898) and is synonymous with "heterophyllous" in this discussion.
[Vol. 43
102 ANNALS OF THE MISSOURI BOTANICAL GARDEN
leaves of P. patulum and P. turneri and the juvenile leaves of P. virgatum. There
is closer similarity between the adult leaves of P. virgatum and the juvenile leaves
of P. umbellatum. In habit, P. pimeleoides is a slender shrub to 2.5 m. tall; P.
umbellatum is a spreading tree to 10 m. tall; and P. virgatum P. patulum, and P.
turneri are trees intermediate in height between these two extremes
In size of inflorescence, flowers and capsules, the species show a similar
gradient from P. umbellatum to P. pimeleoides. In other morphological characters
and in their geographical ranges the five species are distinct; but a comparison of
leaf forms, flowers, and capsules suggests that they may be of common origin and
represent a series in which increase, or more probably reduction, in size has been of
evolutionary significance.
Flowering on semi-juvenile and shade forms has been observed in P. virgatum,
P. turneri, P. patulum, P. pimeleoides, P. pimeleoides var. major, P. divaricatum, P.
obcordatum, P. crassicaule, and P. anomalum, and the phenomenon is not rare in
heterophyllous species??. The possibility that some of the reproductive juvenile
forms have become permanently fixed during the history of Pittosporum is sug-
gested to account for the resemblance in leaf form and leaf ontogeny, and may also
be relevant in the comparative interpretations of plant forms in the family. In
the diagram below I hesitantly present a representation of the hypothesis, adapted
from De Beer*!,
s
»
me
Or Ro
Keng
ONTOGENY
PHYLOGENY
Takhtajan® discussed similar phenomena in higher plants in relation to
phylogeny, and concluded that:
In the Miro of the organism new characters arise as hereditary changes
of the varied stages of their development, beginning from the initials and
acetate it ast ph The most differentiated and Boa ently,
least plastic are the last stages of devel s. nism or org
The | end, particularly the terminating stages, produce small in-
significant variations only. he last phases arise séyrimeerj of the leaf,
zy. hy of the flower, the increase and decrease in T size of the organs,
Es anges in the — stages of development give rise to various
kinds oe regressive pheno: well. Ha UNE set A a che I last stages of develop-
ent Go change can mees move on deep earlier stages of
more
esis. Just in this way great Pile TAE ies of the organs
E take place.
89 Cockayne. D eech of d bens: er Die Vegctation der Erde. ed. 2. 14:140. 1928.
"E were ind en ed.2. 19
82 Trans. Molotov sone Univ. ove 22: Se 1945.
1956]
des
for:
Vote in ‘the
gare
,$ 3 =
COOPER—AUSTRALIAN AND NEW ZEALAND PITTOSPORUM 103
Regarding embryological investigations from the standpoint of comparative
morphology Jepsen, Mayr, and Simpson*?
remarked:
oe ap ger git noe! are agreed on recognizing a principle of the
chrony, alter.
e of ontogenetic stages in the phylogeny of a
structure. Obvio ar his is gene ` a morphologist's phrasing of the Greter
conclusion that mutation may alter the relative velocity of ontogenetic
processes.
dé
AES mi
vert re AA
Tem
A 1 P. PIMELEOIDES null
JUVENILE. FORMS DULT FORMS
2. Variation in leaf forms of five species of Piffosporum: juvenile
at left; un Go at right.
One fourth natural size. Further
THE INFLORESCENCE:
The basic unit of the inflorescence of Pittosporum is the cyme. In P. undu-
latum (fig. 3) the flower-stalks bear one to four flowers.
is the terminal one, after which flowering is frequently from the
flower-stalk upwards, although the order is variable.
The first flower to open
base of each
"7 Genetics, paleontology and evolution. p.71. 1949.
[Vol. 43
104 ANNALS OF THE MISSOURI BOTANICAL GARDEN
Fig. 3. Variation in the inflorescence of P. undulatum,
Australia. X 114.
Gowda** placed the series TRIVALVAE before the series BIVALVAE in his key to
the genus as plants of the former have certain characters (e.g. large 3-valved cap-
sules, many seeds, funicles from the base to near the apex of the capsule valve)
from which the characters of the BrvALvAE were derived. He considered that the
evolution of the inflorescence in the genus “is from the well branched paniculate
[i.e. paniculiform] to the simple cymose type and eventually to the solitary flower
with all intermediate stages present.” But none of the East Asian or New Zealan
species of the TRIVALVAE have well-branched paniculiform inflorescences. In the
Australian and New Zealand species the following trends have been recognised:
1. Reduction from a cyme to a solitary flower, e.g. P. undulatum.
2. Reduction from a fascicle to a solitary flower, e.g. P. tenuifolium.
3. Condensation of a compound inflorescence, e.g. P. dallii.
Presumably the spreading compound inflorescence of P. ferrugineum and its
allies has been derived by aggregation of fertile shoots and reduction of subtending
leaves to cataphylls or bracts. The increase in the number of flowers in the in-
florescence has been balanced by reduction in the size of the capsule and in the
number of seeds.
ute. Wh
PUBESCENCE:
Hairs are most common on young shoots, leaves, and inflorescences; and
persist on petioles and undersides of leaves, pedicels, sepals and ovaries. The two
main types in the Australian and New Zealand species are icacinaceous and mal-
pighaceous hairs as defined by Heintzelman and Howard®. Forms intermediate
8% Joc. cit. p.271. 1951.
85 Amer. Jour. Bot. 35:43. 1948.
1956]
COOPER—AUSTRALIAN AND NEW ZEALAND PITTOSPORUM 105
between the two types are found. The absence of hairs, or the presence of hairs
on parts usually bare, is a criterion for the recognition of particular species or
subspecies. Hair color is also useful in the recognition of several species, e.g. P.
buttonianum, although in other species it is most variable, e.g. P. bicolor.
THE FLOWER:
Kirk®* reported dimorphic flowers in several New Zealand species of Pitto-
sporum, and Petrie? described male and female flowers of P. cornifolium. Several
notes on the sexual behaviour of the Australian species of Pittosporum were pub-
lished by Maiden?5, and the pollination of the New Zealand species was discussed by
son??,
: In most of the Australian and New Zealand species I have found flowers
which appear to be functionally unisexual. The male flowers have weakly capitate
to truncate stigmas, long styles, slender ovaries, oblong anthers 2-4 mm. long
borne on slender filaments. Usually the anthers are level with the stigma or
exserted. The flowers which I think may be female have 2-, 3- or 4-lobed capitate
stigmas, slightly shorter styles, plumper ovaries, apparently abortive, sagittiform
anthers 1-2 mm. long, borne on filaments 1 mm. or more broad at the base, taper-
ing distally. Usually the anthers are borne below the stigma. Flowers with oblong,
apparently functional anthers 3-4 mm. long, capitate stigmas, and plump ovaries
occur (i.e. hermaphroditic), as do flowers with 4 functional anthers 3-4 mm. long
and one apparently abortive anther 1-2 mm. long.
Ross Mitchie, of Kaitaia, N. Z., has observed the flowers of a number of
species over a period of years and considers that some plants have female and others
male flowers, but the female plants sometimes shed a little pollen and the males
occasionally set a few fruit. In a letter to Dr. Woodson” I recently recorded my
own studies as follows:
I have observed four self-sown trees of Pittosporum më der ssp
folium for three years in my garden at Blockhouse Bay, New Zealand, vin
there is an acre of rapidly regenerating coastal scrub and forest. All the trees
flower profusely in the spring (August to October) and two, which bear
flowers with functional anthers and slender ovaries, have not set fruit. The
other two trees, which bear flowers with aborted anthers and plump ovaries,
are laden with fruit each autumn and the seeds germinate freely—there being
of seedlings under both trees.
Over the same period I have watched the behavior of planted trees of P.
obcordatum and P. umbellatum in the Auckland Domain, adjacent to the
useum where I work. The trees appear to be female Ae the flowers tien
apparently ines anthers = plump zx wer fru "^ each ar
but there a “male” in the borhood. Possibly Ka
ege ta fr Po Gr to fertilize EC ovaries, but this irs
ren 7
I have received s ens of flowers and fruit from a planc. of the
Ben an P. wire ée? een in the grounds of the Middlemore Hospital,
near Auckland. The flowers have plump ovaries ge apparently aborted
86 N.Z. Inst. Trans. & Proc. 4:260. 1872.
87 N.Z. Inst. Trans. & Proc. 53:365. 1921.
88 For, Fl. N.S.W. 7:124. 1920.
59 N.Z. Inst. Trans. & Proc. 13:254. 1881, and 57:115. 1926.
9? Personal comm unication.
[Vol. 43
106 ANNALS OF THE MISSOURI BOTANICAL GARDEN
anthers. The fruit are fully developed and the seeds germinate each year.
There is no male plant of = species in the neighborhood, however, and the
source of pollen is not known. More observations on all the Australian and
nes. Misco species of Pittosporum are required to determine the extent of
Cufodontis®! stressed the value of the sepals as a criterion for the recognition
of species and subspecies and described and illustrated five types of sepal arrange-
ment. Only four of these types are found in the Australian and New Zealand
species of Piffosporum: 1, sepals free or slightly coherent at base, not imbricate,
e.g. P. dallii; 2, sepals free and clearly imbricate, e.g. P. bicolor; 3, sepals coherent,
e.g. P. melanospermum; and 4, sepals connate in a tube splitting into 2 parts, one
1- to 2-lobed, the others 3- to 4-lobed, e.g. P. undulatum.
Occasionally in P. phillyracoides and P. cornifolium one or two pairs of sepals
are coherent from base to apex and the arrangement of the sepals varies slightly in
most species examined. Size, shape, and pubescence of the sepals also vary, but
are as valuable as sepal arrangement in the recognition of species.
The usual number of sepals and petals is five, but extra sepals and petals have
been found rarely in P. revolutum and P. pbillyraeoides (Australia), P. erioloma
(Lord Howe Island), and P. crassifolium and P. umbellatum (New Zealand).
Petal color is constant in most of the Australian and New Zealand species of
Pittosporum and is a useful character in the classification of the species. For about
half of the species of the genus the flower color is not known, but in the remainder
the colors usually found are white, greenish-white, and yellow. Red or purple
flowers have been reported in three species in New Guinea, three species in New
Caledonia, one species in the Tonga Islands, one species in Australia, and in most
of the species in New Zealand. In the allied Australian genera there are a number
of species with red, purple, and blue petals, and these colors seem to be associated
strongly with small leaves, heterophylly, and reduced forms.
THE FRUIT:
Gowda® recognized two major series in the Asiatic species which he termed
the BIVALVAE and the TRIVALVAE, according to the carpel number. Cufodontis??
did not discuss valve number but it is clear from his descriptions and illustrations
that all the species he dealt with belong to the BrvaLvaE. The character is some-
what variable, but the group to which a species belongs can be determined if suf-
ficient fruits are available and cognizance is taken of associated characters.
Cufodontis?* considered that valve shape is a valuable character and recog-
nized ge: basic types based on cross-sections of the valves: -
. Valvae convexae—more or less ev. = convex or with a flat middle Ee
x Valvae Ar enu M convex edges lies a concave middle
3. Valva ae—between convex borders is a longitudinal ed je E
4, Velas EM », but with a longitudinal groove down the dorsal thickening.
91 Osterr. Se — 98:109. 1951.
cit.
94 loc. cit. ein 1951.
NEW
1956]
COOPER—AUSTRALIAN AND NEW ZEALAND PITTOSPORUM 107
As the four types represent the variation in cross-sections of two-valved fruits
it was necessary to examine as many three-valved fruits as possible to determine
whether the classification applied to these. Generally the four basic types are
present in the TRIVALVAE but within some species more than one valve type is
found. A rapid survey of the valve shape throughout the range of the genus gives
the impression that the character is useful for the recognition of the Hawaiian
species as a whole; the capsule valves of the Hawaiian species have a distinct lon-
gitudinal thickening with a median longitudinal groove. The character is also of
value in the classification of species and subspecies.
Fig. 4. Ovary sections of P. crassifolium (left) and P. engeni-
oides (right), New Zealand. Resin canals indicated by hollow
circles; vascular strands indicated in black. Lu
Moore and Adams? described and figured a papery endocarp in the New
Zealand P. dallii, P. eugenioides, and P. anomalum. Fruit of P. crassifolium and P.
eugenioides both show (fig. 4) a more or less complete circle of large resin canals
lying outside, rarely inside, the vascular supply which branches to all parts of the
valve. In P. eugenioides the canals fuse more than they do in P. crassifolium and
finally the endocarp, vascular supply, and mesocarp separate in the former species.
Apart from the valve number and the resin canal development, no marked dif-
ferences were found in the anatomy of the fruits of the two species. On boiling
the fruits of P. crassifolium and P. faircbildii, Y have found the endocarp partly or
completely free from the vascular supply and mesocarp, and the separation of the
tissues can be effected without difficulty in several species. No great significance
can be placed on the presence or absence of a papery endocarp, although it is a
useful character for the recognition of P. dallii, P. eugenioides, and P. anomalum.
Pittosporum dalli and P. eugenioides may be related but there is nothing apart
from this character to suggest a close relationship with P. anomalum; in fact all
three are isolated and highly specialized members of the genus.
Gowda® used the placenta and funicles for the recognition of complexes of
95 Trans. Roy. Soc. N.Z. 77: 250. 1949.
96 Joc. cit. 280. 1951.
[Vol 43
108 ANNALS OF THE MISSOURI BOTANICAL GARDEN
East Asian species, and Cufodontis?" has illustrated these characters for the African
species. In the classification of the Australian and New Zealand species they are
valuable, although in most species there is some variation in the length and thick-
ness of the placenta and in the size and arrangement of the funicles, Emphasis is
placed on the placenta and funicles as characters rather than the ovules, because
the funicles persist whereas the ovules are frequently disarranged or missing on
herbarium material.
GEOGRAPHY
The literature on the past history of the Australian and New Zealand flora
and fauna is extensive and the conclusions of the most eminent authors frequently
conflict. Raup?5 and Cain®® have suggested that such varying points of view
might be unified if plant geographers would use more inductive methods. it
this good advice in mind, I have presented below the facts I have regarding the
distribution of Pittosporum and its allies in Australia and New Zealand. I have
included information about the distribution of the Pittosporaceae elsewhere as I
am convinced that the family must be studied on a world basis, not on a regional
one.
DISTRIBUTION OF THE PITTOSPORACEAE:
As at present understood the family Pittosporaceae comprises nine genera and
about 276 species, the distribution of which is illustrated in fig. 5.
The species were defined originally by divers botanists, and it is improbable
that they exercised taxonomic judgment to the same degree. A semblance of
uniformity has been introduced by the general treatments of Pax! and Pritzel!,
but further study will undoubtedly result in some of the species being relegated to
synonymy and others being separated into new categories. Indonesia, New Guinea,
and the Melanesian Islands will probably yield more forms when their floras are
better known, and the revision of the Polynesian Island species will shed further
light on the evolution of the family in the Pacific area.
From the figures available it is possible that the main centre of development
is Australia (9 genera and 48 species) and that important secondary centres are
the Hawaiian Islands (1 genus, 23 species), New Caledonia (1 genus, 46 species),
New Zealand (1 genus, 20 species), southeast Asia (1 genus, 52 species), and
Africa (1 genus, 10 species).
DISTRIBUTION OF HIGHER CATEGORIES:
Bentham??? Pax! and Pritzel!™ recognised two sub-families in the Pitto-
9 Fedde's Rep. Sp. Nov. 551:27. 1952.
98 Assoc
101 Engl. & Prantl, Nat. Pllanzenfem. ed. 2. 18a:265. 1930.
102 In Benth. et Hook.f., Gen. Pl. 1:131. 1862.
104 Joc. cit. 1930.
1956]
COOPER—AUSTRALIAN AND NEW ZEALAND PITTOSPORUM 109
AINUMBERS OF PITTOSPORUM SPECIES
DISTRIBUTION OF SERIES TRIVALVAE 9 A CUM ——
P lean” "m b ne E i r a:
l
so La kd LJ
ee O E
Fig. 5. World distribution of the Pittosporaceae. Figures indicate the number of Pitto-
sporum species in that locality. Letters represent the following genera: A, Billardiera (8
species), B, Bursaria (2 species), C, Cheirantbera (4 species), D, Mariantbus (16 species),
d Pronaya (1 species), F, Sollya (2 species), G, Citriobatus (4 species), H, Hymenosporum
species).
sporaceae, the PITTOSPOREAE, characterized by woody and leathery capsules, and
the BILLARDIEAE, distinguished by succulent berries. In the PITTOSPOREAE, the
genus Pittosporum is wide-ranging, Hymenosporum occurs in Australia and New
Guinea; and Mariantbus, Cheiranthera and Bursaria are known only from Australia.
In the second subfamily, the BILLARDIEAE, the genus Citriobatus occurs in Austral-
ia, Java, Luzon, and the Celebes, while Sollya, Billardiera and Pronaya are known,
so far, only from Australia. The regions where members of the two subfamilies
have been found together are Australia, Java, Luzon, and the Celebes.
In the genus Pittosporum, the series BIVALVAE ranges from Hawaii to Africa
and is present in all regions where the family is found, except the Ryu-kyu, Lord
Howe, and Norfolk islands. The series TRIVALVAE is known from the mainland of
ina, Formosa, the Ryu-kyu and Bonin islands, Japan, South Korea, New Guinea,
New Zealand, Lord Howe and Norfolk islands, Fiji, and Hawaii. The regions
where members of the two series of Pittosporum have been found together are
Upper Burma and Assam, western and southwestern China, Formosa, the Bonin
Islands, New Guinea, New Zealand, Fiji, and Hawaii.
I do not know of any region where both subfamilies of the Pittosporaceae and
both series of Pittosporum occur.
[Vol. 43
110 ANNALS OF THE MISSOURI BOTANICAL GARDEN
DISTRIBUTION OF WIDE-RANGING SPECIES:
The wide-ranging members of the family and the regions in which they occur
are:
Hymenosporum flavum Australia, New Guinea.
Pittosporum ramiflorum Philippines, Java
ar. pa rviflorum New Guinea, Solomon Islands.
eum Australia, New Hebrides, New rome Java, Sumatra, Nicobar Islands.
P. podocarpus——C hina, Indo- o- China, Burma, India
a.
ceylanicum— —Ceylon, southern India.
neelgherrense——Ceylon, es India.
m
m——Fo
var. hainanense——Indo-China, Hainan
napaulense——Nepal, Sikkim, Tibet, Bhotaun, Assam.
var. rawal pindiense——Si kim, Dehra Dun, Punjab.
floribundum ascar.
rrii——China, Indo-China, Siam, Burma.
johnstonianum——Ch ina, northern Burma.
asperm
Sou." ee Kata SALA de
a~
ki
2
R
#
»
5
a
?
Q
viridiflorum
ME Dadi
A (Yon. Ethiopia, central and southern Africa.
Gowdal% considered Piffosporum ferrugineum to be an extremely variable
and poorly understood species, but I have found it too coherent to allow separation
of the Australian, New Guinean, and Indonesian forms with the material available.
At present it is the only species of Pittosporum known which extends from Australia
to Asia but it does not reach the Asiatic mainland. Gowda! found that none of
the eastern Asiatic species of Pittosporum occurred in Indonesia, New Guinea, or
Australia, but Li! believed that Gowda was mistaken and that some species extend
from Asia to Indonesia.
I am tempted to speculate on the past land connections indicated by some of
these ranges, but it is possible that diaspores of the species have been carried to their
present areas by birds, hurricanes, and ocean currents. I must emphasize, too, the
probability that these populations will undergo further revision in the future.
DISTRIBUTION OF INTERGRADING SPECIES:
The diy of Pittosporum found in Hawaii are evidently very closely related.
Sherff198 w
Mu SCH ot the trouble in — a — key for Hawaiian Pittospora lies in
the fact that numerous subspecific ent ities, namely varieties and forms
105 Jour. gne Arb. 32: 322. 1951.
tos loc. cit.
% Jour. Wah. Ac Ad Sci. 43:43.
108 Field Mus. Nat. Hist. Bot. Ser. 22:472. 1942.
1956]
COOPER—AUSTRALIAN AND NEW ZEALAND PITTOSPORUM 111
(formae), occur. While these have their individual criteria by which they
usually can be recognized, they nevertheless afford so many intermediate char-
acters as to make en respective species seem to intergrade or even overlap.
Similar remarks have been made regarding the species of Pittosporum which
occur in New Zealand. J. D. Hooker has been quoted previously; Kirk! com-
mented of P. tenuifolium, P. colensoi, and P. fasciculatum: ‘‘These forms vary
considerably in all their parts, so that is would not be difficult to obtain a con-
nected series of specimens which should include the whole.’
In the taxonomic treatment I have discussed P. ellipticum var. decorum, which
appears to be intermediate between P. ellipticum, P. buttonianum and P. ralpbii
and have mentioned P. intermedium, which resembles large forms of P. tenuifolium
in foliage, while the capsule partakes of the characters of P. crassifolium and the
sepals and petals match P. umbellatum in size and shape.
Pittosporum rigidum, P. divaricatum, P. crassicaule, and P. anomalum contain
a bewildering series of polymorphic and heterophyllous forms most of which occur
in small areas at high elevations. Another series of intergrading forms occurs in
North Auckland, and has been variously treated by different authors; I suspect
that P. reflexum and P. pimeleoides are sun and shade forms of the same species but
have seen too few living specimens to be certain. The large linear-leaved forms of
P. pimeleoides resemble juvenile plants of P. virgatum, and some of the adult plants
of P. virgatum are very similar to juvenile plants of P. umbellatum.
The problem of intergrading forms in the Australian species of Pittosporum is
slight. Small-leaved forms of P. undulatum have been misidentified as glabrous
forms of P. bicolor and some are close to small-leaved specimens of P. revolutum,
but generally the species are easily recognised by their morphological characters.
Gowda!" indicated a number of closely allied species of Pittosporum in eastern
Asia, the most noteworthy being P. kerrii (Siam, China), P. floribundum (south-
ern India, Madagascar) , and P. ripicola (central and eastern Africa). Three groups
of intergrading species are mentioned: (1) P. mapaulense, P. floribundum, and P.
kerrii; (2) P. truncatum and P. heterophyllum; and (3) P. trigonocarpum and
P. sabnianum.
I have been unable to determine the frequency of intergrading forms in the
species of Pittosporum described from other areas, and any conclusions are pre-
mature. So far the evidence indicates that intergrading forms are more frequent
in the Hawaiian Islands and New Zealand than in Australia and Asia. Gowda!!!
recognised 52 species of Pittosporum in southeastern Asia, and the intergrading
species amount to about a seventh of the total. The proportion of intergrading
species in the New Zealand and Hawaiian species is much higher.
Proc. 4:262. 1872.
. Inst. Trans
i Jour. Arnold Aeb 2 284. 1951.
Mi Joe.
[Vol. 43
112 ANNALS OF THE MISSOURI BOTANICAL GARDEN
DISTRIBUTION OF ENDEMIC SPECIES IN AUSTRALIA AND NEW ZEALAND:
All species of Pittosporum in New Zealand are endemic, but only P. tenui-
folium and P. eugenioides are found throughout the country. The other New Zea-
land members of the TRIVALVAE and BIVALVAE occupy smaller areas. For example,
P. turneri is known only from one locality on the Central Volcanic Plateau of the
North Island, and a closely allied species, P. patulum, occurs in a few localities
in the mountains of northwest Nelson, in the South Island. Pittosporum fair-
childii is restricted to the Three Kings Islands, a group of rocks some 30 miles off
the coast, and P. dallii is known only from one locality where about a dozen plants
have been found. A number of the ranges of the New Zealand species are disjunct
and most of the small-leaved heterophyllous species have been found in small
isolated populations. For example, P. obcordatum is known only from three
isolated localities several hundred miles apart, the total population being about 15
trees.
All Australian species of Pittosporum are endemic, except P. ferrugineum
which is also found in the New Hebrides, New Guinea, and Indonesia. Pittosporum
bracteolatum is restricted to Norfolk Island and P. erioloma is known only from
Lord Howe Island. Pittosporum bicolor is most common in Tasmania, but occurs
in Victoria and New South Wales, An allied species, P. phillyraeoides, is found in
every state, except Tasmania. The most widespread of the remaining species is P.
revolutum, which extends from Queensland to Victoria, a distance of 1000 miles.
Within this range are the areas of P. rbombifolium, P. rubiginosum, P. undulatum,
and P. o'reillyanum. | Pittosporum venulosum is found on the coast of Queensland
farther to the north, and P. melanospermum has been collected from the same coast
and from the Northern Territory. No one species occupies the whole of the Aus-
tralian continent, but the majority of species occupy relatively large areas; and all
have continuous ranges, except possibly P. melanospermum. Furthermore, if the
populations of Norfolk and Lord Howe islands are excluded from the Australian
species, the number of individuals comprising an Australian species is far greater
than the number of a New Zealand population.
CLIMATE:
The climatic regions of Australia and New Zealand, according to Thornthwaite's
classification!?, and the number of species of Pittosporum in each are plotted in
fig. 6. The species of Pittosporum in the twelve climatic areas are listed below,
each species being indicated by its number in the present taxonomic treatment.
The concentration of Pittosporum species in eastern Australia (Queensland,
New South Wales, and Victoria) and the relative paucity of species in central and
western Australia, specifically the desert and steppe climate areas, is noteworthy.
Pittosporum phillyraeoides, the only species which is found throughout southern,
central, and western Australia, is xeromorphic as are some of the species of the
112 Geog. Rev. 28: 433. 1933.
1956]
COOPER— AUSTRALIAN AND NEW ZEALAND PITTOSPORUM 113
Climatic type* Australian Species Norfdlk and New Zealand
Lord Howe Islands Species
A A'r 24, 25, 27, 30
B A'w 24, 25,:27,/28, 29, 30
B B'r 9, 10, 24, 26 2 $4 EE e f
1271354413, 16, 17
18, 19, 25,21, 22, 23,
31,32
C Ale E 9, 28
C Bd 9, 11, 24, 26, 29
C B'r 9, 10; 24, 26
C B's 9
D A'w 9
D BA 9
D B's 9
E A'd 9 (coastal only)
E B'd 9 (coastal only)
* The first letter of the climate type indicates the vegetation transitions due to diminished effec-
e rainfall: (A) rain forest, (B) forest, (e mee (D) s teppe, and (E) desert. The second
lee indicates the ee Borges due iminished temperature vi inei (A') tropical
in forest, and (B’) temperate rain forest. e third letter Safe the seasonal distribution of
pini (r) pn vi pirstion at all seasons, (s) scanty rainfall in summer, mr ve in winter,
(w) scanty rainfall in winter, abundant in soke, and (d) dry at all seasons.
allied genera which occur in the desert and steppe. The development of
the allied genera mainly in the southern parts of Australia is linked with the
development of the so-called Australian elements in the flora and arid periods
in the Post-Pleistocene.!!? Stebbins!!* has presented evidence in favour of a hypo-
thesis that environments limiting or deficient in one all-important factor, moisture,
often have promoted rapid evolution. The variety of life-forms in the allied genera
is strikingly different from the general uniformity of Pittosporum and may be
the result of evolution under the stimulus of aridity. The northerly concentration
of the New Zealand species is illustrated by grouping the species according to the
two major geographical units. Until the close of the Pliocene period Cook Strait,
which separates the North and South Islands, was closed and there was no barrier
to the spread of species north and south.
Species found in the North Island only 11
Species found in the South Island only 2
Species common to both islands 7
20
Owing to its relatively small size, New Zealand has only one climatic type,
M3 Crocker and Wood, in Trans. Roy. Soc. S. Austr. 71:91. 1947.
114 Amer, Nat. 86:33. 1952.
> [Vol. 43
114 ANNALS OF THE MISSOURI BOTANICAL GARDEN
but generally the North Island has warmer summers and milder winters than the
South Island. Zotov!!5 studied the correlation between vegetation and climate in
New Zealand and commented:
On the continents where annual variation of temperature is great, the absolute
minimum value of temperature—i.e. degrees of frost ay ome a very
important limiting factor for the distribution of many species. In New Zea-
land, however, the minimum temperature does not appear to be of importance
or the great majority of species, since they reach their summer “limiting tem-
perature" in places well outside the boundaries of limiting winter temperature.
xe "UT" PITTOSPORUM SPECIES
- A i> IN EACH CLIMATIC REGION OF
GC at aes <—|— AUSTRALIA AND 2: SE
SW: A
: nam
d ied 3
e "X S ced 5
1 d > 13 `~ 2
ps". 1 EX ‘
D ea EP'd 1
cr WIS 8
vo AUSTRALIA
so lo [a TEE fool
Fig. 6. Number of species of Pittosporum in each climatic region of Australia and New
Zealand. For explanation see text and table on preceding page.
E
On Thornthwaite's!!8 maps of climatic types the regions occupied by the
species of Pittosporum outside Australia and New Zealand are generally in the rain
forest, forest, or grassland classes, for effective rainfall, and in the tropical and
temperate rain forest classes, for temperate efficiency. Over much of the range
of the genus the isothermal difference between extreme months is relatively small.
In the Pittosporaceae the largest plants are Pittosporum brackenridgei of Fiji
(tree to 25 m. tall), P. undulatum of Queensland, Australia (tree to 30 m. tall),
and P. viridiflorum of Africa (tree to 25 m. tall). These species occur in tropical
rain forests, The smallest plants are those of Marianthus procumbens of Australia
(a low shrub to 30 cm. tall) and its allies, Pittosporum anomalum of New Zealand
(semi-prostrate shrub to 1 m. tall) and its allies, and P. saxicola of Western Szech-
wan (prostrate shrub 0.3 - 2.0 m. tall) and its allies. These species are found in
115 N.Z. Jour. Sci. Tech. 19: 474. 1938.
M6 loc. cit. 1933.
1956]
COOPER—AUSTRALIAN AND NEW ZEALAND PITTOSPORUM 115
various habitats at the extreme northern and southern boundaries of the family
and are dwarfed in all organs.
In the evolution of these forms low temperatures may have been significant,
but it is also possible that aridity has played a role. For example, the small-leaved
xeromorphic P. pimeleoides is found on strongly leached podzolized and skeletal
soils where Agathis australis grows or formerly grew, and the small-leaved xero-
morphic P. divaricatum, P. rigidum, P. crassicaule, and P. anomalum occur on
skeletal mountain and volcanic soils.
GEOLOGICAL HISTORY:
The literature on past land connections of Australia and New Zealand is large
(Benson!", David!!8 119, Marwick!?, Merrill?!, Oliver!??, Skottsberg!?), and the
diastrophic paleontologic, faunal, and floristic evidence is substantial. It is generally
accepted that there have been no land connections between Australia and Asia in
Tertiary and recent times (David*?*, Diels!?5, Merrill?^) , and it is usually implied
or stated that the present land area of the Australian continent has remained above
water throughout Tertiary and Recent periods, save for temporary marginal incur-
sions. The scanty relics of Upper Cretaceous floras are sufficient to show that
flowering plants were established on the Australian land area prior to the Tertiary,
and studies of more recent floras indicate that the flowering plants experienced
significant climatic modifications in the post-Tertiary.
I can find no reference to fossil forms of the Pittosporaceae in Australian
paleobotanical papers. Cookson and Couper!?8 informed me that pollens of the
Pittosporaceae are not used as indicators in Australian and New Zealand Tertiary
pollen studies, and that their absence from published species lists has no signific-
ance. The well-marked morphological characters of the Australian species and the
large geographical areas of most of the populations indicate that the group has
evolved in isolation over a comparatively long period. I suggest that these
morphological and geographical characteristics are indicative, and perhaps typical,
of evolution under continental conditions.
In a recent discussion of the geological history of New Zealand with reference
to the origin and history of the fauna and flora, Fleming? emphasized that the
New Zealand area has long been a relatively isolated archipelago of islands. He
117 Gedenboek, Dr. R.D.M. Verboek, etc. p.53. 1925
115 Geology of the oo of Kael 3 SCH 1950.
ex s, Tonto 2:72.
15; NZ Jour. Sci. & Tech. 11: 202. 1929.
! Far Eastern = 2:66. 1942.
122 Your. Linn. Soc. Lond. Bot. 47: 99. 1925.
va Plant E World 1 18:129. 1915.
2 loc, . 15. 1950.
e GC s on SE in honor of W. A. Setchell. p.194. 1936.
126 loc. cit. 194
7 David, loc. i p. 650. t rage in Austral. Jour. Sci. 15: 47. 1952; Crocker and Wood, in
Trans. Roy Soc. S. Austr. 71:91.
= 28 Personal communications
Tuatara 2:72. 1949.
[Vol. 43
116 ANNALS OF THE MISSOURI BOTANICAL GARDEN
pointed out that the frequent changes in the size and relations of the islands may
have played a significant role in the formation of races species of plants and
animals. Marked climatic changes since the Cretaceous epoch are indicated by
faunal changes in the fossil record!9?, and several post-glacial, climatic fluctua-
tions are reflected in the sequence of pollens found by Von Post and Cranwell!?!
in peat deposits and in profiles of soils studied by Raeside!?2,
Arber??? found two dicotyledons in rocks of Jurassic or lower Cretaceous age,
and Couper" found Angiosperm pollens in coals of early Cretaceous-Eocene age.
Pittosporum leaves have been found in beds of lower Oligocene, lower Miocene and
upper Pliocene age by Oliver!35, and in beds of upper Pliocene or lower Pleistocene
age by McQueen!?6,
In the discussion of the distribution pattern of New Zealand specids of
Pittosporum it was noted that endemism is high, intergrading forms are common,
and geographic ranges are small. Apparently these characteristics are the result
of a long period of evolution in a changing and relatively isolated archipelago
under fluctuating climatic conditions, and I suspect that they are typical of most
species formed under insular conditions. It is possible, of course, that the differ-
ences noted between the species of Pittosporum occurring in Australia and those
found in New Zealand are due mainly to the continental outlook of Australian
taxonomists and the more insular attitude of New Zealand botanists!
In a discussion of the floras of Madagascar and New Caledonia, Good!"
observed that the relative situations of the two groups of islands to the continents
of Africa and Australia respectively are very similar, that the physiographic
structure of the main islands is curiously alike, that the climatic types correspond,
and that both possess floras with a large number of endemic species. Twenty
World (African-Asiatic-Australian) genera have complexes of endemic species in
Madagascar and New Caledonia, and Good cited Pittos porum as the most interest-
ing of them, a citation which I heartily endorse. He suggested, however, that
these curious similarities are the result of past land movements, or continental
drift, and this I must question. If the distribution of the genus Pittosporum is
used as evidence, the range of all species should be considered. Lists of closely allied
species given previously link the various centers of development of the genus and
suggest that extensive movements of the land horizontally are not required to
explain the present distribution of the genus in Madagascar and New Caledonia.
N.Z. Surv. Pal. Bull. 6. 1917. McQueen (Nature 175:177. 1955) said that the slab
studied by Arber came from Upper Cretaceous beds and that the earliest New Zealand angiosperms
are Cretaceous in age.
us NZ. Sci. Rev. 9: 5. 1951, and N.Z. Geol. Surv. Pal. Bull. 22 (in press).
za N.Z. Inst. Trans. & Proc. 23: 276. 1891; and 59: 287. 1928; Tuatara 3: and 1 and 8, 1950.
communication.
187 Blumea 6: 470. 1950. :
1956]
COOPER—AUSTRALIAN AND NEW ZEALAND PITTOSPORUM 117
Chevalier!?3 considered Pittosporum to be a South Oceanic genus and
remarked:
le grande foyer de dispersion du genre Pittosporum ce sont l'Australie et
b iles um Pacifique. Du reste, sur 10 genres que compte la famille des Pitto-
H H
sporées, 8 sont endémiques en Austr oe? be Lu re Pittosporum est aussi en grande
dix
partie pacifico-australien, le iéme e Tribeles phil. (= ane eboa Hook.
f.) avec une unique éspéce Tribeles au iral Philo (=Chalepoa antartica
onfiné à la partie extrêm d de l'Amérique (sud du Chili
sporum ont essaimé des terres australes vers l'Asie, deg part, et vers bat
d'autre part; une seul Pittosporée est parvenue en Amérique australe, à sa
pointe extréme
Engler!?? placed Tribeles in the subfamily EscALLONIOIDEAE of the Saxifraga-
ceae, however, and while that family may be related to the Pittosporaceae, there is
no certain evidence that the Pittosporaceae are represented in South America.
Furthermore, the development of the Pittosporaceae in Australia and New Zealand
is not evidence that the family originated in that part of the world. The deter-
mination of the center of origin for the family must await the progress of paleo-
botany, cytology, and possibly plant exploration. For example, further field work
in the New Guinea-New Caledonia area is likely to yield further novelties and
dr new light on the distribution of the Pittosporaceae.
INTERRELATIONS OF THE SPECIES
Methods by which a pictorialized scatter diagram is developed are explained in
Anderson. 41 and Anderson and Gage!!?. The figures listed in Table I are the
maximum measurements made of capsule width and the averages of the measure-
ments made of petal length, sepal length and petal width for the taxonomic
treatment. The averages are the means of the actual measurements made on the
erbarium material cited in the text.
measurements have been plotted as a pictorialized scatter diagram (fig.
7), to facilitate an understanding of the relationships of the species and subspecies
of Pittosporum. Petal width and capsule width were chosen as abscissa and
ordinate respectively because they varied consistently within each species and could
be measured accurately. The other three characters were indicated by rays from
each dot on the diagram. The number alongside each dot refers to the species in
the taxonomic treatment. Limits of the three grades of each character were chosen
so that extremes associated with higher values for petal width and capsule width
are indicated by long rays and extremes associated with lower values for petal width
and capsule width are represented without rays.
ull. Soc
139 Engl. & Pg be n Pflanzenfam. ed. 2. 18a: 213. 1930.
a A hybridization. 1949.
d Life. 1952.
42 Amer, edu 39:399. 1952.
[Vol. 43
118 ANNALS OF THE MISSOURI BOTANICAL GARDEN
At one extreme, the upper right-hand corner of each diagram, are the species
with petals and sepals which are markedly long and capsules which are three-valved
and markedly broad. In the New Zealand diagram most of the species at this
extreme are in the series TRIVALVAE. In the Australian diagram the species numbered
1 and 2 (P. bracteolatum of Norfolk Island and P. erioloma of Lord Howe Island)
are in the TRIVALVAE; the other species of this extreme are in the BIVALVAE, but
approach the TRIVALVAE closely.
At the other extreme, the lower left-hand corner of the diagram, are the species
with petals and sepals which are markedly short and capsules which are two-valved
and markedly narrow. In the New Zealand diagram most of the species at this
extreme are small-leaved, dwarf members of the BIVALVAE with small fascicles or
solitary flowers. In the Australian diagram most of the species at this extreme are
large-leaved tall members of the BIVALVAE with spreading compound inflorescences.
Species numbered 12, 15, 13, 16, and 14 on the New Zealand diagram (P. umbell-
atum, P. patulum, P. virgatum, P. turneri and P. pimeleoides respectively) show a
proportionate reduction in all characters illustrated, from the upper right-hand
extreme to the lower left-hand extreme, and may connect the series TRIVALVAE and
the small-leaved dwarf members of the svarvar. The variety of leaf-forms in
these five species has been illustrated previously and the suggestion has been made
that the leaf forms indicate the direction of evolution in the New Zealand species.
AUSTRALIA 1 NEW ZEALAND Y
E N
25 Y Y
=
S 20 Eé Y
z ^ i
S MN v
15 a Ki aM
w 27
3 AN ik Me
a y» M T
vu 10 108 y "
Te. me uou `
5 "ne! xe
210
15 20 25 30 35 15 20 25 30 35
PETAL WIDTH MM.
PETAL LENGTH MM. © » * ` TRIVALV
-106 104-80 78+ ava |
SEPAL LENGTH MM e BIVALVAE e
€ D
-5.4 52°32 302
éi Mere Scatter r Suai of information contained in Table I, sein d Meere
of a species of Pittosporum indicated by capsule width and by measurements petals and
Horizontal pé petal w videh: vertical axis, capsule width. Petal length and Ev length are
diagrammed by rays as as explained « on the diagram. Numbers beside dots refer to the position of the
species in the taxonomic treatm
1956]
COOPER—AUSTRALIAN AND NEW ZEALAND PITTOSPORUM 119
TABLE I
AVERAGE MEASUREMENTS OF THE AUSTRALIAN AND NEW ZEALAND
SPECIES OF PITTOSPORUM
Petal Petal Sepal Valve Capsule
Species length width . length number| width
mm. ` mm. cm
1. P. bracteolatum 17.0 4.0 10.7 3 2.5
2. P. erioloma 16.0 3.6 6.2 3 2.0
3. P. fairchildii 11.8 3.4 5.6 3 2.5
4. P. ellipticum 15.5 3 9.3 3 1.7
5. P. crassifolium 13.6 3.9 9.3 3 3.0
6. P. ralphii 14.8 3.2 8.7 3 1.3
7. P. huttonianum 15.6 IL Ké 3 2.0
8. P. tenuifolium
ssp. Zenuifolium 10.8 3.6 4.5 3 2.6
colensoi
9. P. phillyraeoides 10.1 215 2.6 2 1.9
10. P. bicolor 11.2 3.0 4.2 2 1.3
11. P. o’reillyanum 9.2 1.4 1.9 2 12
12. P. umbellatum 11.5 3.7 7.1 2 12
13. P. virgatum 10.0 22 4.6 2 25
14. P. pimeleoides
ssp. pimeleoides 9.7 1.6 4.0 2 8
ssp. major
15. P. patulum 10.0 2.4 4.9 2 1.1
16. P. turneri 8.1 1.9 4.0 2 8
17. P. obcordatum 5.8 12 24 2 7
18 um 9.8 2.0 3.9 2 1.0
9. P. crassicaule 5.4 1.4 L5 2 of
20. P. divaricatum 5.2 13 1.9 2 6
4L P lum 1.7 1,9 2 4
22 ornifolium 11.1 1.8 5.7 2 1.2
23. P. kirkii 17.6 2.9 8.6 2 1.7
24. P. revolutum 15.1 3.7 6.0 2 2.1
25. P. rubiginosum
ssp. rubiginosum 13.0 3.1 6.8 B 1.5
ssp. wingii
26. P. undul 13.7 3.8 7.9 2 1.4
27. P. venulosum 7.9 1.9 4.1 2 1.4
28. P. melanospermum 7.6 2.0 1. 2 1.5
29. P. rbombifolium 6.5 1.9 1.6 2 .8
30. P. ferrugineum 7,5 1.3 2.9 2 1.0
31. P. dalli 8.7 3.3 5.7 2 3
32. P. eugenioides 6.2 1.8 2.3 2 6
SE
Species numbered 22 and 23 (P. kirkii and P. cornifolium) are rather isolated,
very similar in their rays, and are both epiphytic. Species numbered 31, 32 and 21,
(P. dalli, P. eugenioides and P. anomalum), are somewhat isolated and are the
three species in which capsules with a persistent papery endocarp are found.
In the Australian diagram, species numbered 10, 9, and 11 (P. bicolor, P.
bbillyraeoides, and P. o'reillyanum) show a similar proportionate reduction to that
noted for the New Zealand P. umbellatum - P. pimeleoides group. The remaining
Australian species fall into two groups. The first group is characterized by petals
and sepals which are long and broad. Associated characters are few flowers and
lobed, ellipsoid or ovoid capsules. The second group is characterized by petals and
[Vol. 43
120 ANNALS OF THE MISSOURI BOTANICAL GARDEN
sepals which are small and narrow. Associated characters are very numerous
flowers and usually smaller globose or obovoid capsules.
Group 1 Group 2
24. P. revolutum 27. P. venulosum
25. P. rubiginosum 28. P. melanospermum
26. P. undulatum
N
NC
. P. rbombifolium
30. P. ferrugineum
Many years ago, Bentham!*? recognized that P. revolutum, P. rubiginosum and
P. undulatum had characters in common, and that P. rbombifolium, P. melanosper-
mum and P. ferrugineum had some general affinity, but, apart from his key and
arrangement of species, he did not express his views on the relationship of the
species.
The significance of the pictorialized scatter diagram is that it shows a high
degree of correlation among the characters employed, and strongly supports the
key and taxonomic arrangement. Moreover, since the diagram is based on actual
measurements, the results expressed therein are repeatable.
Stupy MATERIALS
Specimens from the herbaria of the following institutions have been studied,
and the symbols !** employed in their citation are indicated,
—Arnold Arboretum of Harvard University, Cambridge, Mass.
AK —Auckland Institute and Museum, Auckland, New Zealand.
BH —Bailey Hortorium, Cornell University, Ithaca, N.Y.
BISH —Bernice P. Bishop Museum, Honol u, Hawaii.
BM ritish Museum (Natural History), London, England
BRI —Depa t of Agriculture, Brisbane, Australia
CANB —Division of Plant Industry, anberra, Australia
CANTY —Canterbu useum, Christchurch, New
CU —Wiegand Herbarium, nell University, Ithaca, N.Y
F hicago Natural History Muse Chicago,
GH —Gray Herbarium of Harvard University, Cambridge, Mass.
HO —University of Tasmania, Hobart, Tasmania.
ILL —Department of Botany, University of Illinois, Urbana, Ill.
—Royal Botanic Gardens, Kew, England.
MASS —Departm f Botany, University of Massachusetts, Amherst, Mass.
MEL — National Herbarium of Vi toria, Melbourne, Australia.
MO —MMissouri Botanical Garden, St. Louis, Mo.
NSW — National Herbari f New South Wales, Sydney, Australia. :
P uséum National d'Histoire Naturelle, Laboratoire de Phanérogamie, Paris.
U —Botanical Museum and Herbarium, State University of Utrecht, Netherlands.
UMEL —Department of Botany, University of Melbourne, Melbourne, Australia.
US — Department of Botany, . National Museum, Washington 25, D. C.
Ww —Naturhistorisches Museum, Wien, Austria.
143 Fl, Austr, 1: 110. 186
863.
144 Lanjouw & Stafleu, Reg. Veg. Index Herbariorum Pt. 1. The herbaria of the world, ed. 2.
1954.
1956]
COOPER—AUSTRALIAN AND NEW ZEALAND PITTOSPORUM 121
ACKNOWLEDGMENTS
I wish to acknowledge my indebtedness to the curators of the herbaria listed
above for the loan of specimens and other courtesies. The Director and staff of
the Auckland Museum, and many Australian and New Zealand friends, have sup-
plied me with specimens and information. I am also indebted to Dr. Egbert
Walker of the U.S. National Museum and to Dr. F. Raymond Fosberg of the U.S.
Geological Survey for sponsoring my visit to the United States. and to the Director
and staff of the Missouri Botanical Garden, and to the Dean and staff of the Henry
Shaw School of Botany, for many kindnesses during my stay, Finally, I must
express my great obligation to Dr. Robert E. Woodson, Jr., of the Missouri
Botanical Garden, who made my visit to the United States possible and for whose
assistance and advice I cannot be sufficiently grateful.
TAXONOMY
PrrrosPoRuM Banks & Soland. ex Gaertn. Fruct. et Sem. 1: 286. £. 59. 1787-88
(T.: P. tenuifolium). Banks & Soland.
Pittosporoides Banks & Soland. ex Gaertn. Fruct. et Sem. 1: 286. 1787-88, nom. nud. in
synon.
Erect trees or shrubs, rarely prostrate or epiphytic, pubescent or glabrous,
unarmed or spinose. Leaves alternate, occasionally subverticillate, usually entire,
petiolate, rarely sessile. Inflorescence terminal, terminal and lateral, or axillary,
few to many-flowered, in fascicles or in umbelliform to paniculiform cymes, or
solitary. Sepals 5, free, coherent or rarely connate. Petals 5, free or connivent,
tips spreading or recurved. Stamens 5, erect; anthers 2-celled, introrse. Ovary
sessile or stipitate, incompletely 2- to 4-celled; stigma capitate to truncate; style
usually short. Capsule globose, ovoid, or obovoid, 1-celled; valves 2 - 5, woody or
coriaceous, with a median longitudinal parietal placenta, Seeds immersed in a resin-
ous viscid fluid.
The synonomy of Pittosporum is given in full by Pritzel!*5.
KEY TO THE SERIES
Capsules 3-valved, rarely 4- or 2-valved; seeds m ee eg
sepals pubescent rats to glabrous in P. Ber cie Norfolk Island, Lord oz
Island, and New Zealan F E iA
Cisia Cat ei ¡rely tkd: seeds few; sepals and petals E small ;
minutely p r glabrous denm in P. rubiginosum, P. Arg re
gatum). rome di Së Nee Dn
145 Engler & Prantl, Nat. Pflanzenfam. ed. 2. 18:273. 1930.
[Vol. 43
122 ANNALS OF THE MISSOURI BOTANICAL GARDEN
Series J. TRIVALVAE
Series I. TRIVALVAE Gowda, in Jour. Arnold Arb. 32:284. 1951.
KEY TO THE SPECIES
a. Petals cream-white or yellow-green; sepals glabrous or scantily pubescent without;
flowers terminal, 1 1-8, fascicled; capsules globose to pyriform; m usualy convex in
e
transverse section with a placenta ud iio from the base to the apex.
and Lord Howe Tile s.
bel
acuminate, 10.0-13.5 mm. long, 2.5-3.0 mm. broad, with a lew hairs e
vein without, glabrous —— petals yellow- -green; capsules globose to pyri
3.5 br
cm. long, 1.5-2.5 cm. broad, without persistent dried sepals. Sch Mai
bb. Shrubs to 5 m. tall; leaves acute, 3—5 cm. long, 1.0—1.8 cm. broad; REN
—2
Norfolk
rees to 7 m. tall; leaves acuminate, 5-11 cm. ime 2.0-3.6 cm. broad; sepals linear,
he ne
teolatum
De
late, acute, 4—8 mm. long, 1.5—2.5 mm. broad, glabrous pace brown-tomentose
vi s E cream-white with a reddish claw; capsules usually globose,
diameter, often with eem ag sepals. Lord How e Hand
1.3-2.0
A P. erioloma
aa. ere purple, reddish-brown, or red; sepals Pubeicmt without "enatis puberulent to
glabrous in P. tenuifolium); Aiken terminal or axillary, 1—10, fascicled or solitary;
capsules subglobose or trigonous, rarely 2- or 4- lobed; valves convex to concave in
transverse section with a pa acenta bearing funicles from thé base to about the midde De
land
the SEN in P. fairchildit). w Zea
terminal ies of 2-10, rarely solitary; capsules relatively large and
di ck.
e Capsules subglobose; valves conv
thick a
ex in t
margins; flowers up to 6 per fcis d pue oae reli brown, or chocolate
Cl bro
d. Shrubs 3-5 m. tall; flowers 2—4; sepals lanceolate, acute, 4-7 ong,
tomentose without vem within}: petals purple; wie eh KE pie?
markedly shickaned apex, bearing minute funicles from the base he apex; —
seeds black, round "Three Kings Islan P. fairchildii
dd. m > oat m. tall; flowers 2-6; sb linear, acumin: 8-11 long
ntose without and to near the base within; petals r silini to
chocolate; capsules E or 2-va dt 2 valves much thickened at apex, be
g flat-
tened funicles up to m. long from the secl to En Lara seeds reddish-bla ck,
irregular. North (ën rte nd, Thames and East Cape .........-----
ce, Capsules trigonous, rarely 2- or 4-lobed; opas Co
IP —
eat
. thick at margins; flowers up i E per fasci cles ëch Séi Ga to el
H Sheubs to small trees 1-10 m. tall; leaves obovate to oblanceolate, 3-8 cm. DNE
1-3 cm. broad, attenuate at Zong de stron ae chicken ed and revolute
b
secondary veins about 5 per side; flowers ee sepals oblong to escher
capsules 1-3 cm.
acute, he edo gos ond withow ut ui regen va d within;
an
long d broad; va y, with a placenta Ee con-
spicuous eeng d Pees ic geass and Thames, on the coast.
5
Kermedec
[59
a
E Gr 2.5 = - m. ima leaves oblong to obovate, 4-12 cm. long, 2-4 cm
s
P. crassifolium
valves about 2 mm. thick, subwoody to cori ind: ac? a nds bearing incon-
i and
spicuous funicle. Wa E nui, Gate: Volcanic Plateau, East Cape
n d in
DE AARP
and in
bb. Flowers in axilary fascicles e? 3, or errs or on very reduced lateral con if
terminal; capsules funds small and thin
Sh
€. Shrubs or trees to 10 m. tall; Vonig leaves, pene and sepals white-floccose-
tomentose; flowers 1—3, usually subtended by o severa cataphylls; pedicels up
o
a Ay cm. s ng; — oblong, acute; petals r e e Paien capsules — 1.2-
ght tion.
n diameter; valves slightly c onvex to flat in transverse
See "island d the Coronada Peninsula.
reat
we? P. buttonianum
1956
COOPER—AUSTRALIAN AND NEW ZEALAND PITTOSPORUM 123
cc. Trees to 10 m. tall;
puberulent to glabrous; flowers 1, rarely 2-3, usually subtended by a leaf; pedice
cm. long; t
up to 1 sepals ovate to oblong, sub-acute to obtuse; petals usually purple,
occasionally maroon, pink, o ite; capsules subglobose, 1.0—1.2 cm. in diameter;
valves convex in transverse section. Throughout New Zealand ................ 8. P. tenuifolium
PrrrosPORUM BRACTEOLATUM Endl. Prodr. Fl. Norf. p. 78. 1833. (T:
Bauer s.n.!).
Trees to 7 m tall; branchlets grayish-brown, the young parts sparsely tomen-
tulose, soon glabrate. Leaves alternate, frequently crowded at the tips of the
branches, lanceolate-oblong to oblanceolate, acuminate, acute to attenuate at base,
entire, 5 - 11 cm. long, 2.0 - 3.6 cm. broad, dark green above, paler beneath, gla-
brous, membranous, margins slightly revolute and undulate, costa immersed above,
raised beneath, secondary veins 8-14 per side, anastomosing, obscure above,
distinct beneath; petioles 4 - 9 mm. long, 1 - 2 mm. broad, sparsely tomentulose
when young. Flowers terminal, 1 - 8, fascicled; pedicels 2 - 3 cm. long, accrescent
and recurved in fruit, sparsely tomentulose when young, usually bearing a linear
bract and subtended by a loose whorl of leaves and caducous, sparsely ciliolate bud
scales about 1 cm. long. Sepals free to the base, linear, acuminate, 10.0 - 15.5 mm.
long, 2.5 - 3.0 mm. broad, with a few hairs on the main vein without; petals linear,
acute, 1.5 - 2.0 cm. long, 4.0 - 4.5 mm. broad, free, spreading from the base,
yellow-green; stamens 7.5 - 10.0 mm. long, anthers linear-oblong or sagittiform,
1.5 - 3.0 mm. long, 0.5 - 0.8 mm. broad. Pistil at anthesis slightly shorter than the
stamens; ovary 4 - 7 mm. long, 1.5 - 5.0 mm. broad, tomentose; style 4.0 - 4.5
mm. long; stigma truncate. Capsules globose to pyriform, 3-valved, 2.0 - 3.5 cm.
long, 1.5 - 2.5 cm. broad, brown-tomentose, glabrate, rugose; valves convex to
slightly sulcate in transverse section, 3 - 5 mm. thick, woody, light yellow within,
with a conspicuous placenta bearing 2 rows of short stout funicles from the base
to the apex; seeds about 70, reddish-black, round to irregular.
Found only on Norfolk Island. Flowers in the winter months (June to
August).
NorroLk Istanp: Bauer s.m. (K); in woods, A. Cunningham 26 (K); Backhouse
669 (K); on the margins of woods, A Cunningham s.n., 138 (K); Mueller s.n. (K);
Maiden & Boorman s.n. (BM, K); Paterson s.n. (BM).
A flower which may have been female had a plump ovary 7 mm. long and 5
mm. broad, stamens 7.5 mm. long, and sagittiform, probably abortive, anthers 1.5
mm. long. The style was missing. Flowers which may have been male had slender
ovaries 3.5 - 4.5 mm. long, 1.5 - 2.0 mm. broad, stamens 8.5 - 10.0 mm. long, and
linear-oblong anthers 2.5 - 3.5 mm. long. The styles were 3.5 - 4.5 mm. long and
the stigma was truncate.
2. PrrrosPORUM ERIOLOMA Moore & F. Muell. in F. Muell. Frag. Phyt. Austr. 7:
139. 1871. (T.: Fullager s.n.!).
Shrubs 2 - 5 m. tall; branchlets grayish-brown, the young parts brown-tomen-
tose, soon glabrate. Leaves approximately verticillate, oblanceolate to obovate,
—
D
[Vol. 43
124 ANNALS OF THE MISSOURI BOTANICAL GARDEN
i
== P. BRACTEOLATUM
l
' p. ERIOLOMA
A P. FAIRCHILDII
3X £4 L /
Yo LAST LONGITUDE 100 WEST LONGITUDE
Fig. 8. P. bracteolatum, P. erioloma, and
P. fairchildii,
acute at the apex and base, entire, or occasionally toothed or lobed when juvenile,
3-5 cm. long, 1.0-1.8 cm. broad, dark green above, paler beneath, glabrous, coriace-
ous, margins ciliate when young, glabrate, thickened and revolute when mature,
costa raised above and beneath, secondary veins 7 - 9 per side, anastomosing, obscure
above, distinct beneath; petioles 2 - 7 mm. long, 1 - 2 mm. broad, brown-tomentose
when young. Flowers terminal, 2 - 7, fascicled; pedicels 1 - 2 cm. long, accrescent
and recurved in fruit, brown-tomentose, subtended by a whorl of leaves and caduc-
ous, ciliate bud scales about 3 mm. long. Sepals imbricate at base, lanceolate, acute,
4-8 mm. long, 1.5 - 2.5 mm. broad, ciliate, glabrous without, brown-tomentose
within; petals lanceolate-oblong, subacute, 15 - 18 mm. long, 3 - 4 mm. broad, free
or coherent at the base, spreading from above the middle, cream-white with a
reddish claw; stamens 7- 11 mm. long, anthers sagittiform or elliptic-oblong,
1.5-3.5 mm. long, 0.5 - 1.00 mm. broad. Pistil at anthesis slightly shorter or
longer than stamens; ovary 3-6 mm. long, 1 - 3 mm. broad, tomentose; style
2.0- 4.5 mm. long; stigma 3-lobed and capitate, or truncate. Capsules globose,
rarely subglobose or 3-lobed, 3-valved, 1.3 - 2.0 cm. in diameter, glabrous, rugose
to slightly rugose, often accompanied by the persistent dried sepals; valves usually
convex in transverse section, 3 - 6 mm. thick, coriaceous, light yellow within, with
a conspicuous placenta bearing 2 rows of short stout funicles from the base to near
the apex; seeds 11 - 23, black, round to irregular.
Mounts Lidgebird and Gower on Lord Howe Island; most common at eleva-
tions above 300 m, in “moss forest". Flowers during September and October.
Loro Howe IstAND: summit of Mt. Lidgebird, Fullager s.n. (MEL); Fitzgerald
$5. (AK); King s.n. (NSW); 600’ to 1800’, Mt. Lidgebird, McComish 136, 1364 (K,
NSW); summit of Mt. Gower, Hedley & Dunn s.m. (NSW); 1000’, Boorman s.
(NSW); Howe’s Island, Moore 9, 23 (K).
Flowers with 3-lobed capitate stigmas which protrude beyond the stamens,
broad filaments, and small sagittiform anthers may be female as the anthers appear
to be abortive. Flowers with truncate stigmas which do not protrude beyond the
1956]
COOPER—AUSTRALIAN AND NEW ZEALAND PITTOSPORUM 125
stamens, narrower filaments, and larger elliptic-oblongoid anthers may be male as
the ovary is small and thin.
Vernacular name: Lord Howe Island's Hedge Laurel.
3. PrrrosPORUM FAIRCHILDI Cheesem. in N.Z. Inst. Trans. & Proc. 20: 147.
1887. (T: Cheeseman s.n.!).
Shrubs 3 - 5 m. tall; branchlets gray to brownish-gray, the young parts white-
tomentose, soon glabrous. Leaves alternate, frequently crowded at the tips of the
branches, obovate, elliptic-obovate or elliptic-oblong, obtuse or acute at apex, acute
at base, entire, 4 - 7 cm. long, 2.0 - 3.5 cm. broad, light green above, paler beneath,
white- to brown-tomentose when young, soon glabrous above but tomentulose
beneath when mature, coriaceous, revolute, costa raised, secondary veins 7 - 11 per
side, anastomosing, distinct; petioles 3 - 8 mm. long, 0.5 - 2.0 mm. broad, white- to
brown-tomentose. Flowers terminal, 2 - 4, fascicled; pedicels 12 - 20 mm. long,
accrescent in fruit, brown-tomentose, subtended by a whorl of leaves and caducous
ciliate bud scales 5 - 10 mm. long. Sepals imbricate at base, lanceolate, acute, 4-7
mm. long, 2.0-3.5 mm. broad, brown-tomentose without, glabrous within, ciliate;
petals lanceolate-oblong, subacute, 9- 14 mm. long, 2.5 - 4.5 mm. broad, free,
spreading from above the middle, purple; stamens 7.0-8.5 mm. long anthers
sagittiform or elliptic-oblong, 1.5 - 2.5 mm. long, 0.5 - 1.3 mm. broad. Pistil at
anthesis slightly longer to slightly shorter than the stamens; ovary 2.5 - 6.0 mm.
long, 2-4 mm. broad, brown-tomentose; style 3 - 4 mm. long; stigma weakly
capitate to truncate. Capsules subglobose, 3-valved, 2.0 - 2.5 cm. in diameter,
tomentose, glabrate, rugose; valves convex in transverse section, 1.5 - 2.0 mm.
thick, coriaceous, with a conspicuous placenta bearing two rows of minute funicles
from the base to the apex; seeds 19 - 22, black, round.
Flowers in August.
EW ZEALAND: THREE KINGS ISLANDs: Great Island, Cheeseman, s.n. (AK, BM,
K); Baylis 22797 (AK); Turbott 23023 (AK); Turbott & Bell 22925 (AK); North East
Island, Buddle 22894 (AK).
Flowers which may be female have sepals up to 6.5 mm. long, petals up to 14
mm. long, sagittiform anthers only about 1.5 mm. long, weakly capitate stigmas
exserted beyond the anthers, thin styles up to 3 mm, long, and stout ovaries 6 mm.
long and 4 mm. broad. Flowers which may be male have sepals 4-7 mm. long,
petals 9 - 13 mm. long, elliptic-oblong anthers about 2.5 mm. long, truncate
stigmas level with or below the anthers, thicker styles 3.5 - 4.0 mm. long, and thin
ovaries about 3.5 mm. long and 2.0 - 2.5 mm. broad.
Cultivated specimens from the Botanic Gardens, Cambridge (England), have
been seen, without collector s.n. (K).
4. Prrrosporum ELLIPTICUM Kirk, in N.Z, Inst. Trans. & Proc. 4: 267. 1872.
(T.: Kirk s.n.!). : RA
Pittosporum ellipticum Kirk ssp. ellipticum Kirk, loc. cit. 1872. (1: Kirk s.n.!).
Pittosporum ellipticum Kirk var. decorum Cheeseman, Man. N.Z. Fl. ed.2. p. 491. 1925.
(T.: Cheeseman s.n.!).
[Vol. 43
126 ANNALS OF THE MISSOURI BOTANICAL GARDEN
Trees to 8 m. tall; branchlets dark brown, covered with appressed rusty tomen-
tum when young, ultimately glabrate. Leaves alternate, crowded at the tips of
the branches, elliptic-oblong to ovate or obovate, acute to obtuse at apex and base,
entire, 3.5 - 9.7 cm. long, 1.5 - 4.8 cm. broad, covered with appressed rusty tomen-
tum when young, soon glabrate above, sparsely tomentose to glabrate beneath,
coriaceous, margins unthickened and slightly revolute, costa raised, secondary veins
7 - 10 per side, anastomosing, obscure above, distinct beneath; petioles 4 - 15 mm.
long, 1.5 - 2.0 mm. broad, rusty-tomentose. Flowers terminal, 2 - 6, fascicled;
pedicels 5 - 14 mm. long, accrescent in fruit, tomentose, subtended by a whorl of
cataphylls and several caducous rusty-tomentose bud scales 1 - 2 mm. long. Sepals
slightly imbricate at base, linear, acuminate, 8- 11 mm. long, 2.0 - 2.7 mm.
broad, rusty-tomentose without and to near the base within, ciliate; petals oblance-
olate-linear, subacute, 14.0 - 19.5 mm. long, 3.0 - 4.5 mm. broad, free, recurved
at the tips, reddish-brown to chocolate; stamens 7.2 - 11.0 mm. long, anthers
sagittiform to oblong, 1.2 - 3.0 mm. long, 0.5 - 1.2 mm. broad. Pistil at anthesis
slightly shorter or longer than the stamens; ovary 2.5 - 7.5 mm, long, 1.5 - 4.5
mm. broad, rusty-tomentose; style 1.5 - 6.5 mm. long, stigma capitate or truncate.
Capsules subglobose, 3- or 2-valved, 1.5 - 2.0 cm. long, 1.3 - 1.7 cm. broad, rusty-
tomentose to glabrate, slightly rugose; valves convex in transverse section, up to
1.5 mm. thick at the margins, much thickened at the apex, almost woody, with a
thin raised placenta fused at the base, bearing flattened funicles up to 1 mm.
long from the base to the middle; seeds 27 - 32, reddish-black, irregular.
Occurs in the hills in the North Auckland, Auckland, South Auckland,
Thames and East Cape districts of the North Island of New Zealand. Flowers
from August to November.
__ NEw ZEALAND: NORTH AUCKLAND: Whangaroa North, Kirk s.n. (AK); Mt. Manaia,
Kirk sn. (A, AK, GH, MO), 122 (K); same locality, Cheeseman s.n. (AK); Whangarei,
Kirk s.n. (A), 623 (BM). AUCKLAND: Waitakere Ranges Cheeseman s.n. (AK, BISH,
K, US); same locality, Meebold 5389 (BISH); same locality, Nihotapu, Matthews,
LeRoy & Carse s.n. (AK); same ocality, eastern side, Bishop s.n. (AK); Titirangi Ranges,
Cheeseman s.n. (GH, K); roadside cliff, Huia, Wood 27210 (AK); Anawhata, West
Coast, Cranwell sn. (K). THAMES DisTRICT: Ohinemuri Gorge near Karangahake,
Cheeseman s.n. (AK) ; same locality, Petrie s.n. (AK); Gordon Settlement, near Te Aroha,
? AK). T CAPE: Hawai River, inland from Opotiki, Field s.n. (AK);
3000”, Aorangi Scenic Reserve near Hikurangi, Williams s.n. (AK).
w
=
Flowers which may be male have truncate stigmas, styles 5.0 - 6.5 mm. long,
ovaries 3.0 - 3.5 mm. long, about 2.5 mm. broad, and long stamens with ‘oblong
anthers up to 3 mm. long. Flowers which may be female have capitate stigmas,
styles about 3 mm. long, ovaries 7.5 mm. long, 4.5 mm. broad, and short stamens -
with sagittiform anthers 1.2 - 1.5 mm. long, and about 0.5 mm. broad. Male
flowers in which the pistil is only 4 mm. long also occur. It is difficult to decide
whether this species belongs to the TRIVALVAE or BIVALVAE; Michie** has found
trees with 3-valved fruits but the specimens I have seen in the Waitakere Ranges
146 Personal communication.
1956] :
COOPER—AUSTRALIAN AND NEW ZEALAND PITTOSPORUM 127
have mostly 2-valved fruits. A yellow-flowered form is in cultivation in New
Zealan
Pittosporum ellipticum Kirk ssp. ovatum Kirk!*" is based on fruiting speci-
mens collected at Whangaroa North over 80 years ago. Kirk!*5 also cited Cheese-
man's specimens from the Manaia Hills and the Titirangi district, but none of these
match the description and the type collection from Whangaroa North. The
Whangaroa district seldom has been visited by botanists and until more material
is available the status of the subspecies is obscure. A specimen labelled var. ovatum
in the Herbarium of the Royal Botanic Gardens, Kew (Kirk 105) resembles a
broad-leaved form of P. virgatum.
Pittosporum ellipticum Kirk var. decorum Cheeseman was based on the col-
lectings of Cheeseman, Petrie, Downard, Field, and Williams, from the Thames and
East Cape districts. Cheeseman wrote:
entum, ing nd
lateral. It has some points of agreement with P. huttonianum but never has the white
floccose tomentum which is such a distinctive character when P. buttonianum is in
oom.
Cockayne identified Petrie's specimens from Karangahake as P. huttonianum,
but a comparison of the characters of P. ellipticum, Cheeseman's var. decorum, P.
huttonianum, and P. ralphii shows that var. decorum has some characters in com-
mon with each of the other three species. Pittosporum buttonianum and P. ellipti-
cum are found to the north, and P. ralphii to the south of Cheeseman's var.
decorum; and this last may possibly be representative of a heterozygous population
derived through introgressive hybridization of the three species. Pittosporum
buttonianum, P. ellipticum, and P. ralpbii occur mainly in areas which were not
submerged in recent geological time. On the other hand, Cheeseman's var. decorum
is found in localities which were beneath the sea during the Castlecliffian (upper
Pliocene) 149, and it is possible that past geographical changes have been significant
in the formation of the three peripheral species and the population represented by
Cheeseman's var. decorum.
5. Prrrosporum CRASSIFOLIUM Banks & Soland. ex A. Cunn. in Ann. Nat. Hist.
4:106. 1839. CT: R. Cunningham s.n.!) .
Pittosporum crassifolium var. strictum Kirk, in N.Z. Inst. Trans. & Proc. 4:266. 1872.
Shrubs to small trees 1 - 10 m. tall; branchlets dark brown, the young parts
White- or rusty-tomentose, becoming black. Leaves alternate, frequently crowded
at the tips of the branchlets, obovate to oblanceolate, obtuse to acute at apex,
attenuate at base, entire, 3 - 8 cm. long, 1 - 3 cm. broad, white- or brown-tomen-
tose when young, soon glabrate above but densely appressed-tomentose beneath,
coriaceous, margins thickened and strongly revolute, costa raised, secondary veins
HINZ. ep? Goen & Proc. 4: 267. 1872. (T.: Kirk s.n.!).
n Vë cit.
[etui in up recie 2: 80. 1949.
[Vol. 43
128 ANNALS OF THE MISSOURI BOTANICAL GARDEN
LONGITUDE
MMMM O
Fig. 9. P. ellipticum. Fig. 10. P. crassifolium
about 5 per side, anastomosing, usually obscure; petioles 4- 14 mm. long 1-3
mm. broad, brown-tomentose. Flowers terminal, 1 - 10, fascicled; pedicels 0.6 -
5.0 cm. long, accrescent in fruit, tomentose, subtended by a whorl of leaves and
numerous caducous, brown-tomentose, ciliate bud scales 3 - 15 mm. long. Sepals
imbricate at base, oblong to linear-lanceolate, acute, 7 - 11 mm. long, 1.5 - 3.0
mm. broad, brown-tomentose without, and to above the middle within, ciliate;
petals oblanceolate to lanceolate, subacute, 10 - 16 mm. long, 3 - 5 mm. broad, free,
recurved at the tips, dark red to purple, becoming black on drying; stamens 5 - 9
mm. long, anthers sagittiform or elliptic-oblong, 1 - 3 mm, long, 0.5 - 1.5 mm.
broad. Pistil at anthesis slightly shorter or longer than the stamens; ovary 3 - 6
mm. long, 2 - 5 mm. broad, tomentose; style 3.0 - 4.5 mm. long; stigma capitate
and obscurely 3-lobed or truncate. Capsules trigonous, rarely 2- or 4-lobed, 3-
valved, rarely 2- or 4-valved, 1 - 3 cm. long and broad, white- to brown-tomentose,
rugose; valves concave in transverse section, 2 - 3 mm. thick, woody, with a
conspicuous placenta bearing short stout funicles from the base to near the apex;
seeds 20 - 36, black, irregular to round.
Common on the coast of Sunday Island in the Kermadec Islands group and on
the coast of the northern portion of the North Island of New Zealand. Flowers in
September and October. ;
KERMADEC ÍsLANDs: SUNDAY ISLAND: northern shore, Cheeseman s.n. (AK, K).
N EALAND: NORTH AUCKLAND: Waya—Rum Bay, Wilkes s.n. (K); on Flat Island,
t - AUCKLAND: Great Omaha, Kirk s.n, (A,
GH, MO), 25 (BM); Kawau Island, Kirk s.n. (AK), 116 (K); David Island, Molesworth
23320 (AK); coast near Auckland, Cheeseman s.n. (AK, BISH, K, NEW); same locality,
Petrie 6417 (A, NSW); Lake Pupuke, Cheeseman s.n. (AK, GH, NSW, US); Auckland,
bapman s.n. (A). THAMES: Tryphena Harbour, Great Barrier Island, Osborne s.m.
1956
COOPER— AUSTRALIAN AND NEW ZEALAND PITTOSPORUM 129
(AK) Great Barrier Island, Matthews 1822 (AK); Little Ee Island, Kirk s.n. (AK),
123 (K); Cape Colville, Kirk sm. (US); Kenn SR Bay mandel, Matthews s.n. (AK,
"E WITHOUT DEFINITE LOCALITY: Allison s SC sg WA Bid vill 198 (K); cone
- (K); Cunningham 43 (U); C. [Cunningham] 2 (NSW); Mueller s.n. (CU, G
Wilkes s.n. (GH); Sinclair s.n. (BM, MO).
Flowers which appear to be female have capitate, obscurely 3-lobed stigmas,
styles about 3 mm, long, and plump ovaries about 6 mm. long and 5 mm. broad.
The anthers are only about 1 mm. long, sagittiform, and the filaments are about
4 mm. long, markedly broad at the base. In the material which I have examined
the flowers are solitary. On Molesworth s.n. two or three fruits occur together,
and Cheeseman”? describes female inflorescences as 1- to 5-flowered. Pedicels
are markedly shorter at anthesis but elongate in fruit. Flowers which appear to be
male have truncate stigmas, styles about 3.5 - 4.5 mm. long, slender ovaries about
3 mm. long and 2 mm. broad. The anthers are 2- 3 mm. long, elliptic-oblong,
and the filaments are 5 - 6 mm. long. In the material which I have examined the
inflorescences are 2- to 8-flowered but Cheeseman’! describes them as 5- to 10-
flowered. The pedicels of "male" flowers are markedly longer than those of
“female” flowers.
Banks and Solander collected specimens of a species of Pittosporum on the east
coast of New Zealand in 1769 and named it P. crassifolium in herb. but did not
publish its description and figure. Allan Cunningham applied the name to speci-
mens collected by his brother Richard, in 1833, "on Flat Island, one of the Cavallos
Group, and at Matauri, etc.” In the Herbarium of the Royal Botanic Gardens,
Kew, the specimens are labelled: —‘On Flat Island, one of the Cavallos, Hamilton
s.n. [Dr. Hamilton of H.M.S. Buffalo, Dec. 1833]; Bay of Islands, R. Cunningham
s.n. [612]; at Matauri, opposite the Cavallos, R. Cunningham s.n. [612]". Unfor-
tunately, these collectings were of a different species from that discovered by Banks
and Solander, Plants similar to those collected by Banks and Solander were sub-
sequently described as P. ralphii by Thomas Kirk (N.Z. Inst. Trans. & Proc. 3:
161. 1871).
Kirk!?? described P. crassifolium var. strictum from Little Barrier Island as a
variety distinguished by having several erect fruiting pedicels clustered together,
and smaller capsules. Solitary and clustered fruits occur on the same plants,
however, and similar small-fruited plants, which grade into the larger-fruited form,
have been found in a number of localities, e.g.: on Hen and Chicken Islands
(Moore $ Cranwell s.n.!, AK) ; Poor Knights Islands (Moore & Cranwell s.n.!, AK) ;
Great Barrier Island (Matthews s.n.!, AK); and Crayfish Island, 3 miles south of
Helena Bay (Olsen zo!, AK). Kirk 123 (K), from Little Barrier Island, is also
remarkable in having pedicels up to 5 cm. long but only one collection is available.
Specimens collected on the Kermadec Islands (Cheeseman s.n.) also have markedly
150 Man. N.Z. Fl. ed. 2. p.493. 1925.
151 loc. cit. 1925.
152 N.Z. Inst. Trans. & Proc. 4: 266. 1872. (T.: Kirk s.n.!).
[Vol. 43
130 ANNALS OF THE MISSOURI BOTANICAL GARDEN
narrow leaves, strict erect fruiting pedicels clustered together, and smaller capsules
as in var. strictum. All these collections may be distinct but in the absence of
flowers of the Kermadec plants and an adequate sample of the Little Barrier popula-
tion they had best be left with P. crassifolium.
Two horticultural varieties are available to gardeners, but are not different
in any important characters. A plant with yellow flowers was found at Tryphena
Harbour, Great Barrier Island, some years ago (Osborne s.n.!) and is known in the
trade as var. album. A form with variegated leaves is available as var. variegatum
(Duncan & Davies Cat, Choice Nursery Stock, 47. 1947), but is not as vigorous
as plants with green leaves. The latter are extensively used in New Zealand, the
south of England, Spain, and California, as ornamentals and hedge plants.
Vernacular name: Karo.
6. PrrrosPoRUM RAaLPHi Kirk, in N.Z. Inst. Trans. & Proc. 3:161. 1871, ex
char. (T.: Ralph s.n.).
Pittosporum crassifolium Banks & Soland. ex Kirk, Students’ Flora N.Z. p. 51. 1899, nom.
j non
Shrubs 2.5 - 4.0 m. tall; branchlets grayish-brown, white- to buff-tomentose,
glabrate. Leaves alternate, frequently crowded at the tips of the branchlets, oblong
to obovate, occasionally oblanceolate or elliptic, obtuse or acute at apex, acute or
obtuse at base, entire, 4-12 cm. long, 2- 4 cm. broad white- to buff-tomentose
when young, soon glabrate above but densely appressed-tomentose beneath, coriace-
ous, margins unthickened, flat, or revolute, costa raised or immersed above, raised
beneath, secondary veins about 9 - 12 per side, anastomosing, distinct in juvenile
leaves, obscure in adult leaves; petioles 8 - 22 mm. long, 1 - 3 mm. broad, white- to
brown-tomentose. Flowers terminal on long leading or short axillary shoots, 2 -
10, fascicled; pedicels 6 - 14 mm. long, accrescent in fruit, tomentose, subtended
by a single leaf or several cataphylls and tomentose bud scales 3 - 18 mm. long,
apparently pedunculate on the shedding of leaves, cataphylls, and bud scales. Sepals
imbricate at base, linear, acuminate, 7 - 11 mm. long, 1.5 - 3.0 mm. broad, white-
tomentose without, tomentulose within, ciliate; petals oblanceolate-linear to
oblong-linear, obtuse to subacute, 11-18 mm. long, 2-4 mm. broad, free,
recurved at the tips, dark red to purple; stamens 4.5 - 9.0 mm. long, anthers sagitti-
form to oblong, 1.0-3.5 mm. long, 0.5- 1.0 mm. broad. Pistil at anthesis
slightly shorter or longer than the stamens; ovary 2.5 - 4.0 mm. long, 1 - 2 mm.
broad, white-tomentose; style 2 - 5 mm. long; stigma capitate. Capsules trigonous,
rarely 2-lobed, 3-, rarely 2-valved, apiculate, 11 - 17 mm. long, 9 - 13 mm. broad,
white-tomentose, glabrate, rugose; valves concave in transverse section, about 2
mm. thick, coriaceous to slightly woody, with a conspicuous placenta bearing
minute funicles from the much-thickened base to above the middle; seeds 9 - 17,
black, irregular to round.
Found from Patea and Wanganui on the west coast to East Cape and Cape
Kidnappers on the east coast of the North Island of New Zealand. Flowers from
October to December.
1956]
COOPER— AUSTRALIAN AND NEW ZEALAND PITTOSPORUM 131
/
ARA
Fig. 11. P. ralphii and P. huttonianum.
no EAST LONGITUDE
New ZEALAND: East cAPE: Hendley Station, Hodgson s.n. (AK); bridge beyond
Okawa, Meebold 18279 (BISH); prope Tolaga, Opuragi and Totaranui, Banks & Solander
sm. ( ; Tolaga Bay, East Cape, Banks & Solander s.n. (AK, MO, US); coast near
Gisborne, T'ownson s.n. AK, BM); Patangata, Hamilton s.n. (AK); Havelock North,
Meebold 4823, 5564 (BISH); Pourere, Kirk s.n. (MO); Pitane, Kirk s.n. (A, GH); 700’,
Te Reinga Falls, near Waikaremoana, Sainsbury s.n. (AK); Wairoa, Sainsbury 164, 622,
623, 624, 708, 709, 710, 711 (AK, MO); Waikaremoana, Cranwell & Moore s.n. (AK);
same locality, near outlet, in forest, Oliver s.n. (AK); road bank near Te Araroa, Moore
Ei Cranwell s.n. (AK); Mokopeka, Meebold 18280 (BISH). wANGANUI: near Taumarunui,
Kirk s.n. (AK); Upper Wanganui River, Allison s.n. (A, NSW); Patea, ex hort., Cheese-
man s.n. (GH); same locality, ex hort., Hector s.n. 176 (K); same locality, ex bort., Kirk
656 (K). wirHour Locauity: Colenso 206, 344 (K); Kirk s.n. (P).
Flowers which may be female have capitate stigmas, styles about 2 mm. long,
and plump ovaries about 4 mm. long and 2 mm. broad. The anthers are probably
sterile, sagittiform, 1.0 - 1.5 mm. long, and the filaments are 3.0 - 3.5 mm. long,
markedly broad at the base, tapering distally. Flowers which appear to be male
have weakly capitate stigmas, styles 3.5 - 5.0 mm. long, and slender ovaries 2.5 -
3.5 mm. long, and 1 - 2 mm. broad. The anthers are oblong, 3.0 - 3.5 mm. long,
and the filaments are 4- 6 mm. long, not markedly broad at the base. On one
specimen (Cheeseman s.n. ex hort), several flowers were found with 4 func-
tional stamens about 9 mm. long and one apparently sterile stamen about 6 mm.
ong. The functional stamens had oblong anthers about 3 mm. long while the
apparently sterile stamens had sagittiform anthers only 1 mm, long. The stigmas
were weakly capitate, the styles long, and the ovaries slender as in male flowers.
Godley153 has informed me that the species is dioecious.
It has long been known that Banks and Solander collected specimens of P.
ralphii at Tolaga Bay on the east coast of the North Island of New Zealand during
Cook’s first voyage to the South Seas in 1769-1772, and named it P. crassifolium in
herb, but their description and plate were never published and the name P. crassi-
folium was later applied by A. Cunningham to specimens of another species col-
lected by his brother, R. Cunningham, in North Auckland. Banks & Solander
Lon, BM) consists, however, of two sheets of P. ralphii Kirk and one sheet of P.
153 2
Personal communication.
[Vol. 43
132 ANNALS OF THE MISSOURI BOTANICAL GARDEN
crassifolium A. Cunn (sterile). The type of P. ralphii is a collection made by Dr.
Ralph at Wanganui about 1870 and forwarded to Kirk for description.
A form with variegated leaves is available in the trade in New Zealand. The
species is cultivated in England (Isles of Scilly, Penzance, Allerford, etc.).
7. PrrrosPORUM HUTTONIANUM Kirk, in N.Z. Inst. Trans. & Proc. 2: 92. 1870.
(T.: Kirk s.n.!).
Pittosporum buttonianum Kirk var. fasciatum Kirk, Students’ Fl. N.Z. p. 48. 1899. (T:
Shrubs or trees to 10 m. tall; branchlets dark brown, covered with floccose
white tomentum when young, glabrate. Leaves alternate, frequently crowded at
the tips of the branchlets, elliptic-oblong to obovate-oblong, acute to obtuse at
apex and base, entire, 4.2 - 12.0 cm. long, 2.1- 5.0 cm. broad, covered with
floccose white tomentum when young, glabrate, slightly coriaceous, margins
unthickened, flat or undulate, occasionally revolute, costa level or sunken above,
raised beneath, secondary veins about 12 per side, anastomosing, obscure above,
distinct beneath; petioles 5 - 15 mm. long, 1.0 - 2.5 mm. broad, covered with ap-
pressed white tomentum when young, glabrate. Flowers terminal and axillary,
solitary or in fascicles of 3; pedicels up to 2 cm. long, accrescent in fruit, white-
tomentose, usually subtended by 1 to several cataphylls and caducous tomentulose
or glabrous bud scales 2 - 10 mm. long. Sepals slightly imbricate at base, oblong,
acute, 6 - 9 mm. long, 2.2 - 3.0 mm. broad, covered with floccose white tomentum
without, glabrous within; petals oblanceolate-linear, obtuse to subacute, 12 - 18
mm. long, 3.0 - 4.2 mm. broad, free, recurved above the middle, red to magenta;
stamens 6-10 mm. long, anthers sagittiform to oblong, 1.5 - 4.5 mm. long,
0.8 - 1.5 mm. broad. Pistil at anthesis slightly shorter or longer than the stamens;
ovary 2.6 - 5.0 mm. long, 1.0 - 3.5 mm. broad, white-tomentose; style 2.0 - 5.5
mm. long; stigma weakly capitate or truncate. Capsules trigonous, 3-valved,
rarely 2-valved, apiculate, 1.2 - 2.0 cm. in diameter, white-tomentose, glabrate,
weakly rugose; valves slightly convex to flat in transverse section, about 1 mm.
thick, coriaceous, with a thin placenta, fused at the base, bearing flattened funicles
up to 1 mm, long from the base to above the middle; seeds 18 - 23, reddish-black
to black, irregular.
Occurs on the Great Barrier Island and the Coromandel Peninsula in the
Bae Island of New Zealand. Flowers from October to November.
W ZEALAND: THA: Great Barrier Mond: Kirk s.n. (A, K, US), 86 (K), 518
(BM); Karan same bakt Kirk s.n. (AK, GH, MO); Kaiwiriki, same locality, K Kirk
sm. (AK); timber track, Whangaparapara, same locality, Molesworth s.n. (AK); same
i E
Bay, Coromandel, Matthews s.n. (AK, MO); 1600', Castle Rock, Coromandel, Cheeseman
GH); same locality, Petrie 6418 (A); same locality, Cranwell & Moore sM-
(AK); Table Mountain behind Thames, Adams s.n. (AK); Pakirerehi, Aion sn. (AK);
same locality, Cheeseman s.n. (US); Steeg, Hector 176 (K).
Pittosporum huttonianum var. fasciatum was characterized by flowers in
terminal cymes and peduncles white with floccose tomentum. The “peduncles”
are the young shoots which have not developed a subtending whorl of leaves or
Di
Py
=
~
-
1956]
COOPER——AUSTRALIAN AND NEW ZEALAND PITTOSPORUM 133
cataphylls. This phenomenon occurs in the development of the inflorescence of
other species and does not merit varietal status.
Flowers which may be female have weakly capitate stigmas on styles about
2 mm. long, plump ovaries 4.8 - 5.0 mm. long, 3.5 mm. broad, and short stamens
6 mm. long, with sagittiform anthers 1.5 mm. long, 0.8 mm. broad. Flowers
which may be male have truncate stigmas on styles 3.5 - 5.5 mm. long, slender
ovaries 2.6 - 4.0 mm. long, 1 - 2 mm. broad, and stamens 6.8 - 10 mm. long with
oblong anthers 2.8 - 4.5 mm. long, 1.0 - 1.5 mm. broad.
8. PrrrosPORUM TENUIFOLIUM Banks & Soland. ex Gaertn. Fruct. et Sem. 1:
286. t. 50. 1787-88. (T.: Banks & Solander s.n.!).
Trees to 10 m. tall; branchlets grayish-brown, tomentose when young, gla-
brate. Leaves alternate, frequently crowded at the tips of the branchlets, oblong,
obovate or elliptical, acute to obtuse at apex and base, entire, 1.5 - 12.2 cm. long,
1.2 - 4.7 cm. broad, green above, sometimes marked with purple, paler beneath,
sparsely tomentose when young, soon glabrate, slightly to strongly coriaceous, mar-
gins unthickened, flat, revolute, or undulate, costa sunken above, raised beneath,
secondary veins 6 - 12 per side, anastomosing, obscure above, distinct or obscure
beneath; petioles 3 - 12 mm. long, 0.5 - 2.5 mm. broad, glabrous. Flowers axillary,
1 - 3, solitary or fascicled, subtended by a whorl of caducous bud scales, occasion-
ally terminal on short axillary shoots and subtended by 1 - 2 cataphylls or leaves,
and several caducous linear bracts, sessile or on pedicels up to 1 cm. long, tomen-
tose, accrescent in fruit. Sepals slightly imbricate at base, ovate to oblong, sub-
acute to obtuse, 3-7 mm. long, 1.4- 4.5 mm. broad; petals oblanceolate-linear
to spathulate and clawed, obtuse, 8 - 16 mm. long, 2 - 5 mm. broad, free, reflexed
from the middle, dark purple, maroon, pink or white; stamens 3.6 - 8.0 mm, long,
anthers sagittiform to oblong, 1 - 4 mm. long, 0.5 - 1.0 mm. broad. Pistil at
anthesis slightly shorter or longer than the stamens; ovary 2.5 - 4.5 mm. long,
1 - 3 mm. broad, tomentose; style 2.0 - 3.5 mm. long, stigma capitate or truncate.
Capsules subglobose, apiculate, 3-valved, rarely 2- or 4-valved, 9-14 mm, in
ter, tomentose, glabrate, weakly rugose; valves 1.0 - 1.5 mm. thick, coriace-
kd Convex in transverse section, with a placenta raised and fused at the base,
g peg-like or flattened funicles from the base to above the middle; seeds
6-31, ‘black, irregular.
KEY TO THE SUBSPECIES
L cm. long, 1-3
cm. broad, membranous to Spe coriaceous, frequently marked with Race: whee young,
Margins often undulate als us , 2.5-5.5 long, 1.5-3. o mm. b AT
tomentose to glabrate. Throughout New Zealand, at lower Ce erm a. P. £. fenuifolium
1.1—4.7 cm. broad, gege: not ma purp
acute, 4, 5-6.0 mm, long, tomentulose to slabrace: ig
Waikato and Volcanic m hand, at higher elevations. ............
[Vol. 43
134 ANNALS OF THE MISSOURI BOTANICAL GARDEN
8a. PrrrOSPORUM TENUIFOLIUM ssp. tenuifolium
Trichilia monophylla A. Rich. in Less. & A. Rich. Voy. l'Astrolabe, Bot. Ess. Fl. N. Zel. p.
306. pl. 34. 1832, ex char. & icon. (T.: d'Urville £d).
Pittosporum translucens Hook. ex Putterl. Syn. Pittosp. p. 13. 1839, nom. nud. in synon.
Pittosporum mayi Hort. Regel, Cat. Pl. Hort. Aksakow, p. 112. 1860, nom. nud.
Pittosporum enderi Regel, in Gartenfl. 17: 226. £. 587. 1868, ex char. & icon.
Pittosporum bidwillianum Regel, loc. cit. 1868, nom. nud.
Pittosporum tenuifolium Gaertn. var. tenuifolium Kirk, Students’ Fl. N.Z. p. 47. 1899.
Pittosporum nigricans Hort. Davy in Bailey, Cycl. Amer. Hort. 3: 1360. 1901, nom. nud.
in synon.
Occurs from the North Auckland district in the North Island to the Otago
District in the South Island of New Zealand, at lower elevations than ssp. colensoi.
Flowers from September to November. >
New ZEALAND: NORTH AUCKLAND: Kaitaia, R. H. Matthews s.n. (AK) ; same locality.
H. B. Matthews s.n. (AK); same locality, Kelly's Bush, R. H. Mattbews s.n. (AK, MO);
Kaiaka, Trig Track, Carse s.n. » MO); near sea level on south bank of Oruru River,
aipa, in remnant of Agathis forest, Cooper 36006, 36014 (AK, MASS, MO); Bay of
Islands, A. Cunningham s.n. (BM); same locality, Wilkes s.n. (K, US); Wangaroa, R.
Cunningham 43 (K); Maungatapere, near Whangarei, Carse s.n. (AK). AUCKLAND:
Omaha, Kirk 83 (GH), 85 (BM); Glenfield, Birkdale, H. B. Matthews s.n. (AK, MO);
Auckland, Hector s.n. (GH); same locality, Kirk 85 (US); same locality, Kirk 54 (AK);
Waitakere, Cheeseman s.n. (US); Titirangi Ranges, Cheeseman s.n. (AK); 1300’, same
locality, Meebold 5410 (BISH, K); Huia, Wood s.n. (AK). THAMES: Coromandel
Pu d :
Buckland, H. B. Matthews s.n. (AK). East cape: Te Whaiti, Urewera, H. B. Matt
s.n. (AK); Lake Waikaremoana, Moore & Cranwell s.n. (AK); Wairoa South, Cheeseman
sn. (AK); 2000’, Blowhard, Meebold 18277 (BISH); 1500’, Tongoio, Meebold 18282
(BISH); Havelock North, Meebold 18283 (BISH). vorcawic PLATEAU: Murimohu,
Ohakune, Attwood s.n. (AK). wamaRAPA: 70 Mile Bush, Colenso s.n. (K). WELLING-
TON: Palmerston North, Allan s.n. (BH); Lower Hutt, Kirk s.n. (NSW); in dry woods,
sheltered valley, Day's Bay, 150', MacDaniels s.n. (CU). MARLBOROUGH: Kenepuru,
Pelorus Sounds, MacMahon 205 (AK). NELSON: Maitai Valley, Mellor 27212 (AK);
500 , same locality, near stream on damp graywacke soil, Neal 10 (BISH). CANTERBURY:
Banks Peninsula, Kirk 54 AK); same locality, Haast 69 (K); same locality, Lyttelton
Hills, Meebold 4739 (BISH) ; same locality, Akaroa, Belligny s.n. (GH, K); Peel Forest,
WE 4096 (BISH); 800', Waiau River, Morrison 33 (A); Ashley Gorge, Meebold
);
. T LOCALITY: Banks & Solander s.n. (AK, BM labelled “prope
Tolaga, Opuragi, Ouhuragi, Totaranui, MO, US”); Colenso s.n. (AK), s.n., 714 (K);
A. Cunningham 11 (CU), s:n., 615 (K); Kirk s.n. (A, MO, NSW, US); Sinclair s.n. (BM,
MO); Raoul s.n. (A).
Flowers which may be female have capitate, obscurely lobed stigmas, styles
about 2 mm. long, ovaries about 3 mm. long, 2 mm. broad, stamens 3.6 - 4.5 mm.
long, slightly shorter than the pistil, and sagittiform, possibly abortive anthers
about 1.4 mm. long. Flowers which may be male have truncate stigmas, styles
2.5-3.0 mm. long, ovaries 2.5-4.0 mm. long, 1.0-1.5 mm. broad stamens
6.0 - 7.5 mm. long, slightly exceeding the pistil, and oblong, probably functional
anthers 2 - 4 mm, long. Godley!5* has informed me that the species is dioecious.
Vernacular names: Mapauriki (Sinclair s.n.), Kobubu, Tawbiwbi.
154 Personal communication.
1956]
COOPER—AUSTRALIAN AND NEW ZEALAND PITTOSPORUM 135
/ ] cJ 7
"o LAST LONGITUDE 180 WEST LONGITUDE oJ
Fig. 12. P. tenuifolium ssp. tenuifolium. Fig. 13. P. £. ssp. colensoi.
8b. PrrrosPORUM TENUIFOLIUM ssp. COLENSOI (Hook.f.) Kirk, in N.Z. Inst.
Trans. 8 Proc. 4: 262. 1872.
Pittosporum dide Hook. f. Fl. N.Z. 1: 22. 1853. (T.: Colenso s.n.!).
j . in synon.
o
T m tenuifolium Cie, ssp. fasciculatum (Hook.f.) Kirk, in N. Z. Inst. Trans. &
Oc. 4: 262. 1872.
P SE tenuifolium Gaertn. var. colensoi (Hook.f.) Kirk, Students’ Fl. N.Z. p.47.
Piscine tenuifolium Gaertn. var. fasciculatum (Hook.f.) Kirk, loc. cit. 1899.
P a tenuifolium Gaertn. var. fasciculatum (Hook.f.) Kirk, sub. var. cymosum
9. j Weit: n.!
Firiaona. buttonianum Kirk var. ër EN Kirk, loc. cit. 1899. (T.: Kirk s.n. d
Pittosporum colensoi Hook. f. var. fasciculatum (Hook.f.) Cheesem. Man. N.Z. Fl.
1906.
Occurs from the Waikato and Volcanic Plateau in the North Island to the
Otago district in the South Island of New Zealand, usually at higher elevations
than ssp. tenuifolium. Flowers in November.
New ZEALAND: WAIKATO: Hora Hora, Petrie 764/4 (K). TARANKI: Urenui,
T eds sn. (AK); roadside near Tarata, Carse 764/6 (K). VOLCANIC PLATEAU:
Rotorua, Petrie & H. B. Matthews s.n. (AK); same locality, Turner 159 (AK); ps
locality, Kaliiruros Creek, iie 54 (AK), 2: 64 (K); Genee Plateau, Cheeseman
(AK); near Taupo, Colen (AK), 2361 (K as P. viridum, type of P. er
tum); in jungle filled sa dominated zë el fer near the Raurimu spiral, Moss
Park, Walker 4 tes (MO, US); 3500', base of Ngauruhoe, Cheeseman s.n. (A K); n
Tongariro, Kirk s (MO); same focality; Hodgkins s.n. (AK); 4000' , Ya m. west “Of
ateau : Attwood s.n. (AK); near Haunted Whare, Waimarino, B.
Matthews s.n (AK); MS oes Plains H. B. Mato 764/ 11 (K); same locality,
Zotov s.n. (BH); Maki Manganui-a-te-ao, Waimarin sn. (AK);
1200”, Ohakune, malas iren (BISH); 2000’, awe bie Meebold 18276 ind
NELSON: between Nelson and Havelock, Sledge 2 6 (K). MARLBOROUGH: Awatere
River, EM A Perrin s.n. (AK); same locality, Gas? s.t. (AK); same locality, 2000.
in bush, Travers s.n. (GH). CANTERBURY: Arthurs Pass, Halpin C:
816 (AK);
anaka District, Hector s.n. (GH); Paradise, near Lake Wakatipu, Turner
Wakatipu, Meebold 4442 (BISH); Soen Valley, Lake Te Anau, 800', Petrie 140 (K);
[Vol. 43
136 ANNALS OF THE MISSOURI BOTANICAL GARDEN
Chalky Bay, Lyall s.n. (K); Otago, not common N. of Dusty Bay, Hector & Buchanan 22
. WITHOUT LOCALITY: Bell s.n. (NSW); Colenso s.n., (P. uniflorum, AK), s.n.
(P. viride, K), 2367 (P. uniflorum, K); Dieffenbacb s.n. (K).
Flowers which may be female have capitate stigmas, styles 1.6 - 2.0 mm. long,
ovaries about 4 mm. long, 2.5 - 3.0 mm. broad, stamens about 4.5 mm. long,
slightly shorter than the pistil, and sagittiform, possibly abortive anthers 1.0 - 1.2
mm. long. Flowers which may be male have truncate stigmas, styles 3.0 - 3.5 mm.
long, ovaries 3.0 - 4.5 mm. long, 1.5 - 2.0 mm. broad, stamens 6 - 8 mm. long,
slightly exceeding the pistil, and oblong, probably functional anthers 2 - 3 mm.
ng.
Vernacular name: Mountain Kobubu.
The two subspecies of P. tenuifolium are used in New Zealand, the United
States (California), France, southern England, and the Scilly Islands, the Channel
Islands (Guernsey), Italy, and Australia, for hedges and ornamental trees. In New
Zealand the two subspecies have been so mixed in plantings and so extensively
planted that it is difficult to determine their original distribution. Furthermore,
they intergrade and it is sometimes impossible to determine the subspecies to which
a specimen belongs. Transplantings of trees selected from wild populations at
different altitudes are required to determine the extent to which variation in the
species is caused by the habitat.
Pittosporum fasciculatum Hook. f. was established to include forms which
have flowers in terminal and axillary fascicles. Both P. £. tenuifolium and P. t.
colensoi occasionally have fascicled flowers, and fascicled and solitary fowers are
frequently found together in the other species of Pittosporum with simple inflo-
rescences. Consequently I do not think that the plants with fascicled flowers
should be treated as a distinct species.
Pittosporum buchanani Hook. f. was based on specimens from a plant culti-
vated at Wellington by Buchanan and reputed to come from near Tongariro. The
type specimen at Kew has only "New Zealand” as the locality. Specimens identi-
fied as P. buchanani were subsequently collected, possibly at Kaitaia, by Buchanan,
on Mt. Egmont, by Hector, and by Kirk in the Wellington district. The specimen
at Kew, collected by Buchanan, as noted ‘... the exact place where found is un-
certain but probably between Hokianga and Bay of Islands.” In the last 50 years
the Kaitaia, Mt. Egmont, and Wellington districts have been thoroughly explored
but no further specimens have been obtained. I have treated the name P. buchan-
ani as doubtful because the specimens are too few and imperfect to determine the
status of it.
Pittosporum intermedium Kirk was based on specimens from a single tree on
Kawau Island. The tree was subsequently destroyed and no others have been found.
In describing the species Kirk!?5 wrote:
155 N.Z, Inst. Trans. & Proc. 4: 266. 1872.
1956]
COOPER—AUSTRALIAN AND NEW ZEALAND PITTOSPORUM 137
I give this well-marked form specific rank with some hesitation; in foliage it M^
large forms of P. tenuifolium, while the capsule partakes of the characters of P. cras
folium and P. ellipticum. Dr. Hooker and Mr. Colenso consider it a new species, stil
it is possible that further observation may show the wisdom of uniting it with one
or other of the above. I have been tempted to attribute its peculiarities to hybridisa-
tion.
A sheet at British Museum is noted by H. H. Allen: “very probably P. ellipti-
cum X tenuifolium”. The few specimens available to me match P. umbellatum
in the size and shape of the sepals and petals, but lack the typical large terminal
inflorescence and lobed capsules. I have treated the name P. intermedium as
doubtful under the circumstances.
The following cultivated forms of P. tenuifolium are available in New
Zealand:
P. tenuifolium “garnettii”. Duncan & Davies, Catalogue of trees, shrubs
and climbers, p. 62. 1950-51.—Leaves edged with white, flecked or tinged with
red,
P. tenuifolium "purpureum". loc. cit. Suppl. list 1950-51.—Leaves deeply
tinged with purple.
P. tenuifolium “variegatum”. loc. cit. p. 62. 1950-51.—Leaves edged with
white or cream.
P. tenuifolium "argenteum". loc. cit. p. 32. 1953-54.—This form has been of-
fered for sale as P. tenuifolium “Silver Matipo”. loc. cit. p. 62. 1950-51, and p. 81.
1954-55, but no description is given It may be the same as P, nigricans var. "Silver
Queen” which is described (R.H.S. Dictionary of Gardening, 3: 1595. 1951), as
having silvery-gray foliage, but I have not seen specimens of "Silver Queen”.
Pittosporum tenuifolium "argenteum" and “variegatum” were introduced into
cultivation prior to 1950 but the references are to the earliest dated catalogues
Which I have found.
Davy" lists P. nigra Hort? as a horticultural name of P. tenuifolium and
Hector noted on a sheet in the herbarium of the Royal Botanic Gardens, Kew
(Hector s.n.) that P. tenuifolium was named P. nigrum by the Sydney Botanic
Gardens in 1862. Lord!5? gives P. nigrescens as another horticultural name for P.
tenuifolium. A form with markedly undulate leaves is available in Europe as
P. mayi, and its origin and status are described in detail by Gadeceau'”*,
5 Davy in Bailey, Cycl. Amer. Hort. 3: 1360. 190
157 Lord, Shrubs & Trees for Australian Gardens, p. om 1948.
158 Bull. Soc, Bot. Fr. 67: 153.
[Vol. 43
138 ANNALS OF THE MISSOURI BOTANICAL GARDEN
Serres II. BIvALVAE
Series II. BrvaLvAE Gowda, in Jour. Arnold Arb. 32:285. 1951
KEY TO THE SPECIES
Inflorescences simple; flowers solitary or in fascicles of few to many (rarely in 3- to
oF flowered cymes in P. peblovercidet):
purple, red or yellow;
nd.
r small trees 6-16 tall; branches ascending; leaves oe
— ~ aga marked rich red to purplish-red; capsule valves cori-
k South Wales, Victoria, and Tasma
b. inne trees or S owers terminal and axillary; pee
capsules to shy eect in P. obcordatum). ralia and New Zeala
c. Plants vichout aee diverse juvenile foliage. Aus
d. Unarmed and shrubs; flowers subterminal LT pe solitary or in
fascicles o (x.
Shrubs or small trees 3—10 m. tall; branches pendulous; leaves glabrous; SCH
goo or white; capsule valves woody, 1.0—3.5 thick. Throu a t
tinental Australia P. Viet
ee. Shrubs o
aceous, less mm. thick. New
0 "P. ` bicolor
dd. Lecce shrubs; flowers axillary or poe on arrested es solitary.
disc at high altitu > o'reillyemum
ec NOS ud e t diverse juvenile Gees (juvenile and adult vues linear in
E vw Zeala nd.
d. Flowers Jae, uud. on normal branchlets, 1-20; capsules 6-16 mm. long;
seeds u black, irregular.
e. Sekten never air divaricate or matted.
f. Trees to 8 m. tall or more; ras: emit acute; capsule valves concave or
sulcate to convex in transverse s
g. Trees to 10 m. tall; 0; sepals, pedicels, young leaves, and
eeng glabrous petals Tek nias nir ¢ Ze ma usually con-
e section. North Auckland and 12; Bee c
gg- Tre rees to Tod Sec flowers 1-6; sepals, este Joa ha md
rusty-tomentose; pet ly dar to purple; capsules obese!
nire sulcate to convex de transverse section. North sd ru and
P. virgatum
ff. Shrubs ape 5—5.0 m. tall; sepals usually acuminate; capsule valves deed to
weakly sulcate in transverse section.
0.5-2.0 m. tall; male flowers 6-9, female flowers 1; petals cream
to yellow with a red stripe; capsules ovoid, acuminate or alm P beaked ` `
at apex. North Auckland. . P. pimeleoides
£g. Shrubs 1-5 m. tall; M ges Se petals purple; capsules aE SA z sub-
globose, . P. patulum
ee Branchlet sharply divaricatis 3 zg gees at juvenile stage or on zeg? Sieg o
adult stem; shrubs or trees 2—9 m. tall; flowers 4—12; petals pink to part)
ca ; Vitéinie Plateau . turneri
terminal, on ested branchlets (terminal on od
m), cad fascicles or solit Get capsules 4.5—
10.0 mm. long; seeds gel der Stee to Sege round to irregu
2—6 m. ta -obl ually piens Seier valves
dehiscing without, leaving a persistent papery ind
f. Adult leaves orbicular to obovate, obcordate to o obrus apex, with 2-4
i ei " ; peti wnally ese winged;
, inal, arres ranchlets; sepals
Ae tomentulose; petals pale Se mc or viis capsules e
sub-ovoid; placentas with funicles deus the base to the middl
^. uckland, Hawkes Bay, and Canterbur 17. P. obcordatum
ff. Adult leaves rarely orbicula ar to games obtuse to subacute at apex, with
obscure eins; petioles without win rings; flowers cut olitary (1-
3 in P. crassicaule), axi la ry beer? ormal or arrested branchlets;
bgl
sepals caducous, glabrous; petals furi: capsules cordate or subg
globose; placentas with falda near the mi
COOPER—AUSTRALIAN AND NEW ZEALAND PITTOSPORUM 139
g. Shrubs to 3 m. y agg pa ascending, not spinose; leaves obtuse at
base; pedicels 0.5 ; flowers solitary, €— or ll on
normal branch Let x "b: er mm. long, 1.0—1. ari d; P
8-12 mm. lon m. broad; ovary tomentose; capsules lobose, 8—
mm. eg Lë mm. broad. Hawkes Bay, Wellington, SS Marlborough.
P. rigidum
gg- Shrubs 4 to 6 m. tall; (— divaricate, almost spinose; ee acute to
attenuate at base; icels minute; flowers 1—3, terminal iem
arrested branchlets; sepals T “2. 5 mm. long, 0.5-1.0 . broad; petals
3.0 long, 1.0-1.5 br osil ovary Wé ee or age
ee "d 4. E 0 mm. men and broa
. Shrubs to 4 main een satis — wie vain
en dee 3.5-17.0 mm. en
fia "rr eiee ith 1-5 welt or vibes Se des sides ibus re
A 1 mm. broad; petals 4.0-7.5 mm. long, 1.2-
e mm tomentulose; op su rais to globose,
rugose. "Marlborough, Nelson Ca terbury, and Westland ....19. P. crassicaule
hh. Shrubs 6 tall; n branches cally Heena adals leaves
a xposed) Haesc-oblanesdla te to linear-oblon ng or ovate, 4-10 mm. long,
broad, re, occ c i crenate; sduit leaves "dide
usually pé 1-3 lobes ath on each side; flowers T —
mm. long, eer mm. aderit tals 4—6 mm. long, 1.5
mm. ih ovary glabrous; capsules im or emendi Sege
rugose. Hawkes ay Volcanic Plateau, Nelson and ege =
0. ». divaricatum
ee. Shrubs completely prostrate in the open, a tangled mass up to 1 m. alli in —
flowers solitary, terminal on short arrested branchlets; petals sti
oblan bulles variously colored; et valves dieing A due leaving a Sa
sis een m SN y endocarp covering the seeds. Volcanic Plateau, Nelson, and
. P. anomalum
bb. Sen shrubs seldom terrestrial; flowers terminal; petals yellow; eure ‘lipsoid.
New
€ She spesa shrub 0.5-1.5 m. tall, polygamous et leaves obovate
ovate-elliptic; male flowers about 10; female resi Zeg psules about 1.7 c
ong, 1.2 cm. broad. North Auckland to Ma riborou P NES in Goa RS pa 22. P. teen
b 1-5 m. tall, m
nui Ee d P. kirkii
aa. ra
b. sules See? a ‘pertistent papery endocarp. Australia
c. gë few (1—32), and relatively "pt sepals 4 4.0-10.5 mm. long; petals
18 mm. long, connate or connivent in a cylindrical or urceolate tube, tips eco
to Pins aad — lobed, woe or ov oid. Ss
d. Shru m. tall; leaf dien or oe sepals slightly imbricate at
GER E aei per i5 ree —2. ie
hrubs ; branchlets je EE entose when young, glabrescent;
bos ge cm. e Zap subcoriac eous to coriaceous, with appresse ty tomen
eath, acute to shortly scuminat, nich corel and sust bud
i in es to
.
ose, sessile; valves woody, 2-3 mm. thick, placentas bearing funicles from
the base to mear the apex; seeds 20-76. Queensland, New DG dag due
Mëtt revolutum
ee. Shrubs 1.56 m. tall; branchlets rusty-tomentose, not glabrescent; leaves
3.5 cm. long, membranous with sparse erect r hairs beneath, rusty -tomen-
tose on the veins, acutely ege ée to cuspidate, m argins eeneg peg
sometimes irregularly revolute; bud scales caducous; infloresc 32-
flowered; sepals tomentose; capsules elli psoid to ovoid, 2-valved, gef Mondo
1.3—2.1 cm. long, minute b rugose, —— Bee — ec? T^ Tun
f to the middle; see
Les ANS wem | dus A8, P. rubiginosum
dd. Shrubs or trees 5-13 m. tall; leaf ma H "markedly “undulate; frequently
connate in a tube splitting into 2 parts, one 1- t o 2-lobed, the other 3- to 4-lobed;
[Vol. 43
140 ANNALS OF THE MISSOURI BOTANICAL GARDEN
tals white; e Ea 2-valved, 1.0—1.4 cm. in diameter. Queenslan
mm South Wales and Vic 26. P. undulatum
cc. pri very numerous and celatively — sepals 1.1—3.5 mm. lon tals 5
mm. long, connivent in a cylindrical tube, or free and patent; ni dibus
to dipl asm id.
d. Sepals oblong to ovate, obtuse to ert capsules subglobose to spera m valved,
rarely 3-valved, stipitate; valves woody or coriaceous, about 1 mm.
e. Leaves lanceolate to ovate- el sepals oblong, obtuse, rusty- geed mentosé;
capsules subglobose to ea —2 cm. long, 2- or rarely eve ed; valves 1—
mm. thick, woody, c Y transverse section; placentas much thickened
and fused at base, Paria E nicles about the middle; « —— to 10 m. tall;
petals white; seeds 6—14, diia to black. Queensland............... 27. P. venulosum
ee. Leaves oblanceolate, oblong. , Or ovate; sepals : acute to —
white-tomentulose or gla Peri E es obovoid, E -valved; valves less than
mm. thick, woody, pana sulcate in transverse section; placentas much thic e
ened at base and apex and fuse s more or less completely to -— two chambers,
bearing several funicles from the base to the middle or at t ase.
f. Shrubs or small trees to 6 m. tall; leaves oblanceolate e oblong, entire;
petals yellow; o. 6-15 mm, in diameter; valves with a placenta set ar-
ing several short stout funicles esie een the bue m de — seeds 2
black. Een eege and Northern Queensland ............ P. ee
. Trees 13-30 tall; leaves rhombic to ovate, me apen ee petals
amet Geer about fee mm. long, 5-8 mm. broad; valves o a
plac "ta bearing 1 or nute fuhickn niz the base; seeds e: ack.
cien and and si alls New South Wales 29. P. bua s
dd. Sepals en to linear-lanceolate, acute to acuminate; capsules globose to "-
bose, wes, a 7-10 mm. in diameter; valves coriaceous, less than 1 mm.
thick, =. Wee placentas: with funicles from the ch to the
beri prias: ; black; trees or shrubs 8—20 m. tall; ye —
Quee and z Malay riens P. ferrugineum
bb. E pd hie a pan tent papery pete sed Zealan
; leaves coarsely serrate to entire; ME condensed, about
sry Dé Ce white with ri ad veins; pov kerja about 15 mm. long, 9 mm. broad;
placentas bearing about 12 funicles from the base to near the apex. Northwest
e P. dallii
"^
"^
Trees tall; leaves geng eg pes spreading, 30- to Pul qs
petals M ier 9-10 lon mm. a placentas i
funicles near the middle. Now. Agena im to Otag P. E
9. PITTOSPORUM PHILLYRAEOIDES DC. Prodr. 1:347. 1824. (T.: Leschenault
EMT)
Pittosporum angustifolium Lodd. Bot. Cab. 19: t. 1859. 1832, ex icon. & char
Pittosporum ligustrifolium A. Cunn. ex Loud. Hort. Brit. Suppl. 1: 597. 1832, nom. nud.
ittosporum longifolium Putterh Syn. Pittosp. p.15. 1839, oi char. Ga n.).
ittosborum roeanum Putterl. loc. cit. p.16. 1839, ex char. .
Pittosporum ligustrifolium A. Cunn. ex Putterl. loc. cit. p. SE ME (T.: A. Cunning-
D
o
o
bam s.n. [ Is.).
Pittosporum oleaefolium A. Cunn. ex D loc. cit. p. 17. 1839. (T.: A Cunningham
s.n. [31, 32, 143] ! Dick Harto Is.).
pese acacioides A. Cunn REN Nat. Hist. 4: 109. 1839. (T.: R. Brown s.n.
Pittosborum ice Lindl. in Mitch. Jour. Exped. Int. Trop. Austr. p. 97. 1848. (T.:
Neo 273 !).
tosporum lanceolatum A. Cunn. in Mitch. loc. cit. p. Hz 291. 1848, nom nud.
Seet bbillyraeoides DC. var. microcarpa S. Moore in Jour. Bot. 35: 163. 1897.
(T.: Spencer Moore s.n. !).
Shrubs to small trees 3-10 m. tall; branches pendulous; branchlets grayish-
brown, tomentose when young, soon glabrate. Leaves alternate, linear-oblong to
-oblanceolate, rarely elliptic-oblong, acute and apiculate at apex, attenuate
at base, entire, 2.5 - 12.6 cm. long, 2 - 15 mm. broad, tomentose when young, soon
1956]
COOPER—AUSTRALIAN AND NEW ZEALAND PITTOSPORUM 141
glabrate, very coriaceous, margins flat or revolute, costa raised to sunken above
and below, secondary veins usually obscure; petioles 4- 15 mm. long, 0.5 - 1.0
mm. broad, glabrous. Flowers pseudoterminal and axillary, 1 - 4, solitary or fas-
cicled, rarely in 3- to 4-flowered cymes, clustered at the ends of leafy branchlets;
peduncles and pedicels 3 - 15 mm. long, accrescent in fruit, sparsely tomentulose,
glabrate, subtended by a single leaf and a whorl of caducous bud scales 1 - 2 mm.
long. Sepals imbricate or coherent at base, 1 - 2 pairs occasionally fused to apex,
lanceolate, subacute to acute, 2 - 3 mm. long, 1.1 - 2.0 mm. broad, glabrous; petals
linear-oblong to linear-oblanceolate, sometimes slightly pandurate, obtuse, 8 - 11
mm. long, 2.0 - 3.5 mm. broad, coherent in a cylindrical or somewhat urceolate
tube for about two-thirds of their length, tips spreading to reflexed, yellow to
white; stamens 4.2 - 8.0 mm. long, anthers sagittiform, ovoid or oblong, 1.0 -
3.3 mm. long, 0.8 - 1.3 mm. broad. Pistil at anthesis slightly shorter or longer
than the stamens; ovary 3.0 - 5.5 mm. long, 1 - 3 mm. broad, tomentose, sometimes
Blabrate above; style 1.5 - 3.0 mm. long; stigma capitate, sometimes obscurely
lobed. Capsules ovoid to globose, obtuse to subacute, 2-valved, rarely 3-valved,
apiculate, 1- 2 cm. long, 9.5 - 19.0 mm. broad, glabrate, smooth, dark red or
orange without, yellow within; valves convex to sulcate in transverse section,
1.0 - 3.5 mm. thick, woody, with a conspicuous placenta, fused at the base, bearing
short stout funicles from the base to above the middle; seeds 4 - 24, red, irregular.
Occurs throughout Australia. Flowers from July to October.
Fig. 14. P. pbillyraeoides.
[Vol. 43
142 ANNALS OF THE MISSOURI BOTANICAL GARDEN
AUSTRALIA: NORTHERN TERRITORY: 60 miles N.E.C.2. [near Merde Ranges,
between Stuart Bluff Range and the Lander], Hill 283 (MEL); Northwest Petermann
Ranges, Basedow 159 (K); Petermann Ranges Basedow de 1 (K); eier locali, Hill
98 (NSW). QUEENSLAND: Burdekin ec sn. (K); rs fs ong s.n. (A
K); on stony ridges between Aramac mé" Muttaburra, Brass v W bite T Ki between
Bowen Downs and Mueller's Range, Birch s.n. (MEL); Texas, Boorm n. (A); Md
Edwards s.n. (A); Barcoo, Schneider s.n. (MEL); Eidsvold, Beeroti i s.n. (A); in
brown fine sand, Glenber, C. 40 miles S.E. Charlesville, Warrego District, Everist 3397
(K); Barunga, north of Miles, Belson s.n. (A); in brigalow scrub in heavy soil, C. 890',
Wandoan, Hubbard 5024 (K); Wallumbilla, Jale s.n. (A, K); Dalby, White 9032
(A, K); Moreton Bay, Mallard s.n. (GH); Ipswich, Nernst 35 (MEL); Bumble Station,
70 mil north of Mungindi, Cambage 4404 (NSW); 558', in heavy black soil in open
Eucalyptus forest, Dirranbandi, Hubbard 5572 e K); Noondoo, Maranoa District,
Everist 7 dà v K); EE epiphytic i " e Eucalypt (E. corymbosa?) near Warwick,
E. Tbom 6 (K); Silverwood, Wbite 1712 (A); Inglewood, Boorman s.n. (NSW);
Gonia. See s.n. Wa NEW SOUTH WALES: Moru-Mungundi District, Kerry
s.n. (NSW); Uriseno-Thurloo Downs, Boorman s.n. (NSW); Middle Warrego, Bullock
s.n. (MEL); between 40 and 50 miles N.W. of Collarenebri, Jackson s.n. (NSW); AD.
pee $5. ( D; New Angledool, Paddison s.m. (A); Narrabi W., Boorman s.
B W
Nyngan Exp. Farm, Downing s.n. (BISH, NSW, US); Co bbora-Boomley Rd., 1700,
quartzite in rocky scrub- pu Jobnson & Erans s.n. (NSW); Currabubula, Camb-
age 3575 (NSW); Cobar, Hadley s.n. (NSW); Trangie, Maclean 6 (NSW); Barrier
Range, Irvine s.n. (BM, MO); Ricken. Hill, d s.n. (NSW); in open die Mundi
Mundi Station, Broken Hill, Constable s.n. (N xu sandy flat, "Wilmatha, Condobolin,
Hadley 29 (A, NSW); Wralone: Boorman s.m. (NSW); same locality, House s.m. (K,
MO); Wollongong, Wilkes s.n. (US); E. Mirrool, Campbell s.n. (NSW); in barren
country S.W. of the Lachlan River, A. Cunningham 35 (K); in granite soil at side of =
Ardlethan District, Constable s.n. (AK, K, NSW, US); Temona, Dwyer s.n. (MO, N W);
junction of Dering and Murray rivers, Holding s.n. (MEL); Wagga, Helms s.n. (NSW). d
Yenda, Searcy s.n. (NSW); without definite locality; Fraser s.m. (A, K); VICTORIA:
Mildura, ler: s.n. (UMEL); Lower Murray River, epee s.n. (BM, GH, K, MO);
Box Ridge, Robinvale, Murray River, Froggatt s.n. (NSW); Benjeroop, Murray River,
C. French Jr. s.n. (NSW); Swan Hill District, C. French p s.n. (GH, MEL); Deniliquin,
Crawford s.n. (NSW); Dimboola, d'Alton 31 (NSW); Wimmera, Mueller 18 (GH
same locality, Williamson s.n. (NSW); same locality, Walter sm. (GH ee scrub,
amarooka, near Bendigo, Paton s.n. (MEL); Mason C. Frencb Jr. s.n. W); ranges
bearing S.W. from Camp 29 [vicinity of Mt. Cole, near Ararat], bi? s.n. (BM).
Ree AUSTRALIA: Gov deum E e Basedow 112 (NSW); Elder Expedition,
Camp 10, Helms s.n. (K, ME NSW); Musgrave Range, Lord s.n. Gei Ooldea, Ker-
sbaw s.n. (MEL) ; Commonw anm Hill, near Ooldea, Basedow 84 (NSW); Mt. poseer
Kocb 7 (K); Flagstaff Point, near Streaky Bay, Rogers s.n. (NSW); äre Bay, Crocker
CA
s.n. ( ; Port Augusta, Lea s.n. Sarl Port Lincoln, Browne C. 29 (MEL); same
locality, Schomburgk s.n. (US); Memory Cove, near Port Lincoln, SE s.n. (NSW);
shores of Bay III, South Coast, R ecd 5447 (BM, K, MO); 10 miles E. of Mann
Bir
Australia, Sturt s.n. (BM); without locality, Conservator of Forest s.n. (NSW); WESTERN
AUSTRALIA: in stream-bed, Nickol ak Gregory Expedition s.n. (MEL ); Sharks Bay,
Mueller s.n. (MEL); barren sandy downs, Dirk Hartog's Island, A. Cunningham s.n.,
32 (K), 143 (BM); Murchison River, Oldfield s.n . (GH, K, MEL); mie Flat, Gray
s.n. (MEL); Tabletop Hill, Northampton, Comp ll 18 (K); South Island, Houtman's
Abrothos, Gilbert 98 (BM O); W. , S. of Geraldton, Morrison 13305 (K);
saltbush plain south of tank vd Sech Zeng s.n. (BM); Arrino, Fitzgerald sn.
(NSW); arid shores of Rottnest land, A. Consissbem s.n., 30 (K), 142 (BM); ™
1956]
COOPER—-AUSTRALIAN AND NEW ZEALAND PITTOSPORUM 143
arenosis sylvae orae occidentalis insulae Rottnest, Preiss 1297 (GH, MO, P); Swan River,
Drummond s.n. (K), 31 (BM, K, MO); 57, 76 (K),76 (BM); R. des Cygnes, Drummond
76 (P); same locality, Preiss 1297 (P); limestone rocks, face of cliff, Claremont, Andrews
37 (BM, K); Kellerberm, Vachell s.n. (NSW); Merredim, Koch 2983 (K, MEL, NSW);
near rock outcrop about 6 miles N.W. s Wie Bailey 182 (CANB); 70 miles N.
of Coolgardie, C. A. W bite s.n. e: n Camp, Kurrawang, Maiden s.n. (NSW); E.
of Kalgoorlie, Transcont. Ry. Survey, e s.n. ( ); Point Eeron; on ns dunes near
sea, Burbidge 1962 (CANB); on "aub flat in plain, 7 miles west of Dea irt s.n.
(CANB) ; Benner Bay, Menzel 56 ( NSW); eink definite iliy, Vim 6 (MO).
WITHOUT LOCALITY: Exped. Baudin, Leschenault s = M); N. Holl., Lhotsky $.n. (P);
Mitcbell s.n., 228, 229, 273 (K); [228, 229, on the B ando River near 147^ X 22^, 27',
Mantuan Dawns bieiin Kilsyth and valley ^ Nogoa rms 1475? X 24V?.]
Flowers which may be female have pistils slightly longer than the stamens,
Sp capitate stigmas, styles about 1.5 mm. long, ovaries 4.0- 5.5 mm. long,
2.5 - 3.0 mm. broad, and sagittiform to ovate anthers 1.0 - 1.5 mm. long, borne
on filaments only 3 - 4 mm. long. Flowers which may be male have pistils slightly
shorter than the stamens, smaller capitate stigmas, styles 3.0 - 3.5 mm. long,
ovaries 3 - 4 mm. long, 1 - 2 mm. broad, and oblong anthers 2.0 - 5.3 mm. long
borne on filaments 3.5 - 6.0 mm. long. Intermediate forms with large anthers
and plump ovaries occur.
Pittosporum phillyraeoides is grown as a hedge, windbreak, and as an orna-
mental shrub in the United States (California, Arizona, and Florida), France,
Australia, and New Zealand; and in the drier parts of Australia it is used as a
stock food
The variability of the leaves was noted by Bentham!””, who commented:
his species, apparently spread over the whole desert country of Aus o cague
rtions
ss confounded with any other Ar: 2 — or the pro; of
its leaves, flowers and fruit. In some of the E are MY
e inches long, and fully an inch aide e, bibi in in a rae a ber wie ern and s
estern ones they attain 4 or 5 inches in length with a eg of veli 2-or. 3 gang
In the material I have examined there is a specimen from Shark Bay, western
Australia (Mueller s.n. MEL), with elliptic-oblong leaves about 3.5 cm. long and
up to 1.7 cm. broad, covered with appressed tomentum beneath, but this extreme
is connected with the more common form by a number of intermediates, and I am
unable to define any varieties. The variability of the capsules in size and shape
must be mentioned. Again the extremes are connected by intermediates and I am
unable to recognize var. microcarpa S. Moore.
Vernacular names: Butter Bush, Willow or Willow Tree, Berrigan, Locket
Bush, Butterwood, Bell’s Orange, Quinine Tree, Weeping Pittosporum, Native
Willow, Poison-berry Tree, Apricot Tree, West Australian Willow, Macla (Paroo
River Blacks) , Derrine (Dubbo Blacks).
159 Fl. Austr. 1: 113. 1863.
[Vol. 43
144 ANNALS OF THE MISSOURI BOTANICAL GARDEN
10. PrrrosPoRUM BICOLOR Hook. in Jour. Bot. 1: 249. 1834. (T.: Lawrence
$.n.).
Pittosporum buegelianum Putterl. in Endl. Nov. Stirp. Dec. 43. 1839, ex char. (T.: Hügel
s.m.).
Pittosporum discolor Regel in Gartenflora 1:133. 7. 15. 1852, ex icon.
Shrubs to small trees 5 - 16 m. tall; branchlets ascending; branchlets grayish-
brown, the young parts hoary or rusty-tomentose, soon glabrous. Leaves alternate,
rarely subverticillate, oblong-lanceolate to linear, obtuse or apiculate at apex, acute
or attenuate at base, entire, 2 - 7 cm. long, 3 - 18 mm. broad, glaucous-green and
glabrous above, white-, yellow- to brown-tomentose beneath, coriaceous, revolute,
costa and secondary nerves usually distinct above, costa distinct beneath, but
secondary nerves obscure; petioles 2-5 mm. long, 1-2 mm. broad, tomentulose
when young. Flowers pseudoterminal and axillary, 1 - 4, solitary or fascicled,
clustered at the ends of leafy branchlets; pedicels 3 - 19 mm. long, accrescent in
fruit, brown-tomentose, subtended by numerous caducous bud scales about 1 mm.
long. Sepals imbricate at base, lanceolate, acute, 2-6 mm. long, 1-3 mm. broad,
puberulous; petals lanceolate-oblong, subacute, 9 - 15 mm. long, 2 - 4 mm. broad,
free or coherent at the base, spreading from above the middle, yellow with red
veins, yellow marked with red, or reddish purple; stamens 3 - 9 mm. long, anthers
elliptic-oblong, 1 - 3 mm. long, 0.5 - 1.0 mm. broad, Pistil at anthesis as long as
the stamens, rarely almost twice as long; ovary 3 - 5 mm. long, 2 - 3 mm. broad,
villous; style 1.5 - 5.0 mm. long; stigma capitate and obscurely 4-lobed on short-
styled pistils, weakly capitate to truncate on long-styled pistils. Capsules sub-
globose, 2-valved, rarely 3-valved, 7 - 13 mm, in diameter, tomentose, glabrate;.
valves convex in transverse section, less than 1 mm. thick, coriaceous, with a con-
spicuous placenta bearing short stout funicles from the base to just above the
middle; seeds 10 - 20, yellowish-red to reddish-black, irregular.
Fairly common from sea-level to 1300 m. in Tasmania, less common in eastern
Kee and southern New South Wales. Flowers from July to December.
TH WALES: Bibbenluke, near Bombala, Edwards s.n. (NSW);
ane State E oe Beazcville LI (NSW); Brown Mountains near Littleton, Betche s.n.
H5 g erbrook Forest, Lothian s.m. (GH); Blacks Spur, Deane s.n.
BE NSW); same locality, Morris 1441 (NSW); Milla Milla, Clinton s.n. (NSW);
Bonang, Baeunton s.n. (NSW); near Mt. Ellery, Merrab s.n. (MEL, NSW); Genoa
District, Baeunton s.n. (MEL, NSW); along Dividing Range, Stirling s.n. (NSW); same
B 4
(NSW) ; Mt. Disappointment d Dandenong, Mueller s.n. (MEL, NSW); near Melbourne,
Cecil 219 (K); p Gully, Staer s.n. (NSW); Emerald, ra ái. (BM); dps
Cookson s.n. (UMEL) ; Korumburra, Baker s.n. (MEL); East Gippsland, C. French Jax
(GH); same et Mueller s.n. (US); same E Stirling s.n. (BM); Corner Ka
Mueller s.n. Xn W); Sealer’s Cove, pre s.n. (BM, K, NSW); near Cape Otway, Lucas
s.n. (MEL, SY). TASMANIA: g's vd pei gas (MEL, NSW); talas Head,
s.n. (HO); Chilton, Surrey Hills, Gunn s.n. ad May A Meus 3000-3500’, in mr
open situations on the sides or tops of mountains, [Gunn] 651 (K); gum forest assoc
2700', Dove Valley, Weindorfer s.n. (K); De hora e See s.n. (NSW); Deb. to to St
1956]
COOPER—-AUSTRALIAN AND NEW ZEALAND PITTOSPORUM 145
Helens, Ford s.n. (NSW); near = Patrick’s River, eperdi ILI9 des NSW); same
locality, Gunn 154 (NSW); top of Western ves 3500', Gunn 651 (K); Launceston,
Gunn 154 (MO); same loc Mir. Gunn dog (BM. K); same d. Stuart s.n.
(MEL, NSW); Cataract Gorge, Rupp 31 (NSW); Cora Linn, N. Esk, Gunn 154/1842
i 0
an. );
Grand s.n. (MEL); same locality, in damp shaded woods on the low shor unning-
bam 33 (K); beech forest between Lake Huegel and Lake St. Clair, “Burbidge 3359
(CANB); 3500’, Lake Fenton National Park, Lord s.n. (HO, K); rocky slopes above Lake
Fenton, Mt. Field National Park, Burbidge 3280 (CANB); Maria Island, summit of
mountain, c. 2000”, Gibbs 6395 (K); Hobart Town, Siemsen s.n. (MEL, 2225 Lenah
valley near Hobart, White 8406 (A, BM); p prope Derwent, Brown 5448 (BM, K ; MO);
Diabase Hill, Collins Vale in Fairy Glen, Long 1008 (HO); open places in nont forest,
2000”, Collinsvale, Comber 1550 (K); 300’, Waterworks, Hobart, Olsen s.n. (HO); 800’,
near waterworks, Atkinson 93 (CANB, HO, K); 1000’, z^ Peak, Hobart, Curtis s.n.
(BH); rocky hillside above springs, Mt. Wellington, Long 3 55 (HO); Mt. Wellington,
Gunn 154/1842 (NSW), 651/1842 SC SE locality, Verreaux 1961 (K, P); same
locality, Maiden & E age s.n. p. Saf ers og bank, New Town side, Mt.
Wellington, Long 375 (HO, K); AU . Wellington, ME 355 (A);
Kangaroo Bottom, Hooker 844 eg Eder Valley, y hera 842 (K); Mt. Nelson, Rod-
way s.n. (HO); Little Plain, Simson s.n. (AK); AS Bay, [Bligh, e Aë EI Gs
s.n. (BM, K); Recherche Bay, Maiden. 3991 (BH, NSW). wirHouT LocaLiTY: Archer
s.an. (NSW); Blyth sn. (NSW); Caley s.n. (A); Gunn 154, 651 (GH, K); Hannaford
sn. (NSW); La Billardiére s.n. (BM, GH, MO); Lawrence s.n. (K).
The variation in plant form and leaf size in this species was noticed by
Hooker!, who remarked:
One of the most rers plants in the pana eegene), dod in low grounds, a
tree 30 - 40 feet high; with a trunk a n diameter, but gradually
Pr stunted scrubby bush on sending t em ntains. The Shanes of habit
characters on ascending are so gradual that I: haser rigidly define the varieties,
and
of which the fo tawing are the best Lé? on low ground, branches lax, twiggy,
with ig tnt flat, linear-obovate or oblong leaves, 1-2 inches long, with massid
olu scendi he branches
short, dei , robust, erec esti de e nsely eda erect, shorter, randi uh
bla coriaceous margins very revolute, hated below with ferrugineous silky and
The Re also vary in size and form. Flowers which may be "Tische have
Capitate stigmas, styles 1 - 2 mm. long, plump ovaries, filaments 2- 3 mm. long,
and possibly abortive anthers about 1 mm. long. Flowers which may be male
ve weakly capitate to truncate stigmas, styles 3 - 5 mm. long, slender ovaries,
filaments 3 - 5 mm. long, and functional anthers 2- 3 mm. long. Pedicels of
female flowers are usually shorter than those of male flowers. Between the two
extremes described, intermediate forms have been found; e.g. Lord s.n. from 3500'
at Lake Fenton National Park has a truncate stigma, style 3 mm. long, filaments
4.5 - 5.0 mm. long (i.e. it appears to have male flowers) but the anthers are only
1 mm. long, apparently abortive; Olsen s.n. from 300' at the waterworks, Hobart,
has a capitate stigma, style 1.5 mm. long, and anthers 1.5 mm. long (i.e. it appears
to have female flowers) but the filaments are 4.5 mm. long as in male flowers.
Pittosporum bicolor is cultivated in California and England.
Vernacular names: Whitewood, Tallow-wood, Tolosa-wood, Dragon’s Blood
Pittosporum (Tasmania), Cheesewood (Victoria), Banyalla (Victoria, aboriginal).
160 EL Tasman. p. 38.. 1860.
[Vol. 43
146 ANNALS OF THE MISSOURI BOTANICAL GARDEN
d
a
Ja
PEE E
D AR
i Ka m
-
em ton
—— -———
m 2 ae e EE |
Fig. 15. P. bicolor.
Fig. 16. P. o'reillyanum.
11. PrrrosPoRUM o’REILLYANUM White, in Proc. Roy. Soc. Queensland 47:53.
1936. (T.: White 6173!).
Shrubs 1 - 2 m. tall; branchlets grayish-brown, spinose, brown-tomentulose
when young, soon glabrate. Leaves alternate, sometimes clustered on arrested
branchlets, ovate to elliptic-oblong, acute to cuspidate at apex, obtuse at base,
entire, 2.5 - 20.0 mm. long, 2 - 5 mm. broad, green above, paler beneath, sparsely
tomentulose when young, soon glabrate, margins flat or recurved, costa immersed
above and beneath, secondary nerves 2 - 4 per side, anastomosing, distinct or
obscure; petioles up to 1.0 mm. long, sparsely tomentulose when young, soon
glabrate. Flowers axillary or terminal, on arrested branchlets, solitary; pedicels
1 - 2 mm. long, accrescent in fruit, brown-tomentose, subtended by 1 to several
leaves and caducous brown-tomentose bud scales 0.5-1.0 mm. long. Sepals
irregularly connivent at the base, subulate, acute, 1.7 - 2.0 mm. long, 0.5 - 0.8
mm. broad, tomentulose; petals linear to linear-oblanceolate, subacute to acute,
6.0 - 9.5 mm. long, 1.2- 1.5 mm. broad, connivent in a cylindrical or slightly
spreading tube with reflexed tips; stamens 6.8 - 7.0 mm. long, anthers oblong to
ovate, 1.8 - 2.0 mm. long, 0.7 - 1.0 mm. broad, filaments filiform, coherent to the
petals. Pistil at anthesis equal to the stamens; ovary 3 - 4 mm. long, 1 mm. broad,
tomentulose; style 2.5-3.0 mm. long; stigma subcapitate, obscurely 2-lobed.
Capsules globose, 2-valved, apiculate, 7 - 12 mm. broad, glabrous, smooth; valves
convex in transverse section, less than 1 mm. thick, coriaceous, with a placenta
bearing 3 - 4 strap-like funicles, up to 2 mm. long, from the middle to the base;
seeds about 6, light red, irregular.
1956]
COOPER— AUSTRALIAN AND NEW ZEALAND PITTOSPORUM 147
Occurs at high altitudes in southeast Queensland. Flowers in September and
October.
USTRALIA: QUEENSLAND: very common in rain eise usually weighted down
na masses of liverworts and mosses, 4000', Mt. Hobwee, Lam gton N National Park, White
6173 (A, BM, K, MO); in Notbofaus Kaes 3700', Mt. Mer , MacPherson Range
Jobnson s.n. (NSW); MacPherson Range, Perry 512 (CAN B).
12. PrrTOSPORUM UMBELLATUM Banks & Soland. ex Gaertn. Fruct. et Sem. 1:
286. 1.50. 1787-88. (T.: Banks & Solander s.n.!).
Pittosporoides umbellata Soland. ex Gaertn. loc. cit. 1787-88, nom. nud. in synon.
Small trees 4 - 10 m. tall; branchlets grayish-brown, the young parts sparsely
brown-tomentose when young, soon glabrate. Leaves subverticillate, dark green
above, paler beneath, glabrous, coriaceous, margins thin, flat and brown-pilose
when young, thickened, revolute, and glabrate when mature, costa raised above and
beneath, frequently sparsely brown- to white-tomentose at base, secondary veins 8 -
15 per side, anastomosing, obscure above, distinct beneath. Juvenile leaves obovate
to oblanceolate, usually crenate, lobed, or parted; adult leaves elliptic to oblong-
lanceolate, entire, or rarely crenate, acute or acuminate at apex, acute to attenuate
at base, 2.5 - 10.0 cm. long, 1.5 - 5.0 cm. broad; petioles 6 - 22 mm. long 1-2
mm. broad, brown-tomentose when young, glabrate. Inflorescences terminal or
sometimes becoming lateral on development of a leading shoot from an axillary
bud, in umbels up to 20-flowered; pedicels 4 - 25 mm. long, accrescent in fruit,
brown-tomentose, subtended by an approximate whorl of leaves and numerous
caducous, glabrous, ciliate bud scales up to 12 mm. long. Sepals slightly imbricate
at base, lanceolate, acute to acuminate, 4.5 - 9.0 mm. long, 1.5 - 3.0 mm. broad,
sparsely ciliate; petals linear-oblong, subacute to obtuse, 11.0- 12.5 mm. long,
2.5 - 5.0 mm. broad, loosely coherent at the base, spreading from above the middle,
dull red; stamens 4 - 8 mm. long, anthers sagittiform to elliptic-oblong, 1 - 3 mm.
long, 0.6 - 1.3 mm. broad. Pistil at anthesis slightly shorter or longer than the
stamens; ovary 2.5 - 3.5 mm. long, 1 - 3 mm. broad, villous; style 3 - 4 mm. long;
stigma capitate and obscurely 4-lobed, or truncate. Capsules tetragonous or 4-
lobed, 2-valved, 7 - 12 mm. in diameter, green to black, sparsely pubescent; valves
concave in transverse section, less than 1 mm. thick, with much thickened rims,
coriaceous, with a conspicuous placenta bearing short stout funicles from near the
base to just above the middle; seeds 9 - 14, black, irregular.
Occasional from the North Cape to the Coromandel Peninsula, North Island
bf New Zealand, in forest remnants on the sea coast and coastal islands. Reputed
to occur as far south as Gisborne in Poverty Bay, but I have not seen specimens
from south of the Thames district.
EW ZEA NORTH KLAND: summit of Ounuwhao, 950', Spirit's Bay,
Cooper 24478 ( AK); DE Stee near sea-level, q odo ue Spirit's Bay, Cooper
24448 (A Spirit's Bay, does sm. (AK);
Bay, Wilkens 53 ep flag M US); in Agatbis Greg, fore Saies, agr seti Harbour,
Cooper 35960 (AK, MASS MO); Totara North, Whangaroa Harbour, Carse s.n. (
Koukoumiko, eg des fle; Raoul s.n. (P), 81 (K); Bay of Islands, Wilkes s s.n. (US);
[Vol. 43
148 ANNALS OF THE MISSOURI BOTANICAL GARDEN
same locality, A. Cunningham 29/1826 (K); same locality, Fraser s.n. (K); same locality
and Tippona, Wilkes s.n. (K); Opua, Bay of Islands, Moore & Cranwell s.n. (AK); Helena
Baylis s.n. (K); same locality, Turner s.n. (AK); same locality, Turbott s.n. (AK);
aranga (Hen Island), Moore & Cranwell s.n. (AK). THAMES: Great Barrier Island, Kirk
s.n. (AK), 90 (K); same locality, Hynes s.n. (AK); same locality, near State forest Service
Station, Lloyd & Anderson 36309, 36310 (AK); same locality, coastal forest, Whangapara-
para, Moleswortb s.n. (AK); Little Barrier Island, Kirk 90 (K); same locality, Cheeseman
sn. (AK, US); same locality, Shakespeare s.n. (AK); same locality, Adams s.n. (AK);
same locality, 1000', Turner s.n. (AK); Cape Colville, Kirk s.m. (AK); Te Moehau, near
Stony Bay, c. 1000 - 1100', Moore & Cranwell s.n. (AK, K); Shag Bay, Coromandel
Peninsula, Moore & Cranwell s.n. (AK, K); Coromandel, sea-level, Mackie s.n. (AK);
Kennedy Bay, Matthews s.n. (AK); in sylvis prope Opuragi, Banks & Solander s.n. (BM);
Mercury Bay, Kirk 6 (A, BM, GH, MO, US); North of Buffalo Beach, W better 1980
(AK); Tapu, Thames Coast, Molesworth s.m. (AK); Thames, Cheeseman s.m. (AK);
same locality, Adams s.m. (AK); same locality, near sea, Petrie s.m. (A). WITHOUT
ocaLrrY: Banks & Solander s.n. (AK, MO, US); Colenso s.n. (K); A. Cunningbam 30
(U); R. Cunningham 192, 613 (K); Hooker, 350, 351 (K); Kirk s.n. (K); Wilkes s.n.
(GH).
7
Fig. 17. P. umbellatum.
Heterophylly in this species has been described and illustrated in the discussion
of taxonomic criteria. Flowers occur which are probably female. They have
capitate, 4-lobed stigmas which are exserted 1 - 2 mm. beyond the stamens, and
sagittiform anthers only 1.0 - 1.5 mm. long, which appear to be abortive. The
pedicels are usually short (4 - 14 mm.). Flowers which are perhaps male have
truncate stigmas, styles 1.0 - 1.5 mm. shorter than the stamens, elliptic-oblong
anthers 2 - 3 mm. long, and ovaries which are slender and appear to be abortive.
The pedicels are usually long (10 - 25 mm.). Intermediatė forms occur with
weakly capitate stigmas and pistils which are more or less equal in length to the
stamens. I have not found flowers with capitate and truncate stigmas in the same
inflorescence, or on the same specimen, but field studies are required to determine
the significance of these morphological differences.
A form of the species having leaves gradually narrowed below and cordate
capsules has been described as P. umbellatum var. cordatum by Kirk in N.Z. Inst.
Trans. & Proc. 4: 264. 1872. (T.: Kirk s.n. A, AK, GH, MO, 90K.) The original
,
1956]
COOPER—AUSTRALIAN AND NEW ZEALAND PITTOSPORUM 149
specimens came from the Great and Little Barrier Islands where there are several
puzzling forms, and until further studies are made of these populations the status
of the variety is obscure.
A cultivated specimen from the Scilly Islands has been seen, Hill s.n. (K).
Vernacular name: Haekar
13. PrrrosPORUM VIRGATUM Kirk, in N.Z. Inst. Trans. & Proc. 4: 264. 1872.
(T.: Kirk s.m. !).
Pittosporum virgatum var. crataegifolia Kirk, loc. = a 1872. déi "dle s.n. !).
Pittosporum virgatum var. serratum Kirk, bác: cit. 1872. (T.: Kirk s.
plots Ag in virgatum var. sinuatum Kirk, Dti Fl. NA. p. et. 1899. (T.: Kirk
Pon matthewsii Petrie, in N.Z. Inst. Trans. & Proc. 52: 17. 1920. (T.: Matthews
s.n. Y).
Small trees to 8 m. tall; branchlets brown, rusty-tomentose when young, soon
glabrate. Leaves alternate, frequently somewhat crowded at the tips of the
branchlets; juvenile or lower leaves linear, entire or variously lobed or parted,
1.0 - 3.8 cm. long, 1 - 7 mm. broad, rusty-tomentose, glabrate; intermediate leaves
lanceolate to oblong or obovate, usually lobed or parted, 1.3 - 5.2 cm. long, 0.9 -
2.7 cm. broad; adult or upper leaves oblong, occasionally linear, oblanceolate, or
elliptic-oblong, entire, rarely sinuate or lobed, 1.8 - 7.0 cm. long, 0.4 - 2.1 cm
broad; margins flat to slightly undulate, sometimes a little thickened and revolute,
costa immersed above, raised beneath, tomentose, secondary veins 7 - 9 per side,
anatomosing, obscure above, distinct beneath; petioles 1 - 7 mm. long, 0.5 - 1.0
mm. broad. Flowers terminal, 1 - 6, fascicled or solitary; pedicels 5 - 9 mm. long,
accrescent in fruit, rusty-tomentose, subtended by an approximate whorl of leaves
and several minute rusty-tomentose caducous scales. Sepals not imbricate at base,
oblong to linear-lanceolate, acute, 3.5 - 6.5 mm. long, 1.0 - 2.5 mm. broad, rusty-
tomentose; petals linear-oblanceolate to linear-oblong, acute, 6 - 13 mm. long, 2 -
3 mm. broad, coherent in a tube to above the middle, tips reflexed, dark red to
purple, rarely yellow, pink or white; stamens 4- 7 mm. long, anthers sagittiform
or oblong-ovate, 1 - 2 mm. long, 0.4 - 1.0 mm. broad. Pistil at anthesis equal to
or slightly longer than the stamens; ovary 2 - 4 mm. long, 1.0 - 2.5 mm. broad,
rusty-tomentose; style 1 - 5 mm. long; stigma capitate and obscurely 2- or 4-lobed
on short-styled pistils, truncate on long-styled pistils. Capsules subglobose to
subpyriform, 2-, rarely 3-, valved, apiculate, 11 - 16 mm. long, 10 - 13 mm. broad,
rusty-tomentose to BCC obscurely rugose; valves sulcate to convex in trans-
verse section, about 1 mm. thick, coriaceous, with a thickened placenta bearing
stout flattened funicles up to 2.5 mm. long from the base to above the middle or
apex; seeds 1 - 16, black, irregular.
Occurs from the hills behind Ahipara on the west coast to Coromandel
Peninsula on the east coast of the North Island of New Zealand. Flowers in Sep-
tember and October.
[Vol. 43
150 ANNALS OF THE MISSOURI BOTANICAL GARDEN
Fig. 18. P. virgatum.
New ZEALAND: NORTH AUCKLAND: Kaitaia, Matthews s.n. (AK, MO); Taumata
Mahoe, near Kaitaia, Matthews s.n. (AK); range west of Okahu, Kaitaia, Matthews s.n.
(AK); at edge of forest, high country, Pukepoto, Matthews S Carse s.m. (AK); Whang-
aroa, Kirk 96 (K). THAMES: Great Barrier Island, Kirk s.n. (GH), 521 (BM); same
locality, Kirk 521 (US); same locality, Matthews & Petrie s.n. (AK); same locality, Hynes
26568 (AK); Port Fitzroy, Great Barrier Island, Kirk s.n. (AK), 87, 88 (K); back of
Whangarparapara, Molesworth 22171 (AK); Coromandel, Cheeseman s.m. (AK, BISH,
GH, ); Kennedy T Coromandel, Matthews s.n. (AK, MO); Waipuna, Kennedy
)
The plants recognized by Kirk as var. cra/aegifolia, var. serratum and vat.
sinuatum are juvenile froms. The distinguishing characters of P. matthewsii from
Kennedy Bay were stated to be the smaller and narrower linear-oblong leaves and
the subpyriform capsules. Among the large collection left by Matthews are speci-
mens of P. virgatum from both Kaitaia and Kennedy Bay with subpyriform cap-
sules; and Michie has sent me material from the Kaitaia area with linear leaves
which match those of P. matthewsii, collected at Kennedy Bay (the type locality).
As the morphological differences are slight and both of the main populations show
them I have not recognized P. matthewsii as a distinct population.
The flowers are of two kinds. Flowers which may be female have capitate,
2- or 4-lobed stigmas, styles 1.0 - 2.5 mm. long, ovaries about 4 mm. long and 2.5
mm. broad, short stamens 4 - 5 mm. long, with sagittiform, possibly abortive
anthers about 1 mm. long. Flowers which may male have truncate stigmas,
styles 3.5 - 5.0 mm. long, ovaries 2.0 - 3.5 mm. long, 1 - 2 mm. broad, stamens
5 - 7 mm. long with oblong-ovoid, apparently functional anthers 1 - 2 mm. long.
14. PrrrosPORUM PrMELEOIDES R. Cunn. ex A. Cunn. in Ann. Nat. Hist. 4:
108. 1839. (T.: R. Cunningham 40, 6181).
Shrubs 0.5 - 2.0 tall, erect or prostrate; branchlets slender, brown, the young
parts white or brown-tomentulose, soon glabrous. Leaves alternate, sometimes
subverticillate, elliptic, elliptic-obovate, oblanceolate, linear-oblong, or linear,
acuminate to obtuse at apex, acute to attenuate at base, entire or obscurely crenu-
late, 0.5 - 4.3 cm. long, 0.5 - 13.0 mm. broad, pale green above, lighter beneath,
sparsely tomentulose and ciliolate when young, coriaceous, margins thickened, flat,
1956]
COOPER—AUSTRALIAN AND NEW ZEALAND PITTOSPORUM 151
costa raised above and beneath, secondary veins anastomosing to form large areoles,
obscure above, obscure or distinct beneath, sessile or with petioles up to 0.5 mm.
long. Flowers terminal or sometimes becoming lateral on development of a leading
shoot from an axillary bud; male flowers 6 - 9, fascicled; female flowers solitary;
pedicels filiform, 2 - 9 mm. long, accrescent in fruit, puberulent, subtended by an
approximate whorl of leaves and several caducous, glabrous, ciliolate bud scales
2-3 mm. long. Sepals not imbricate at base, linear, acuminate, 2.5 - 5.0 mm.
long, 0.5 - 1.0 mm. broad, glabrous; petals linear, acuminate, 7.5 - 11.5 mm. long,
1.0 - 2.2 mm. broad, coherent in a tube to about their middle, spreading to reflexed
above, cream to yellow with a central red stripe; stamens 2.5 - 6.5 mm. long,
anthers sagittiform, or ovate to oblong, 0.5 - 2.0 mm. long, 0.2 - 0.8 mm. broad.
Pistil at anthesis equal to or slightly longer than the stamens; ovary 1 - 2 mm.
long, 1.0 - 1.5 mm. broad, villous; style 1.5 - 3.5 mm. long; stigma capitate and
obscurely 2-lobed, or almost truncate. Capsules ovoid, 2-valved, acuminate, 6 -
12 mm. long, 4 - 6 mm. broad, green to brown, rugose, pubescent when young,
glabrate; valves convex in transverse section, less than 1 mm. thick, coriaceous,
with a slightly thickened placenta bearing flattened funicles up to 3 mm. long
from base to near the middle; seeds 7 - 17, black, irregular.
KEY TO THE SUBSPECIES
Shrubs usually eb 0.5—2.0 m. p HV. Ca linear-oblong or uen 0.5-4.3 c.
. lon,
long, 0.5—9 . br vM Sepa 54.0 ong; petals 7.5-9.0 m Zeg Nort!
Aad. $ itai. aia to the Ka i P. p. Goen
€ prostrate; leaves dio: or Ree 9-13 mm. long, 4.5-7. e mm. broad i
un, 1.0-2.7 cm. long and 6-13 mm. broad in shade. Beds 4. 5-5. 0 mm. mE ium
"n 0-11.5 mm. long. North Auckland: Kerr. Point, North Cape P. p. major
14a. PITTOSPORUM PIMELEOIDES ssp. pimeleoides
pir un pimeleoides A. Cunn. ex Putterl. Syn. Pittosp. p. 15. 1839, ex char. (T.: R.
ingham s.n.
P A Re Sei R. al ex days hg Pittosp. p. 15. 1839. (T.: bri se)
ee Sien um R. Cunn. ex A. Cunn. in Ann. Nat. Hist. 4: 108. 1839. (T.:
ingham 40, 617
Sg
Pittosporum radicans R. Cunn. ex A. Cunn. loc. cit. 1839. (T.: R. Cunningbam 200,
P meet sepia R. Cunn. ex A. Cunn. var. reflexum (A. Cunn.) Hook. f. Fl.
v. Zel. 853.
P Baton pe ese Kirk, in N.Z. Inst. Trans. & Proc. 1: 143. 1868. (T.: Kirk s.n.,
aT).
tee arios ear bd R. Cunn. ex A. Cunn. ssp. reflexum (A. Cunn.) Kirk, in N.Z.
ns. & Proc. 4: 265. 187 : f :
P itosporum pra vides R. Cunn. ex A. Cunn. var. gilliesianum (Kirk) Kirk, loc. cit. 264.
Occasional from Kaitaia to Kawakawa in North Auckland, New Zealand.
Flowers SC March to e
Kaitaia, H. B. Mattbews s.n. (AK); in Kauri
D:
ée attbews t$ Carse s.n. (AK, MO );
New ZEALAND TH A
t ; , R. H. M
ea j^ Y fore Fairburn, Man sar eg southwest of “Tape, 100’ above sea-leve ei in
remnant of pis trai forest, Cor 36009 (AK, MASS, MO); same locality,
[Vol. 43
152 ANNALS OF THE MISSOURI BOTANICAL GARDEN
/ l /
P. PIMELIOIDES ssp. MAJOR
P. PIMELIOIDES
ssp. PIMELIOIDES
TE DL
LAST LONGITUDE
Fig. 19. P. pimeleoides ssp. major
and ssp. pimeleoides. Names on map
should read pimeleoides.
Powell 26437 (AK); hills north of Mangonui Harbour, Cheeseman s.n. (AK, BM); same
locality, Kirk s.n. (US), 515 (BM); a "pe near Mangonui, Cheeseman s.n
near the head of the EO Kawa, R. Be ed 200, 619 (K, as P. SCH WITHO
LOCALITY: Buchanan ign ¿ein ei (K); A. Cunningham 618 (K); Edgerley ey
s.n. (K); "Auckland", Kirk (K); a Nd D woods, Hooker 352 (K); Wilkes
s.n. (US); ora Ecce is Hai 25/2, 26 (R. Cunningbam) (W).
Flowers which appear to be female have capitate obscurely 2-lobed stigmas,
styles only about 1.5 mm. long, and plumper ovaries 2 mm. long by 1 mm. broad.
The pistil is exserted 1 mm. or more beyond the stamens, which have sagittiform
probably sterile anthers only 0.5 mm. long. Flowers which appear to be male
have weakly capitate stigmas, styles 3.0 - 3.5 mm. long, and thinner ovaries 1.0 -
1.5 mm. long and 0.5 - 1.0 mm. broad. The pistil is level with the anthers, which
are ovoid, apparently functional, and 1.0 - 1.5 mm. long. The pedicels of “fe-
male” flowers are markedly shorter than those of “male flowers”.
The ranges of P. pimeleoides and P. reflexum are not distinct, and I cannot
find any constant morphological characters to separate P. reflexum as a species Or
variety. I suspect that most of the variation in leaf size and shape is caused by
ecological factors, but have seen too few living plants to be certain. In the dis-
cussion of P. eugenioides, later in this work, the priority of the specific names of
A. Cunningham (Ann. Nat. Hist. 4: 108. 1839) over those of Putterlick (Syn.
Pittosp. p. 15. 1839) is discussed.
14b. Ze PIMELEOIDES ssp. major (Cheeseman) R. C. Cooper, stat.
Pitostoram bimeleoides R. Cunn. ex A. Cunn. var. major Cheeseman, Man. N.Z. Fl. p.60.
(T.: Cheeseman s.n. !).
1956]
COOPER—AUSTRALIAN AND NEW ZEALAND PITTOSPORUM 153
ay only on Kerr Point, North Cape, New Zealand. Flowers in June.
w ZEA : NO ree Kerr Point, North Cape, Cheeseman s.n. (AK);
same DIAM Micbie 80/45 (AK
Kerr Point was Heres an island but now is joined to the North Island of
New Zealand by a sand bar some 70 miles long, built probably during the Castle-
clifian (upper Pliocene) period. Several species of plants are endemic to Kerr
Point, but I hesitate to recognize ssp. major as a distinct species as it is very similar
to ssp. pimeleoides in the characters of the flowers and fruits.
15. PrrrosPoRUM PATULUM Hook. f. Handb. N.Z. Fl. p. 19. 1864 (CT: Sin-
clair s.n. !).
Shrubs 1 - 5.0 m. tall; branchlets grayish-brown, the young parts brown-
tomentose, soon glabrous. Leaves alternate, 2 - 6 cm. long, 3 - 10 mm. broad;
juvenile leaves usually longer and narrower than the adult, linear, deeply lobed to
pinnatifid, the lobes variously toothed; adult leaves linear-oblanceolate, entire to
crenate-serrate, obtuse to acute at apex, attenuate at base, dark green above, paler
beneath, glabrous, coriaceous, margins flat, rarely revolute, costa raised above and
below, side veins 10 - 15 per side, anastomosing, usually obscure when juvenile
but distinct above when adult; petioles 2 - 7 mm. long, brown-tomentose when
young, glabrate. Flowers terminal, 3 - 7, fascicled; pedicels 7 - 15 mm. long,
contracted in fruit, brown-tomentose, subtended by a whorl of leaves and numer-
ous caducous, sparsely tomentose bud scales 1 - 2 mm. long. Sepals rarely imbri-
Cate at base, ovate-lanceolate, irregularly toothed above, acuminate, 4 - 6 mm.
long, 1.5 - 2.5 mm. broad, glabrous; petals oblong, obtuse, 9 - 11 mm. long, 2 -
3 mm. broad, free or coherent at the base, spreading from above the middle, purple;
stamens 4 - 7 mm. long, anthers about 1 mm. long and broad. Pistil at anthesis
slightly exserted; ovary 2.5 - 3.0 mm. long, 1 - 2 mm. broad, glabrous; style 1.5 -
3.5 mm. long; stigma capitate and 2-lobed or truncate. Capsules cordate, globose
to subglobose, 2-valved, 6 - 11 mm. in diameter, glabrous; valves convex to weakly
sulcate in transverse section, less than 1 mm. thick, coriaceous, with a conspicuous
placenta bearing 2 rows of short stout funicles from the base to just above the
middle; seeds about 18 per capsule, black, irregular.
LLL
TTS
Fig. 20. P. patulum. Fig. 21.
[Vol. 43
154 ANNALS OF THE MISSOURI BOTANICAL GARDEN
Rare and local in the northern ranges of the South Island of New Zealand.
ZEALAND: NORTH AUCKLAND: Kerr Point, North Cape, Cheeseman s.n. (AK);
near Collingwood, Gibbs s.n. (BM); Lake Rotoiti, Buchanan s.m. (AK); same locality,
Cheeseman s.n. (AK); Spencer Mountains, Kirk sm. (GH MO); Spencer Mountains,
Glacier Gully, Laing s.n. (NSW); woods, Upper Waiau, Travers s.n. (K); 5000’, same
locality, Sinclair s.n. (
16. PrrrosPORUM TURNERI Petrie, in N.Z. Inst. Trans, & Proc. 55: 95. 1924.
(T.: Matthews & Carse s.n. !).
Shrubs or small trees 2 - 9 m. tall; branchlets dark gray or brown, sharply
divaricating and matted at juvenile stage or on lower part of stem, fastigiate in
adult stage or on upper part of stem, white-tomentulose when young, soon glabrate.
Leaves alternate, clustered at the tips of short branchlets, brownish-green above,
paler beneath, tomentulose when young, soon glabrate, coriaceous, with slightly
thickened and revolute margins, costa obscure above, raised beneath, secondary
veins 10 - 13 per side, anastomosing, obscure above, distinct beneath. Juvenile
leáves orbicular, obovate, or linear, entire or variously lobed and parted, 2 - 15 mm.
long, 1 - 5 mm. broad, adult leaves obovate to oblanceolate, obtuse to acute at
apex, attenuate at base, entire or obscurely crenate, 1 - 4 cm. long, 6- 12 mm.
broad; petioles 0.5 - 2.5 mm. long, 0.5 - 1.5 mm. broad, tomentulose when young,
soon glabrate. Flowers terminal, 4 - 12, fascicled; pedicels 1 - 3 mm. long, ac-
crescent in fruit, brown-tomentose, subtended by an approximate whorl of leaves
and numerous caducous tomentose-ciliate bud scales 2 - 7 mm. long. Sepals
slightly imbricate at base, lanceolate, acute to acuminate, 3.5 - 5.0 mm. long, ! -
2 mm. broad, ciliolate, tomentulose; petals oblong, subacute to obtuse, 6 - 9 mm.
long, 2.0 - 2.5 mm. broad, free spreading from above the middle, pink or purple;
stamens 4.0 - 5.5 mm. long, anthers ovate-elliptic, 1.5 - 2.0 mm. long, 0.5 - 1.0
mm. broad. Pistil at anthesis slightly shorter than or equal to the stamens; ovary
2.5 - 3.5 mm. long, 1.0 - 1.5 mm. broad, tomentulose; style about 2 mm. long;
stigma weakly capitate to truncate. Capsules globose, 2-, rarely 3-, valved, apicu-
late, 5 - 8 mm. in diameter, slightly rugose; valves convex or sometimes sulcate in
transverse section, less than 1 mm. thick, coriaceous, with a conspicuous placenta
bearing several pairs of short stout funicles between the base and the middle; seeds
3 - 10, black, irregular.
Found only on the Central Volcanic Plateau of the North Island of New
Zealand, at forest margins between Erua and Waimarino, about 1000 m. above sea-
level. Flowers in November and December.
ZEALAND: VOLCANIC PLATEAU: Waimarino, Matthews t$ Carse s.n. (AK,
r
3000’, Erua and Waimarino, Attwood s.n. (AK, K, MO); forest margins, Waimarino,
Allan s.n. (GH, K); on stream side near Erua, Allan s.n. (K); same locality, Moore &
MO)
1956]
COOPER——AUSTRALIAN AND NEW ZEALAND PIT CG 133
17. PrrrosPoRUM OBCORDATUM Raoul, in Ann. Sci, Nat. III, 2: 121. 1844.
(T.: Raoul s.n. !).
Shrubs to small trees 2 - 5 m. tall; branchlets divaricating, gray to reddish-
brown, tomentulose when young, soon glabrate. Leaves alternate at seedling stage
and on young branchlets, later restricted to the tips of arrested branchlets 1 - 2
mm. long, tomentulose to glabrous, submembranous when young, coriaceous when
adult, margins entire or crenate, flat or revolute, sparsely ciliolate, costa immersed
above, raised beneath, secondary veins 2 - 4 per side, anastomosing, obscure above,
distinct or obscure beneath; juvenile leaves at first oblong to elliptic, entire, 5 - 9
mm. long, 2.5 - 4.0 mm. broad, soon linear to spathulate, variously lobed, toothed
and parted, rarely entire, 1.3 - 3.6 cm. long, 2 - 12 mm. broad; adult leaves or-
bicular to obovate, obcordate to obtuse at apex, attenuate at base, 4 - 14 mm. long
and broad; petioles 0.5 - 5.0 mm. long, tomentulose to glabrous, usually narrowly
winged. Inflorescences axillary or terminal, on minute arrested branchlets, 1- to
5-flowered, umbelliform; pedicels up to 2 mm. long, accrescent in fruit, tomentu-
lose, subtended by 1 - 5 leaves and numerous caducous sparsely ciliolate and tomen-
tose bracts 1-2 mm. long. Sepals slightly imbricate at base, lanceolate-subulate
acute, 1.5 - 3.0 mm. long, 0.5 - 1.0 mm. broad, ciliolate, tomentulose; peta
linear-oblong, obtuse, 4.0 - 6.5 mm. long, 0.7 - 1.5 mm. broad, connate in a
Cylindrical tube with spreading or reflexed tips, pale purple, yellow, or white,
sometimes with a reddish-purple stripe; stamens 2.5 - 4.5 mm. long, anthers sagitti-
orm to ovate, 0.5 - 1.0 mm. long. Pistil at anthesis slightly shorter or longer
than the stamens; ovary 1.5 - 3.3 mm. long, 0.5 - 1.5 mm. broad, tomentose;
style 1 - 2 mm. long; stigma capitate and obscurely 2-lobed or truncate. Capsules
ellipsoid to subovoid, 2-valved, apiculate, with persistent sepals at base, 6.5 - 10.0
ong, 5 - 7 mm. broad, frequently dehiscing laterally by one opening, green
to black, slightly rugose, sparsely tomentose, glabrate; valves convex in transverse
section, less than 1 mm. thick, coriaceous, with a slightly thickened placenta
bearing 2 - 4 stout flattened funicles up to 1 mm. long from the base to just above
the middle; seeds 4 - 7, reddish-black to black, irregular.
Found in three isolated localities in New Zealand. Flowers in October and
November,
New ZEALAND: NORTH AUCKLAND: near Lake Tongonge, Kaitaia, H. B. Matthews
s.n. (AK, CANTY); same locality, R. H. Matthews s.n. (AK, BM); in open on river
ank, west of wireless station, Kaitaia, H. B. Matthews & Carse s.n. (AK, CANTY); in
alluvial. land subject to flooding, in shade = river bank below Kaitaia, R. H. Matthews
9 Carse s.n, (AK, MO); Kaitaia, H. B. Matthews s.n. (AK, MO); 1855 (K). EAST
CAPE: Wairoa River, Sainsbury s.n. (AK, pos TY); Hurumua, r, Wairoa, Hodgson
t», (CANTY). CANTERBURY; presqu'ile de Banks, Raoul s.n. (P), 8
The type material was collected by Raoul between 1840 and 1842 at Banks
Peninsula in the South Island of New Zealand, but the plant has not been found
again in that locality. In 1901 R. H. Matthews found two or three plants at
Kaitaia in the extreme north of the North Island, and about 1923 G. O. K. Sains-
bury discovered twelve or fifteen plants near Wairoa on the east coast of the North
[Vol. 43
156 ANNALS OF THE MISSOURI BOTANICAL GARDEN
Island. The plants of these widely separated stands are not floristically d
and Laing & Gourla!?! described the Kaitaia population as P. obcordatum v
taiaensis but their description lacks a Latin diagnosis and is therefore invalidi
The two authors mentioned separated the Kaitaia plants as a variety on their seed-
ling form, but I do not consider that their evidence is adequate. Cheeseman’™
noted that specimens from Kaitaia had slightly larger young leaves, and Sainsbury
pointed out that the bark of the Wairoa plants is reddish-gray rather than gray,
but the differences in the material which I have seen are so slight that I do not
consider that varietal names are warranted.
The flowers of Kaitaia and Wairoa plants appear to be unisexual. Flowers
which may be female have capitate, 2-lobed stigmas, short styles, plump ovaries 3
mm, long, 1.0 - 1.5 mm. broad, stamens 0.5 - 1.3 mm. shorter than the pistil and
sagittiform probably abortive anthers under 1 mm. long. Flowers which may be
male have weakly capitate to truncate stigmas, longer styles, more slender ovaries
1.5 - 2.5 mm. long, 0.5 - 1.0 mm. broad, stamens almost equal to or slightly longer
than the pistil, and ovate probably functional anthers about 1 mm. long. The
single Akaroa specimen has flowers which appear to be male. Godley!9* has in-
formed me that the species is dioecious.
no LAST LONGITUDE 10 WEST LONGITUDE
Fig. 22. P. obcordatum. Fig. 23. P. rigidum.
18. PrrrosPoRuM RiGmuM Hook. f. Fl. Nov. Zel. 1: 22. £. IO. 1853. (T::
Colenso s.n. !).
Shrubs to 3 m. tall; branchlets erect, not interlaced or divaricating, grayish-
brown, rusty- to gray-tomentose, glabrate. Leaves alternate, dark green above,
paler beneath, costa raised or immersed above, raised beneath, secondary veins
usually obscure; juvenile leaves obovate, oblanceolate, or elliptic-oblong, acute,
6l Laing & oe in Trans. R. p Me m 65: 47. 1935.
162 Int, Code Bot. Nomencl. 1952.
163 Man, N.Z. Fl. pri r» vn
165 Personal communica
1956
COOPER—AUSTRALIAN AND NEW ZEALAND PITTOSPORUM 157
rarely obtuse at apex, attenuate at base, variously incised, toothed or crenate, occa-
sionally entire, 5 - 13 mm. long, 3- 10 mm. broad, tomentulose, soon glabrate,
membranous to coriaceous; adult leaves elliptic-oblong, occasionally oblanceolate
“to obovate, rarely lanceolate, obtuse to subacute at apex, obtuse at base, entire,
rarely toothed, 5 - 24 mm. long, 3-13 mm. broad, rusty-tomentose, glabrate,
coriaceous, slightly revolute; petioles 1-3 mm. long, rusty-tomentose. Flowers
terminal or axillary, solitary; pedicels 0.5 - 4.0 mm. long, accrescent in fruit, sub-
tended by one to several leaves and a whorl of caducous brown-tomentulose bud
scales 1 - 2 mm. long. Sepals not imbricate, lanceolate-oblong, acute, 3.5 - 6.0
mm. long, 1.0 - 1.5 mm. broad, sparsely ciliolate with scattered hairs; petals linear-
oblong, obtuse, 8 - 12 mm. long, 2 mm. broad, free, patent with reflexed tips,
dingy purple; stamens 4.0 - 6.5 mm. long, anthers sagittiform or ovate, 0.6 - 1.8
mm. long, 0.3 - 1.0 mm. broad. Pistil at anthesis slightly shorter than the stamens;
ovary 2.0 - 2.5 mm. long, 1 - 2 mm. broad, tomentose; style 2.5 mm. long; stigma
Capitate to truncate. Capsules subglobose, 2-valved, apiculate, 8 - 10 mm. long,
6 - 10 mm. broad, tomentose, glabrate, rugose; valves convex in transverse section,
less than 1 mm. thick, coriaceous, with a slightly thickened placenta bearing 1 - 5
thick strap-like funicles up to 1.5 mm. long near the middle; seeds 3 - 9, black,
irregular.
Occurs from the East Cape of the North Island of New Zealand to Marl-
borough in the South Island, along the mountain ranges between 650 and 1400 m.
Flowers in November and December.
EW ZEALAND: EAST CAPE: Mt. Hikurangi, Adams s.n. (AK); 4000’, same local-
ity, Petrie s.n. (AK); 4300’, undershrub in Nothofagus (beech) forest, ors cy gn
cl Moore t$ Cranwell s.n. (AK). WELLINGTON: Mt. Holdsworth, 3500’, Town
- (BM); same locality, at forest line, Cockayne 9076 (A); same locality, Aston 3.0.
(AK, NSW); same locality, Jones s.n. (MO); 3000'—4000', Mt. Hector, Petrie s.n.
(AK); same locality, Attwood s.n. (AK); Mt. A sepe, Attwood s.n. (AK); Mt.
Omega, Turner 156 (AK); Field je Moore & Cranwell s.n. (AK). MARLBOROUGH:
s Stok urner s.n. ( ; our LOCALITY: Colenso s.n., 45, 64, 924 (K);
T. L. Travers s.n. (K)
The flowers are of two kinds: those which appear to be male have truncate
stigmas, styles about 2.5 mm. long, slender ovaries 2.5 mm. long, 1 mm. broad,
stamens 5.5 - 6.5 mm. long, and probably functional ovoid anthers 1.5 - 2.0 mm.
long, 1 mm. broad. The flowers which appear to be female have capitate stigmas,
styles about 2 mm. long, plump ovaries 2.0 - 2.5 mm. long, 2 mm. broad, stamens
about 4 mm. long, and probably abortive sagittiform anthers 0.6 - 0.8 mm. long,
0.5 - 0.5 mm. broad.
The flowering material available is inadequate but in the specimens seen the
male flowers had pedicels up to 4 mm. long while the female flowers had pedicels
only 0.5 mm. long. Godley** has informed me that the species is dioecious.
Cockayne!®* divided P. rigidum Hook. f. into two species, and kept the name
165 Personal c
166 N.Z. Inst. Tow k k bui: 47:111. 1915.
[Vol. 43
158 ANNALS OF THE MISSOUR! BOTANICAL GARDEN
P. rigidum for the East Cape and Tararua population, as Colenso gathered the type
material in the East Cape district.
19. PrrTOSPORUM CRASSICAULE Cockayne ex Laing & Gourlay, in Trans. Roy.
Soc. N.Z. 65: 50. 1935. (T.: Laing & Gourlay s.n. !). T
Pittosporum lineare Laing & Gourlay, loc. cit. 57. 1935. in part. (quoad MacMahon 351).
Shrubs to 4 m. tall; main branches usually erect, often closely compressed;
branchlets divaricating, opposite or whorled at the nodes, stout, often spinose at
the tips, white-tomentose. Leaves alternate at seedling and juvenile stage, re-
stricted to the apex of short, stout, sometimes minute branchlets at adult stage,
submembranous when juvenile, coriaceous when adult, green above, paler beneath,
sparsely ciliolate and tomentulose when young, glabrate, costa slightly raised above
and beneath or immersed, secondary veins obscure; juvenile leaves obovate to ob-
lanceolate, variously lobed and parted, lobes irregularly dentate, 4.5 - 12.0 mm.
long, 3 - 8 mm. broad; adult leaves linear or elliptic-oblong to oblanceolate, obtuse
to subacute at apex, attenuate at base, entire, occasionally lobed or toothed, espe-
cially in shade forms, 3.5 - 17.0 mm. long, 1 - 4 mm. broad; petioles 1.0 - 1.5 mm.
long, glabrous. Flowers terminal, 1 - 3, solitary or fascicled, sessile on minute
arrested branchlets, subtended by 1 - 6 leaves and numerous persistent tomentulose
bud scales about 1 mm. long. Sepals imbricate at base, lanceolate-ovate, 1.2 - 1.5
mm. long, 1 mm. broad, glabrous, ciliolate ; petals linear-oblong, obtuse, 4.0 -
7.5 mm. long, 1.2 - 1.5 mm. broad, coherent in a cylindrical tube, tips spreading,
later recurved, purple; stamens 1.5 - 4.5 mm. long, anthers ovate, 0.6 - 1.0 mm.
long, 0.4 - 0.6 mm. broad. Pistil at anthesis slightly shorter or longer than the
stamens; ovary 1.5 - 2.5 mm. long, 1 - 2 mm. broad, tomentulose; style 1 - 2 mm.
long; stigma capitate or truncate. Capsules subglobose to globose, 2-valved,
apiculate, 4.5 - 7.0 mm. long, 5.5 - 7.0 mm. broad, tomentulose or glabrate, rugose;
valves convex in transverse section, less than 1 mm. t ick, almost woody, with a
placenta thickened from the middle to the base, bearing 1 - 2 slender peg-like
funicles near the middle; seeds 1 - 3, black, round to irregular.
Occurs in Marlborough, Nelson, Westland, and Canterbury, mainly on the
western side of the South Island of New Zealand. Flowers in October and
November.
New ND: MARLBOROUGH: Pelorus Valley, MacMabon 35 (AK). NELSON:
500’, Maitai Valley, Cheeseman s.n. (AK); Rocky River, Bainham, Collingwood, Wall s.n.
(CANTY); Buller Valley, Townson 445 (AK); same locality, Cheeseman s.n. (AK, MO).
WESTLAND: 380 m. in scrub on banks of Teremakau River, near Railway Settlement,
i . CANTERBURY: 730 m. north side of Rough
Creek, near railway cottages, Arthur's Pass, Laing & Gourlay s.n. (CANTY); same local-
ity, Laing s.n. (K); same locality, near Jack's Hut, Laing s.n. (K); roadside, same locality,
Cooper 24318 (AK, US).
Flowers which may be female have capitate stigmas, and stamens 1.5 - 1.6
mm. long, with anthers 0.6 mm. long and 0.4 mm. broad. Flowers which may be
male have truncate stigmas, and stamens 3.9 - 4.6 mm. long with anthers 1 mm.
1956]
COOPER——AUSTRALIAN AND NEW ZEALAND PITTOSPORUM 159
long and about 0.5 mm. broad. I cannot find any consistent differences in the
size of the ovaries.
NI Pac uo Hol
T PII.
no EAST LONGITUDE 180 WEST LONGITUDE LAST LONGITUDE WEST LONGITUDE
Fig. 24. P. crassicaule. Fig. 25. P. divaricatum.
20. Prrrosporum DIVARICATUM Cockayne, in N.Z. Inst. Trans, & Proc. 47: 111.
1915. (T.: Cockayne 8551 !).
Pittosporum divaricatum Cockayne, in N.Z. Inst. Trans. & Proc. 44: 20. t. 3. f. 2 @ t. 8.
1912, nom nud; in Austral. Assoc. Adv. Sci. Proc. 13: 219. 1912, nom. nud.
Pittosporum lineare Laing & Gourlay, in Trans. Roy. Soc. N.Z. 65: 57. 1935, in part.
(quoad Laing & Gourlay s.n. 1).
Shrubs to e m. tall; branches thick and woody, interlaced; branchlets divari-
cating, opposite or whorled at the nodes, stout, often spinose at the tips, tomentu-
lose when young, soon glabrate. Leaves alternate at seedling stage and on young
branchlets, later restricted to the tips of arrested branchlets 1 - 5 mm. long, sub-
membranous when juvenile, coriaceous when adult, green above, paler beneath,
glabrous, costa immersed above, sunken beneath, secondary veins obscure; juvenile
leaves oblong, obovate to lanceolate, or almost linear, margins with 1 to several
lobes or teeth on'either side, occasionally crenate, 6 - 9 mm. long, 1 - 4 mm. broad;
adult leaves linear-oblanceolate to linear-oblong, or ovate, acute to obtuse at apex,
acute at base, 4 - 10 mm. long, 2 - 7 mm. broad, smaller and entire in sun, larger
and variously dentate, crenate, or lobed in shade; petioles about 1 mm. long,
Blabrous. Flowers terminal on short arrested branchlets, solitary, sessile or with
minute pedicels, subtended by several leaves and caducous ciliolate bud scales about
1 mm. long. Sepals not imbricate at base, lanceolate, acute, 1.5 - 2.5 mm. long,
0.5 - 1.0 mm. broad, glabrous, ciliolate; petals linear-oblong, obtuse, 4 - 6 mm.
long, 1.0 - 1.5 mm. broad, coherent in a cylindrical tube, tips spreading, purple;
stamens 1.5 - 2.6 mm. long, anthers ovate or sagittiform, 0.5 - 1.4 mm. long,
0.3-0.7 mm. broad. Pistil at anthesis slightly exserted; ovary 1.0- 2.2 mm.
long, 0.5 - 1.0 mm broad, glabrous; style 1.0- 1.5 mm. long; stigma capitate to
truncate. Capsules cordate or subglobose, 2-valved, acute and apiculate, 6 mm.
long, 5.5 - 6.0 mm. broad, glabrous, weakly rugose; valves convex in transverse
[Vol. 43
160 ANNALS OF THE MISSOURI BOTANICAL GARDEN
section, less than 1 mm. thick, coriaceous, with a slightly thickened placenta bear-
ing 1 - 2 pairs of peg-like funicles near the middle; seeds 1 - 6, black, round to
irregular.
Occurs from the Ruahine Range and Central Volcanic Plateau of the North
Island to Arthur's Pass on the Main Divide of the Southern Alps of New Zealand.
Flowers in October and vae e?
EW ZEALAND: EAST CAPE: Pukatitiri, eastern l| of Ruahine Range, Hodg-
son sm. (AK, CA ae VOLCANIC : arioi, vicinity of karan
Attwood s.n. (AK, MO). WELLINGTON: Tararua Yi “ig Petrie s.n. (AK).
BOROUGH: Picton, MacMahon s.n. (AK, NSW); Koromiko, Laing & Gourlay [The > label
of the Kew sheet of the Koromiko plant is noted by H. H. Allan as “col. H. Jenkins end
of Oct. 1932."] s.n. (CANTY, K). NELSON: De s ale Sainsbury s.n. (CANTY);
angapeka River, Cheeseman s.n. (AK); 2000’, in bog forest on >- — E
Cockayne 8551 A, 3 without locality, Bidwill 96 (K). CANT
Morrison 39 (A, CANTY); Cass River, Kirk s.n. (GH, NSW); SSC dde SC "oni
Creek, Arthur's E Laing & Gourlay s.n. (CANTY).
It is with hesitation that I treat P. lineare as a synonym of P. divaricatum and
P. crassicaule. Laing and Gourlay, the two authors of P. lineare, cultivated and
studied the small-leaved New Zealand species of Pittosporum for many years before
publishing their results, but in the material cited by them there appear to be two
distinct entities: the Koromiko plants (Laing & Gourlay s.n.) which have flowers
with sepals 2.0-2.5 mm. long, capitate stigmas, styles 1.0- 1.2 mm. long,
glabrous ovaries 2.0 - 2.2 mm. long, and stamens about 2 mm, long; and the
Pelorus Valley plants (MacMahon s.n.) which have flowers with sepals 1.0 - 1.5
mm, long, truncate stigmas, styles about 1.8 mm. long, tomentulose ovaries 1.5
mm. long, and stamens nearly 4 mm. long. The linear entire leaves of the Koro-
miko plant give it a very different appearance from most specimens of P. divari-
catum, but the flowers are within the range of variation of P. divaricatum, and
similar plants with linear entire leaves and bearing the distinctive cordate glabrate
fruits of P. divaricatum occur on the Volcanic Plateau (Attwood s.n.), and in
cultivation (Cooper 36299). The tomentulose ovary of the Pelorus Valley plants
is a feature of P. crassicaule, P. rigidum, and P. obcordatum, but the other char-
acters of the flower, fruit, and foliage are similar to those of P. crassicaule. Con-
sequently with some misgivings I have included the collection in that species.
In the material examined there are slight differences in the size and form of the
flowers. Flowers which may be female have capitate stigmas, ovaries 1.5 - 2-2
mm. long, 1 mm. broad, stamens 1.5 - 2.5 mm. long and sagittiform to ovate
anthers 0.5 - 0.9 mm. long, 0.3- 0.5 mm. broad. Flowers which may be male
have truncate stigmas, ovaries 1.0 - 1.5 mm. long, 0.5 - 0.6 mm. broad, stamens
about 4.0 mm. long, and ovate anthers 1.0 - 1.5 mm. long, 0.6 - 0.7 mm. broad.
21. PrrrosPORUM ANOMALUM Laing & Gourlay, in Trans. Roy. Soc. N.Z. 65:
54. 1935. (T.: Laing & Gourlay s.n. !).
Shrubs, prostrate in the sun and with rigid, almost spinose branches, semi-
prostrate in shade and forming subglobose masses up to 1 m. tall and 2 m. in
1956]
COOPER—AUSTRALIAN AND NEW ZEALAND PITTOSPORUM 161
diameter with non-spinose, interlaced branches; branchlets brownish-gray, to-
mentulose when young, soon glabrate. Leaves alternate on juvenile plants and
young branchlets, later restricted to the tips of arrested branchlets, oblong-linear
to oblanceolate-linear, obtuse to subacute at apex and base, submembranous when
young, coriaceous when adult, green in summer, brownish-red in winter, paler
beneath, sparsely tomentulose when young, soon glabrate, costa immersed above,
raised beneath, secondary veins obscure; juvenile leaves dentate-serrate, 8 - 12 mm.
long, 2 - 3 mm. broad; adult shade leaves deeply doubly dentate or serrate to
crenate, occasionally entire, 5 - 10 mm. long, 1.0 - 2.5 mm. broad; adult sun
leaves entire, occasionally crenate, 2 - 5 mm. long, 1.0 - 1.5 mm. broad; petioles
up to. 1 mm. long, glabrous. Flowers terminal on short arrested branchlets,
solitary, sessile, subtended by several leaves and caducous ciliolate bud scales 2 mm.
long. Sepals not imbricate at base, lanceolate, 2 mm. long, 0.5 - 1 mm. broad,
glabrous, ciliolate; petals obovate to oblanceolate, obtuse to subacute, 2.5 - 4.0 mm.
long, 1.2 - 2.0 mm. broad, spreading, creamy yellow, purple at the edges and tips;
stamens 2 - 3 mm. long, anthers ovate to orbiculate, 0.5 - 0.8 mm. long, 0.3 - 0.8
mm. broad. Pistil at anthesis equal to the stamens; ovary 1.0- 1.3 mm. long,
0.8 - 0.9 mm. broad, glabrous; style 0.5 - 12.2 mm. long; stigma capitate and 2-
lobed or truncate. Capsules ovoid, 2-valved, acute to acuminate, apiculate, 5.5 -
7.0 mm. long, 3 - 4 mm. broad, glabrous; valves convex in transverse section, less
than 1 mm. thick, subcoriaceous, splitting longitudinally on dehiscence and leaving
a papery endocarp covering the seeds; placenta unthickened, bearing 1-2 short
stout funicles about the middle; seeds 2 - 4, black, round to irregular.
Occurs on the Central Volcanic Plateau of the North Island, and at Nelson
and Arthur's Pass in the South Island of New Zealand. Flowers in December and
January.
New ZEALAND: VOLCANIC PLATEAU: shrub steppe, Mt. Hauhangatahi, Carse &
Matthews s.n. (CANTY); Hauhangatahi, Moore & Cranwell s.n. (AK, MO); 3000’, in
tussock near Chateau Tongariro, Mt. Ruapehu, Godley 24963 (AK). NELSON: woode
peak, Gibbs s.n. (CANTY). CANTERBURY: 3000’, Arthur's Pass, Laing & Gourlay s.n.
(CANTY); same locality, "Jack's Hut”, Laing s.n. (K).
Flowers which may be female have capitate 2-lobed stigmas, styles 0.5 - 0.7
rm LJ
Fig. 26. P. anomalum. Fig. 27. P. cornifolium.
[Vol. 43
162 ANNALS OF THE MISSOURI BOTANICAL GARDEN
mm. long, and ovate anthers 0.5 mm. long, 0.3 mm. broad, on filaments about
1.5 mm. long. Flowers which may be male have truncate stigmas, styles 1.0 - 1.2
mm. long, and orbiculate anthers 0.7 - 0.8 mm. long, 0.5 - 0.8 mm. broad, on
filaments 1.5 - 2.2 mm. long. There is little difference in ovary size in the speci-
mens which I have seen
22. PrrrOsPORUM CORNIFOLIUM A. Cunn. ex Hook. in Bot. Mag. f£. 3101.
1832. (T.: A. Cunningham 616 !).
Pittosporoides verticillata Banks & Soland. ex A. Cunn. in Ann. Nat. Hist. 4: 107. 1839,
nom. n in
Shrubs 0.5 -2.5 m. tall, epiphytic, rarely terrestrial; branchlets forked or
verticillate, dark brown, glabrous. Leaves verticillate, obovate- to ovate-elliptic, -
acute to subacuminate at apex, acute to obtuse at base, entire, 2 - 10 cm. long,
1-5 cm. broad, light green above, paler beneath, ciliate when young, soon gla-
brate, coriaceous with thickened slightly revolute margins, costa raised above,
immersed beneath, secondary nerves obscure above, distinct beneath; petioles
0.5 - 3.0 mm. long and 0.5 - 2.0 mm. broad, glabrous. Inflorescences terminal, 1-
to 10-flowered, usually umbelliform; pedicels 2 - 15 mm. long, accrescent in fruit,
brown-tomentose, subtended by a whorl of leaves and numerous caducous bud
scales about 1 mm. long. Sepals not imbricate, narrow-lanceolate, acute, 4-7
mm. long, about 1 mm. broad, spreading, sparsely puberulent; petals linear-
lanceolate, acute to acuminate, 8- 12 mm. long, 1.5 - 2.0 mm. broad, coherent
in a tube with reflexed tips, yellow; stamens 4 - 6 mm. long, anthers sagittiform
to elliptic-oblong, 1 - 2 mm. long, 0.5 - 1.0 mm. broad. Pistil at anthesis slightly
longer to slightly shorter than the stamens; ovary 1.5 - 3.0 mm. long, 0.5 - 2.0
mm. broad, villous; style 2.5 - 4.0 mm. long; stigma capitate and 2-lobed to
truncate Capsules ellipsoid, 2- or 3-valved, about 1.7 cm. long, 1.2 cm. broad,
glabrate; valves convex in transverse section, less than 1 mm. thick, coriaceous,
with a short basal placenta bearing several strap-like funicles up to 5 mm. long;
seeds 4 - 8, black, irregular.
Occurs from sea-level to 650 m. throughout the North Island and northern
part of the South Island of New Zealand; common in the northern part of the
North Island, rarer to the south. Flowers from June to September.
x ALAND: NORTH AUCKLAND: Kaitaia, Matthews s.n. (AK, MO); Okahu,
Kaitaia, Matthews s.n. (AK); Whangaroa, Kirk s.n. (GH); Bay of Islands, Wilkes s.n.
(AK); Poor Knights Islands, Cranwell s.n. (AK). AUCKLAND: Te Pahi, Kirk s.n. (A);
Omaha, Kirk s.». (AK, F, US); Leigh, Buddle 27204 (AK); same locality, Turner s.n.
: K ); same locality, Petrie s.n. (CU); Birk-
4 anges, Cheeseman s.n. (BISH, NSW, US);
same locality, Petrie s.n. (NSW); epiphytic on Metrosideros tomentosa, Huia, Wood s.n.
1956]
COOPER——AUSTRALIAN AND NEW ZEALAND PIT UM 163
(AK); Titirangi, Cheeseman s.n. (GH, NSW, US), 11 (K); Auckland, Cranwell s.n.
(AK); Buckland, Matthews s.n. (AK, MO.) rHAMESs: Cape Colville, Kirk s.n. (MO);
Coromandel Peninsula, Adams s.n. (AK); Shag Bay, Te Mad Moore & S s.n.
(AK, K); Pakirarahi, Adams s.n. (AK); Tararu, Kirk sn. (AK); Thames, Adams s.n.
(US); in sylvis prope Tolaga [East Cape], Opuragi DT Bay], Banks & agio
BM
sn. (BM). EAST CAPE: Lake Waikaremoana, Sainsbury 15 (AK); Dra rois
Turner 162 (AK). WANGANUI: Parapara, Mangowhero, Waimarino, Attwood s. K);
Turakina, Allison s.n. (A, NSW). WELLINGTON: e iphytic on Leger Ped
orest margin, Fielding, Allam s.». (BH); Mungaroa, Kirk 136 (BM); Ofungekoago;
H d
Kirk 130 (GH). MARLBOROUGH: terrestrial, Endeavour Inlet, Queen Charlotte Sound,
MacMabon s.n. (AK); Tennyson va erm een Dec As (A Kier
OCALITY: B K, M
Tileresceices Ed e to 10 flowers appear to be male. The flowers have
truncate to weakly capitate stigmas level with or shorter than the anthers, styles
3.5 - 4.0 mm. long, and ovaries 1.5 - 2.0 mm. long. I think that they are male
as the ovaries are very thin, apparently abortive, and the anthers are elliptic-oblong,
apparently functional. Inflorescences with only 1 or 2 flowers appear to be female.
The flowers have capitate 2-lobed stigmas level with or exserted beyond the anthers,
styles 2.0 - 2.5 mm. long and ovaries 3 - 4 mm. long. I think that they are
female as the ovaries are plump, apparently functional, and the anthers are sagitti-
form and abortive. Kirk!" and Petrie!99 described the flowers of P. cornifolium
as unisexual, and Cheeseman! regarded the plants as polygamous or dioecious.
Field studies are required to determine the significance of the morphological
differences.
The species was described by W. J. Hooker from material grown at the Royal
Botanical Gardens, Kew, and from Allan Cunningham's specimens and notes made
by him in New Zealand in 1826. Two "type" sheets in the herbarium of the
Royal Botanic Gardens, Kew, bear five labels, two sterile specimens, a fruiting
specimen and fragments of flowers. One label is dated 1826, one 1833, two 1838,
and one is undated. As the species was described in 1832 only part of the material
can have been available to W. J. Hooker. Fortunately the species is distinct and
the muddle of labels and specimens does not have serious consequences.
Vernacular name: Cornel-leaved Pittosporum.
23. Prrrosporum KRKI Hook. f. ex Kirk, in N.Z. Inst. Trans. & Proc. 2: 92.
1869. CT: Kirk s.n. !).
Shrubs 1-5 m. tall, epiphytic, occasionally terrestrial; branchlets reddish-
Purple, glabrous. Leaves subverticillate, linear-obovate to oblong-elliptic, acute
or obtuse at the apex, acute at the base, entire, 4 - 11 cm. long, 0.7 -
broad, pale green above, lighter beneath, glabrous, coriaceous, with Me ined and
slightly revolute margins, costa raised above, flattened beneath, secondary veins
d gg Students! Fl. N.Z. p. 49.
68 N.Z. Inst.
Trans. & Proc. 53: ei 1921.
hice Bate ed. 2 p. 495. 1925.
i [Vol. 43
164 ANNALS OF THE MISSOURI BOTANICAL GARDEN
about 18 per side, anastomising, obscure; petioles 3 - 16 mm. long, 1 - 2 mm.
broad, glabrous, purple. Inflorescences terminal, 3- to 10-flowered, umbelliform;
pedicels 5 - 10 mm. long, contracted in fruit, glabrous, subtended by an approxi-
mate whorl of leaves and by numerous caducous glabrous bud scales 1 - 2 mm.
long. Sepals imbricate at base, lanceolate, acuminate, 6 - 10 mm. long, 1.5 - 4.0
mm. broad, glabrous; petals oblanceolate, acuminate, 1.5 - 2.1 cm. long, 2.5 - 3.5
mm. broad, fused slightly above the base to the middle, sharply recurved above,
yellow; stamens 6 - 7 mm. long, anthers elliptic-oblong, 2.5 - 3.0 mm. long.
Pistil at anthesis equalling or slightly longer than the stamens; ovary 3-4 mm.
long, 1.5 mm. broad, tomentulose; style about 3 mm. long, stout, stigma capitate.
Capsules ellipsoid, 2-, rarely 3-, valved, apiculate, 2.6 - 3.9 cm. long, 1.1 - 1.7 cm.
broad, green to yellow, glabrate; valves convex in transverse section, about 1.5
mm. thick, coriaceous, with a placenta bearing slender strap-like funicles up to
4 mm. long, from the base for three-fourths the length of the valve; seeds about
40, black, irregular.
Occasional from 250 to 1000 m. in North Auckland, Auckland, Thames,
including Great Barrier Island, and Wanganui districts of New Zealand.
Flowers from November to January.
New ZEALAND: ORTH AUCKLAND:
2000’, ele Mangonui, Cheeseman
s.n. (AK, K); epiphytic on a dead log, Wara-
wara State Forest, North Hokianga, Cooper
35572 (AK, MO); Hokianga, Petrie s.n. (K);
same locality, Otaua, Berggren s.m. (BM);
epiphytic ded tall tree, Waipoua kauri forest,
coute t 419 (A); same locality, Turner
3 (AK); same locality, Jessup 26640 (AK);
bete. use s.n. (AK); ridge near Pu-
tutu, Takah
ke e side, Petrie s.n (AK,
AUCKLAND Titirangi Ranges, Cheeseman jn
: (AK, GH, K, US), 12 (K); Huia, Mackie
Pech uh E sn. ( Ps ; Huia dam, epiphytic in second-
growth Agathis australis forest, page sn. (AK). AMES: epiphytic, Kaeoruruwahine
forest, Great Barrier Island, aves «(B GH ; MO); "2000, Great Barrier Island, Kirk 63
(K); Cape Colville, Kirk s. (AK), same locality, 2300’, Hector s.n. (BM); same local-
ity, Adams s.n. (AK); Miete and Thames Goldfield, 2500’, Kirk s.n. (A, AK, K), 63
E same locality, Adams s.n. (AK); acer forest, 1500’, ^V ddivcigomet Mine, Te Aroha,
Gemen 1298 (BM); epiphytic, c. 2000', Te Moehau, Moore & Cranwell s.n. (AK,
zy. D ANUI: Maungaturuturu, Manager -a-te-ao, Attwood s.n. (AK)
[epe us name: Kirk's Pittosporum.
24. PrrrosPoRUM REVOLUTUM Aiton, Hort. Kew. ed. 2. 2:27. 1811. (T.: ex
Hort. Kew. !).
Wee sto — i ~ Trans. Linn. Soc. Lond. 10: 298. #. 20. 1811. (TFN
n.!—The type sheet at AEN is marked “1. New Holland. Fleming").
Pittosporum fomentosum Bonpl. Descr. Pl. Rar. Malm. p.56. 4.21. 1813. (Ta "Bonpland
1956]
COOPER——AUSTRALIAN AND NEW ZEALAND PITTOSPORUM 165
oo hirsutum Link, Enum. Hort. Berol. 1: 233. 1821, ex char. (T.: ex Hort.
E feiere Aiton var. tomentosum (Bonpl.) Bailey in Queensland Agr. Jour.
95; 19
Shrubs to 5 m. tall; branchlets grayish-brown, rusty-tomentose when young,
glabrescent. Leaves alternate, usually crowded at the tips of the branchlets,
elliptic-ovate to elliptic-oblong, acute to shortly acuminate at apex, attenuate at
base, entire, 4 - 19 cm. long, 2 - 7 cm. broad, dull green above, glabrate or covered
with fine appressed red hairs beneath, subcoriaceous, margins thickened and revo-
lute, occasionally undulate, costa and side veins sunken above, raised beneath, side
veins 4-10 per side, anastomosing; petioles 5 - 18 mm. long, 1-2 mm. broad,
rusty-tomentose. Inflorescences terminal, rarely axillary, 1- to 11-flowered,
cymose-umbelliform; peduncles brown-tomentose, subtended by several leaves
and a whorl of persistent tomentose bud scales 2 - 5 mm. long. Sepals slightly
imbricate at base, lanceolate, acuminate with recurved tips, 4 - 8 mm. long, 1.5 -
3.0 mm. broad, sparsely tomentulose and ciliolate, glabrate; petals linear-oblanceo-
late to linear and pandurate, spreading to reflexed at the tips, obtuse to subacute
at apex, clawed, 12.5 - 18 mm. long, 2.5 - 5.0 mm. broad, connivent from the
base for two-thirds their length in a cylindrical or somewhat urceolate tube,
yellow; stamens 6.4 - 12.0 mm. long, anthers sagittiform to oblong, 1.9 - 4.0 mm.
long, 0.5 - 1.5 mm. broad. Pistil at anthesis equal to or slightly longer than the
stamens; ovary 3.5 - 6.0 mm. long, 2 - 4 mm. broad, tomentose; style 3.5 - 7.0
mm. long; stigma capitate and obscurely 2- to 4-lobed. Capsules ellipsoid to sub-
globose, 4- to 6-lobed, 2- to 4-valved, apiculate, 1.5 - 2.8 cm. long, 1.1 - 2.1 cm.
broad, green to brown, coarsely rugose, tomentose or glabrate; valves convex to
sulcate in transverse section, 2 - 3 mm. thick, woody, with a placenta bearing
peg-like funicles up to 4 mm. long from the base to near the apex; seeds 20 - 76,
red to reddish-brown, round to irregular.
Occurs from northern Queensland to Victoria, Australia. Flowers from July
to October, earlier in Queensland, later in New South Wales and Victoria.
I LIA: QUEENSLAND: Dalrymple Heights, South Kennedy District, Clemens
rubby Creek, Herberton Range, White
Jf ; ockingham
Y» y, Dallecky s Ss (F, MEL, MO); Herbert River, Eaton s.n. (BRI); pee ene
Daintree e sm. (MEL); Port Mackay, Mueller s.n. (MEL); c creek alluvium, Rose
a Gm Pialba, White s.n. (A, BRI); in scrub in sandy valley, Fraser Island, Hubber
vi same y n. Petrie 120 (BRI); Wide Bay, Eaves s.n. ref
E) KE Simmonds s.n. (BRI ^
fre country, Mudgeeraba E 41 $0 (K); Moreton Bay, vita collector s.n.
ueller
soil, bank of
ne ,ina pv:
Gegen River, See POG Habban 3701 (A; BM, BRI, K); Cor-
rumbin, Longman s.n. (K); brown loam, rain forest, 3000’, Springbrook, Macpherson
[Vol. 43
166 ANNALS OF THE MISSOURI BOTANICAL GARDEN
{ NV ds |
em, ER
É Y, Ss VK
Ju
Fig. 29. P. revolutum.
1800', Mixinis Creek valley, Mate est sien . (NSW); eg of es
ca. 2000’, Macpherson Range, Bailey s.n. dent ‘Tambourine M Longman & White
(BRI); Point SE Schneider s.n. (BRI); Mt. Lindsay, border ui and New
South Wales, White s.n. (A). NEW SE WALES: Tweed, Guilfoyle s.n. (MEL); Bruns-
wick River, Maiden & Boorman s.n. (NSW); Stanthorpe, Davidson 158 (BRI); Richmond
š : immon
same "zer ës Goat oe? Cheel s.n. (NSW); ; erg in i — on river bank, Tanner 8
Stuart s.n. (MEL); Coffs Ha ege Boorman s.n. (BISH, N SW); Mackasy Rive, Beckler
s.n. (MEL); Hastings — Mueller (U); Gloucester Buckets, Maiden (NSW);
same locality, bush at foot of — Gregson s.m. (NSW); Upper Wi illiams River,
Fraser & Vickery s.n. (NSW ); same locality, WP oia in brush forest, White 11474
(A, BRI); Bulahdelah SC Forestry Officer 20 (NSW); banks of Hunter’s, William’s
and Paterson’s rivers, Brown 0 (BM); Hunter’s River, Brown 5450 (BM, K); Box
Point to Kangaroo River, Maiden s.n. — ; shaded woods on the coast, qe
District, Port Jackson, Cardunnee, A. Cun ingbam 18 (BM, K); Port Jackson, R
Sn., 08 (BM, K, MO); same locality, Can feld s.n. (BISH, US); same locality, ‘Carey
s.n. (GH, K); prope Sydney, Anderson (BM); Manly, Helms s.n. (NSW); Cronulla,
Steenbobm s.n. —: near Sydney, Bede s.n. (NSW); Paramatta, Caley s.n. (A);
Nelson's SE Boorm n. (US); poa, Compete sm. (US); Mt. Ke mbla, Se
1956]
COOPER—AUSTRALIAN AND NEW ZEALAND PITTOSPORUM 167
695 (NSW); Nepean River, without collector s.n. (K); River Grose, Brown 545 0 (BM);
Grose Vale,Vickery s.n. (NSW); Liverpool, Moore s.n. (BM); Bulli Pass Road, Carne $
Hudson sm. (NSW); Cambewarra, Rodway 871 (K); near Nowra, Barnard 109
y K
iver, without collector s.n. (MEL). wirHouT LocALmY: N. S. Wales, Fraser s.n. (A),
114, 115 (BM); N. S. Wales, Caley s.n. (A, BM); N. S. Wales, A. Cunningham s.n. (K);
cote meridionale, Baudin s.n. (P) ; A. Cunningham 248, 249, 265 (U); Fleming 1 (BM);
Macarthur s.n. (K). CULTIVATED: Jardin de la Malmaison, Bonpland s.n. (P), ex Hort.
Kew s.n. (BM)
Pittosporum hirtum (Willd. Enum. Hort. Berol. p. 261. 1809) is sometimes
cited as a synonym of P. revolutum Ait. but I have not seen specimens and the
home of the plant is given by Willdenow as the Canary Islands.
Pittosporum revolutum is not common in cultivation but I have seen speci-
mens from Italian, Californian, and Australian gardens, The leaves of specimens
from New South Wales are frequently much smaller than those from Queensland,
but the extremes are connected by numerous intermediates and I am unable to
recognize distinct forms.
Vernacular names: Wild Yellow Jasmine (Lagoon Pocket, Lowe 14) ; Yellow-
flowered Brisbane Laurel; Genoa Pittosporum (Victoria).
25. PrrrosPoRUM RUBIGINOSUM A. Cunn. in Ann, Nat. Hist. 4:108. 1832.
(T.: A. Cunningham s.n. !).
Pittosporum wingii F. Muell. in South. Sci. Rec. n.s. 1: 49. March 1885. (T.: Dallachy
s. 1).
Shrubs 1.5 - 6.0 meters tall, branchlets brown and rusty-tomentose. Leaves
basically alternate but rather irregularly subverticillate, elliptic to lanceolate or
oblanceolate, acutely acuminate to cuspidate at apex, attenuate at base, entire,
6.0-33.5 cm. long, 2.5-13.0 cm. broad, dark green above, pale green
to reddish-purple beneath, sparsely tomentose to glabrescent above, sparsely to
densely tomentose beneath, especially on the veins, the hairs dark red and erect,
membranous, margins flat or sometimes irregularly revolute, costa and secondary
veins sunken above, raised beneath, rusty-tomentose, secondary veins 8 - 13 per
side, anastomosing; petioles 2.5 - 12.0 mm. long, 1 - 2 mm. broad, tomentose.
Inflorescences variable, consisting of a shoot 4.0 -11.5 cm. long, red-tomentose,
subtended by a whorl of leaves and bud scales and a sequence of cataphylls which
become foliar above, sometimes producing 1 to several solitary flowers in the
axils of the uppermost cataphylls and sometimes being terminal, 3- to 32-flowered,
umbelliform; peduncles and pedicels red-tomentose, accrescent in fruit, subtended
by numerous caducous foliar to scarious bracts. Sepals free or slightly imbricate
at base, linear-lanceolate, acute to acuminate with recurved tips, sparsely ciliate
with scattered hairs to densely ciliaté, tomentose, 4.0 - 8.5 mm. long, 1 - 2 mm.
broad; petals oblanceolate to linear-oblong, sometimes pandurate, obtuse to sub-
acute, 11 - 17 mm. long, 2 - 4 mm. broad, coherent from the base to the middle in
[Vol. 43
168 ANNALS OF THE MISSOURI BOTANICAL GARDEN
a somewhat urceolate to spreading tube, or free and patent; stamens 4.5 - 11.5
mm. long; anthers sagittiform to ovate-elliptic, 1.0 - 3.5 mm. long, 0.5 - 1.0 mm.
broad. Pistil at anthesis slightly shorter or longer than the stamens; ovary 2.0 -
5.5 mm. long, 1 - 3 mm. broad, villous; style 3.5 - 6.5 mm. long; stigma capitate
and obscurely 2- or 4-lobed or truncate. Capsules ellipsoid to ovoid, 2-, or rarely
3-, valved, 1.3 - 2.1 cm. long, 6 - 15 mm. broad, with an evident stipe 0.5 - 4.0
mm. long, yellow to orange, densely to sparsely rusty-tomentose or glabrate,
minutely rugose; valves convex to slightly sulcate in transverse section, less than
1 mm. thick, coriaceous, with a placenta thickened at the base and bearing peg-
like flattened funicles up to 2.5 mm. long from the base to about the middle;
seeds 3 - 14, dark rusty-red to red, irregular.
KEY TO THE SUBSPECIES
Sege 1.5—2.5 m. HH leaves sparsely rusty-tomentose to ás beneath; ri bag?
e with scattered hairs; petals coherent to the middle i somewhat urceolat
cylindric al ibi, pah blok al sometimes pandurate; stamens 6.0-11.5 mm. ori et
ease capsules sparsely ceat ose to glabrate. Queensland: Cooktown to
Pipe P irn 25a. P. r. rubiginosum
Shrubs to 6 m. SC? leaves densely rotes tomentose beneath; sepals densely ciliate and rusty-
tomentose; petals —€— to SEN middle in a spreading tube, linear-oblong to resin ion
stamens 4. 16 0 mm anthers 1.5 -3.5 mm. dons: capsules densely rusty-tom .
Bay Sat?
25a. PrrrosPORUM RUBIGINOSUM ssp. rubiginosum
Known from Cooktown to south of Cairns, Queensland, from near sea-level
to 16 geg m. dies from July to mee
LAND: sum of Mt. Cook, Endeavour River, A. Cunningbam
26 (BM, MO); Powy Gë on des Sie of Mt. Cook, A. Cunningham 26 (K); need
— Rose 109 (MEL, NSW); Mossman River, Sayer 186 (MEL); same locality, L
MEL); in hillside forest in the lowlands, Daintree River, Brass & White 279 (BRI);
i tiia Tp Daintree, k District, Blake 14996 (BRI); same locality, Brass 2220 (A,
SH. D
LIS ma
Toog $ Zoch Freshwater ees near Cairns, Francis s.n. (BRI, K); same locality,
. and
Johnstone River, AE 9, 30, 117 (BRI); same Last eier s.n. RI, K); State
Forest Reserve 185, Danbulla, Doggrell A.5. (BRI); Mt. Alexandra, [be 13 (BRI).
Specimens collected near sea-level have leaves much longer than those from
high elevations; e.g. Brass & White 279 from lowlands near the Daintree River
has leaves 19 cm. long and 6.5 cm. broad, and Blake 14982 from 100’ above sea-
level at The Intake, near Cairns, has leaves up to 20 cm. long and 6.5 cm. broad.
Brass 2090 from 2500’ on Mt. Dimi has leaves up to 11.5 cm. long and 3 cm.
broad, and White & Brass 227, from about 4000’ on Thornton Pi Peak, has leaves up
1956]
COOPER—AUSTRALIAN AND NEW ZEALAND PITTOSPORUM 169
to 10 cm. long and 4 cm. broad. Changes in leaf size are gradual and I cannot
define distinct forms.
The flower parts also vary in size. Flowers which may be female have capi-
tate and obscurely 2-lobed stigmas, styles 4 - 5 mm. long, plump ovaries 4.0 - 5.5
mm. long, 3 - 4 mm. broad, short stout filaments 4 - 6 mm. long, probably abortive
sagittiform anthers 1 - 2 mm. long, and an almost urceolate corolla-tube from
which the stigma is exserted. Flowers which may be male have truncate stigmas,
styles 5.0 - 6.5 mm. long, slender ovaries 2.0 - 4.5 mm. long, 1-2 mm. broad,
slender filaments 6 - 9 mm. long, probably functional ovate-elliptic anthers 2.5 -
3.0 mm. long, and a tubular corolla in which the stigma is included Flower color
is described as "white tinged with cream" (Blake 14982), and “yellow, paler
toward the base" (Blake 15242).
Vernacular name: Rusty Pittosporum.
°F. RUBIGINOSUM ssp. RUBIGINOSUM
> | YS DA *
§
p 5 |
A P. geen.
| e
de
Na
Fig. 30. P. rubiginosum ssp. rubiginosum and ssp. wingii.
25b. Prrrosporum RUBIGINOSUM ssp. Wingii (F. Muell.) R. C. Cooper, stat.
nov.
Pittosporum wingii F. Muell. in South. Sci. Rec. n.s. 1: 49. March 1885. (T.:Dallachy
s.n
"een from the Atherton Tableland to Rockingham Bay, Queensland.
Flowers in September.
[Vol. 43
170 ANNALS OF THE MISSOURI BOTANICAL GARDEN
AUSTRALIA: QUEENSLAND: top of the Coast Range, under the rocks, Rockingham
Bay, Dallachy s.n. (GH, K, MEL MO); 800 m., Lake Barrine, Atherton Tableland,
Kajewski 1341 (A, BRI, K, MEL, NSW).
Variation in flower size and form has also been found in this variety. The
Rockingham Bay flowers have truncate stigmas, styles 5 mm. long, ovaries 3 - 4
mm. long, 2.5 - 3.0 mm. broad, filaments 5 - 7 mm. long, and probably functional
anthers 3.0 - 3.5 mm. long. The sepals are 7.5 - 9 mm. long, and the petals 13
mm. long. The Atherton Tableland flowers have 4-lobed stigmas, styles 3 - 4 mm.
long, ovaries 3.5 - 4.0 mm. long, 2.5 - 3.5 mm. broad, filaments 3 - 4 mm. long,
and apparently abortive anthers only 1.5 - 2.0 mm. long. In one flower two large
functional anthers and three small aborted anthers were found. The sepals are
5 - 9 mm. long, and the petals 9 - 11 mm. long. It is possible that the Rocking-
ham Bay flowers have been collected at a later stage in development than those
from Atherton Tableland, but the slender ovaries, larger styles, truncate stigmas,
and functional anthers of the former suggest that the flowers are male, while the
plumper ovaries, shorter capitate stigmas, and apparently abortive anthers of the
latter suggest that the flowers are female.
26. PrrrosPORUM UNDULATUM Vent. Descr. Pl. Nouv. Jard. Cels, #. 76.
[1802 ]1"° ex char. & icon.
Shrubs or trees 5 - 13 m. tall; branchlets gray, the young parts tomentose,
soon glabrate. Leaves alternate, elliptic-oblong to oblanceolate, acuminate at apex,
attenuate at base, entire, 6 - 16 cm. long, 1.5 - 5.0 cm. broad, green above, paler
beneath, tomentulose when young, glabrate, membranous, margins undulate or
flat, sometimes recurved, costa sunken or raised above, raised below, secondary
nerves about 12 per side, anastomosing, distinct; petioles 7 - 26 mm. long, 1 - 2
mm. broad, tomentulose when young, glabrate. Inflorescences terminal, 4- to 15-
flowered, subumbelliform; peduncles and pedicels 1.9 - 3.1 cm. long, accrescent
in fruit, tomentose; peduncles subtended by a whorl of leaves and numerous
caducous, brown-tomentose, ciliolate bud scales 1 - 2 mm. long. Sepals frequently
connate in a tube which splits into 2 parts, one 1- to 2-lobed, the other 3- to 4-
lobed, lanceolate, acuminate, 6.5 - 10.5 mm. long, 1.2 - 2.5 mm. broad, tomentulose,
faling before the petals; petals linear-oblanceolate to linear-oblong, obtuse, 11 -
17 mm. long, 3 - 4 mm. broad, coherent at the base, spreading to recurved above,
white; stamens 5 - 11 mm. long, sometimes reduced to sterile rudiments 0.5 - 1.0
mm. long; anthers lanceolate-oblong to sagittiform, 2.5 - 5.0 mm. long, 1.0 - 1.5
mm. broad. Pistil at anthesis slightly shorter or longer than the stamens; ovary
4 - 6 mm. long, 1.8 - 4.0 mm. broad, tomentulose; style 2.0 - 4.5 mm. long;
stigma capitate and obscurely 2- to 4-lobed or almost truncate. Capsules sub-
globose, 2-valved, 10 - 14 mm. in diameter, yellow to brown, smooth to slightly
rugose, glabrous; valves convex in transverse section, less than 1 mm. thick,
170 W. T. Stearn (Jour. Soc. Bibl. Nat. Hist. 1:199-201. 1939) gives the dates of publication of
this work as 1800-1802, 7.76 being published in 1802.
1956]
COOPER— AUSTRALIAN AND NEW ZEALAND PITTOSPORUM 171
coriaceous, with a placenta thickened at the base and bearing 2 - 3 rows of short
stout peg-like funicles from the base to just above the middle; seeds 12 - 22, red
to dark red or black, irregular.
Occurs from the Macpherson Range, in southern Queensland, to the Victorian
Alps, along the ranges of New South Wales. Flowers in August and September.
Fig. 31. P. undulatum.
USTRALIA: QUEENSLAND: Bunya Mts, Bailey s.n. (BRI, NSW); same locality,
White s.n. (BRI); same EH a s.n. (BRI); 2800', at edge of scrub in partial
shade, perd Mistake, Hubbard 5225 (A, K); same locality, "Shirley s.n. (A); Eumundi,
Si ü í ; n in
, Sou
SÉ s.n SW "itd: localit
GE ele locality adi onds sm. (A); near Canungra, Brass s. (NSW);
. Park, Moore És. (BRI); Beechmont, White 6179 (BRI); Acacia Creek via
Killarney, Been 129 (NSW); near Emu Vale, Macpherson Range, Francis s.n. (BRI);
ingham's Gap, Bailey s.n. (BRI); on Rhyolite at edge of rain forest in wet sclero-
phyll forest, ca. u— Nixon's Creek, Upper Falls, je cpherson Range, Jobnson 106
(N NEW SOUTH WA s in n.
= SCH Tilba, Bate 9 edge near Tentafield, Stuart s.n. (K); Conjola, Heron
sm. (N ; Lismore, Maiden 17 (U); New England, Stuart s.m. dues d) in bed of
creek, ca. pé. miles N.N.E. of Boggabi, towards Nandewar Range, Swain s.n
N
Barraba, Kifford s.n. (NSW); Hastings iie Beckler s.n. d U); Gloucester Buckets,
t branch, Tisi 5451
; occasional in pae
on granite, 690 m., Ruby Creek, Mt. Werong, ca. 30 Se south ot Oberon, Johnson &
Constable s.n. (K, N ur Ci upper reien River; Fraser & Vickery s.n. (A, BISH, BH,
MO, NSW, US); on old sand dunes, peninsula north of the entrance to Tuggerah, via
[Vol. 43
172 ANNALS OF THE MISSOURI BOTANICAL GARDEN
woods on the sea-coast, NH Port P A. NOM s.n. (K); de Séi Port
zeg Brown 5451 (BM); Port Jackson District, Brown 5451 (K, MO); same locality,
Siemssen 97 (MEL); same locality, Steel s.n. (NSW); same locality, Helms s.n. (BRI);
same locality, Camfield s.n. (MO) ; near Sydney, Weber s.n. (NSW); rs Set
s.n. (NSW); Petersham, Steel s.n. (AK); Sydney, Haswell s.n. (BRI); same pelin
Backhouse s.n. (BM); same locality, Wilkes s.n. (K, US); same locality, McWilliam
(K); same locality, Verreaux 406 (K); Illawarra, Macarthur 26 (K); pá, Mountain,
Maiden s.n. (NSW); Sugar Loaf Mountain, Braidwood, Baeunton s.m. (M SW);
Queanbeyan, Breakwell s.n. (NSW); 1200’, between Nowra and Kangaroo v Wi
bewarra Range, Rod way Á 690 (A, K); Nowra, Rodway 361, 847 (K); Shoslhavt River,
Wingello, Murphy 9 (NSW); Comerong Is., same localiey, Rodway 874 (K); Candelo,
White s.n. (MEL); Bud. n" Father s.n. (A); same locality, Mueller s.n. (MEL).
VICTORIA: Genoa River, Reader s.s. (MEL); Broadrib, or Broadribb River, Mueller s.n.
(K, MEL); in rupibus prope Bachan ad flumen E Mueller s.n. (MEL); Snowy
River, Mueller s.n. (K, MEL); Stony Creek, Smith s.n. (MEL); East Gippsland, M Mueller
s.n. (GH, MEL); Dead-Cock Creek, E. of Lindenow, Green s.n. (MEL); in small es
gully above sea cliff, Mt. Martha, E. Port Phillip, Johnson s.n. (NSW); in moist and-
stone gully, Epping, Ford s.s. (NSW); Wilson's property, near oes Serzlecki
anges, without collector s.n. MED. WITHOUT DEFINITE LOCALITY: Australia, Ver-
reaux 48 (BM), 44P (P, US); East Australia, Schomburgk s.n. (M EL); | South Australia,
cbe ei s.n. (NSW); Australia, Lbofsky s.n. (BM); Australia, Sieber 221 (A, BM,
» Be O); . Wales, A. Cu de ws or "Anderson and otbers 321 (U);
N. ` Wales, Caley s.n. (B M).
Pittosporum undulatum is widely cultivated as an ornamental, hedge, or
shelter plant, and I have seen specimens from New Zealand, Australia, Chile,
Colombia, Bolivia, Jamaica, Bermuda, the United States (California and Florida),
the Azores, the Canary Islands, France, Israel, India, Ceylon, China, and Hawaii.
It has become naturalized in Bermuda, the Canary Islands and Hawaii. A form
with variegated leaves is available in the trade in Australia"! and a small-leaved
plant, labelled ““Pittosporum hybrid," and “a glabrous-leaved form of P. bic
have been collected from the Strzlecki Ranges, Victoria, (without collector s.n.
MEL) and from South Australia (Schomburgk s.n. NSW). The last form is
connected with the large-leaved plants by intermediates, and I am unable to
recognize it as a distinct variety. According to Bentham,"? the height of P.
undulatum is "about 40 feet, or according to M'Arthur 60 to 90 feet". Recent
collectings are from smaller trees, but a plant at East Alameda Plaza, Santa Bar-
bara, California, is stated to be about 60 feet tall and 50 feet in crown diameter
(Moran 2349 BH), so it is possible that trees of 60 to 90 feet were found in the
early days of Australian botany.
The variation in the morphology of the flower has been described and illus-
trated by Maiden,173 Gowda,!"4 and Cufodontis. Flowers which may be female
171 Pittosporum undulatum "variegatum", commonly known as the Variegated Sweet Pittosporum
deel Shrubs and Trees for Australian Gardens. p.34. 1948).
115 Österr. Bot. Zeitschr. 98: 114. 1951.
1956]
COOPER—AUSTRALIAN AND NEW ZEALAND PITTOSPORUM 173
have capitate obscurely lobed stigmas, styles 2 - 3 mm. long, and ovaries 4.0 - 4.5
mm. long and 2 - 3 mm. broad. The stamens are rudimentary, gland-like structures
0.6 - 1 mm. long. Flowers which may be male or bisexual have weakly capitate
to truncate stigmas, styles 3.0 - 4.5 mm. long, ovaries 3.5 - 5.0 mm. long, 1.5 -
2.0 mm, broad, stamens 8.5 - 11.0 mm. long, and functional anthers 3.5 - 5.0
mm. long and 1.0 -1.5 mm. broad. Intermediate forms occur in which the stamens
are reduced, the most remarkable being Ford s.n. in which the stamens vary in
length from 5 - 7 mm. in the same flower. Flowers also occur with 1 - 2 extra
sepal lobes, and petals.
Vernacular names: Wave-leaved Pittosporum, Cheesewood, Engraver's wood,
New South Wales Box Tree, New South Wales Mock Orange, Native Laurel,
Victorian Laurel, Sweet Pittosporum (Victoria), Wallundundeyren, Bart-Barb
(Aboriginal).
27. PrrrosPoRUM VENULOSUM F. Muell. Frag. Phyt. Austr. 6: 186. 1868. (T.:
Dallacby s.n. !).
Trees to 10 m. tall; branchlets gray, rusty-tomentose when young, soon
glabrate. Leaves alternate to subverticillate, lanceolate to ovate-lanceolate, acute
to acuminate at apex, attenuate at base, entire, 4.5 - 10.5 cm. long, 1.7 - 4.2 cm.
broad, rusty-tomentose when young, especially on costa, glabrate, coriaceous,
margins undulate, irregularly revolute, costa immersed or sunken above, raised
below, secondary veins 8 - 12 per side, anastomosing, distinct; petioles 6 - 21 mm.
long, 1 - 1.5 mm, broad, rusty-tomentose when young, soon glabrate, Flowers
terminal, in umbelliform cymes; peduncles subtended by a loose whorl of leaves
and by numerous caducous rusty-tomentose bud scales 2 - 6 mm. long; peduncles
and pedicels accrescent in fruit, rusty-tomentose, soon glabrate. Sepals not im-
bricate, oblong, obtuse, 4.0 - 4.5 mm. long, 1.5 - 2.0 mm. broad, ciliate, rusty-
tomentose without, glabrous within; petals oblong, obtuse, 8 - 11 mm. long, 1.5 -
2.0 mm, broad, free, spreading at the tips, white; stamens 4-5 mm. long,
anthers sagittiform, 2.0 - 2.5 mm. long, 0.5 - 1.0 mm. broad. Pistil at Ge:
longer than the stamens; ovary 4.0 - 4.5 mm. long, 2 mm. broad, rusty-tomentose
style 1.5 - 2.0 mm. long; stigma capitate, obscurely 4-lobed. Capsules ll
to obovoid with an evident stipe, 2-, rarely 3-, valved, apiculate, 1 - 2 cm. long,
1.0 - 1,4 cm. broad, glabrate, rugose; valves convex in transverse section, 1 - 2
mm. thick, woody, with placentas much thickened and fused at the base, bearing
6-8 peg-like funicles up to 2 mm. long about the middle; seeds 6-14 per
capsule, reddish-black to black, irregular.
Occurs in the coastal ranges of Queensland, between Rockingham Bay and
ast Range, at edge of scrub, M Bay,
Ka s.n. ». (BM, x, MEL, NSW); ae Range, Francis s.n. (BRI ; rare, at
edge of light rain forest on rather steep hillsides, Byfield near Keppel Bay, White 8142 (A,
BRI) ; Mt. Spec, White 8974 (A, BH).
I have not seen adequate flowering material and have described the inflores-
cence from immature specimens.
[Vol. 43
174 ANNALS OF THE MISSOURI BOTANICAL GARDEN
Fig. 32. P. venulosum. Fig. 33. P. melanospermum.
28. ME MELANOSPERMUM F, Muell. Frag. Phyt. Austr. 1: 70. 1859.
(T.: Holt s.n.
cue y siae Bailey, in Queensland Agr. Jour. 1: 451. 1897. (T.: Barclay-
rs.n
Pitlostoreá SEH Domin, in Bibliog. Bot. 22: 718. 1925. (T.: Schultz 758 !).
Shrubs or small trees to 6 m. tall; branchlets reddish-gray, glabrous. Leaves
alternate, sometimes crowded, oblanceolate to oblong, rarely obovate, obtuse to
acuminate or mucronate at apex, attenuate at base, entire, 4 - 14 cm. long, 1.1 -
4.5 cm. broad, glabrous, coriaceous, margins thick, flat to slightly undulate, costa
immersed above, raised below, secondary veins 13-18 per side, anastomosing,
raised on both surfaces; petioles 1.4 - 4.0 cm. long, 1 mm. broad, glabrous. Flowers
terminal, very numerous, in spreading paniculiform cymes; peduncles subtended
by an approximate whorl of 3 - 4 leaves and minute caducous bud scales, and
bearing branches which are subtended by caducous cataphylls, and which are again
divided, the ultimate divisions each bearing several pedicels; peduncles, branches,
and pedicels accrescent in fruit, white-tomentulose, soon glabrate. Sepals slightly
imbricate to coherent in a shallow cup, ovate, subacute to acute, 1.5 - 2.5 mm.
long, 1 - 2 mm. broad, white-tomentulose becoming glabrate; petals obovate to
lorate, obtuse, 7 - 9 mm. long, 2.0 - 3.5 mm. broad, free, patent, yellow; stamens
6.5 - 10.5 mm. long, anthers elliptic-ovate, 1.5 - 2.0 mm. long, up to 1 mm. broad.
Pistil at anthesis slightly shorter than the stamens; ovary 2.5 - 3.5 mm. long,
0.5 - 2.0 mm. broad, white-tomentose, on a glabrous stipe about 1 mm. long;
style about 1.5 mm. long; stigma truncate. Capsules obovoid to subglobose, 2-
1956]
COOPER— AUSTRALIAN AND NEW ZEALAND PITTOSPORUM 175
valved, apiculate, 6 - 15 mm. in diameter, frequently with a stipe 0.5 - 1.5 mm.
long, glabrate, brownish-yellow; valves convex to sulcate in transverse section,
less than 1 mm. thick at sides, coriaceous, with a placenta much thickened at base
and apex, and occasionally fused to form 2 chambers, bearing several short stout
peg-like funicles between the base and middle; seeds 2 - 6, black, irregular.
Known from northern Queensland and the Northern span
ALIA: NORTHERN TERRITORY: Arnhem’s ec Mueller deve north E
);
AND: Cape Y Yotk: Pesini: Eier At Hann 98 (K); Walsh River;
Barclay-Millar s.n. (A, BM, BRI, K); common in rain forest, 500 m., Lake Barrine,
Atherton Tableland, Kajewski 1352 (A, BRI, P).
29. PrrTOSPORUM RHOMBIFOLIUM A. Cunn. ex Hook. Icon. Pl. 7: 7. 62I. 1844.
(T.: A Cunningham 29 !).
Trees 13 - 30 m. tall; branchlets light brown, glabrous. Leaves alternate,
sometimes crowded at the tips of the branchlets, rhombic to ovate, acuminate to
obtuse at apex, attenuate at base, coarsely serrate to entire, 4 - 10 cm. long, 1.5 -
7.0 cm. broad, glabrous, coriaceous, margins frequently flat or revolute, costa
sunken above, raised beneath, secondary veins about 12 - 14 per side, anastomos-
ing, raised above and below; petioles 0.9 - 2.7 cm. long, 1 - 2 mm. broad, glabrous.
Flowers terminal, many, in spreading umbelliform cymes; peduncles and pedicels
up to 5 cm. long, accrescent in fruit, glabrous, peduncles subtended by a single
leaf and 1 or 2 minute caducous bud scales. Sepals coherent at base, ovate, acute
to subacute, 1.1 - 3.0 mm. long, 0.5 - 2.0 mm. broad, glabrous; petals elliptic to
oblanceolate, obtuse to subacute, 5.5 - 8.5 mm. long, 1.5 - 3.0 mm. broad, free,
patent, white, with short claws; stamens 5.5 - 8.0 mm. long, anthers lanceolate,
1.7 - 2.4 mm. long, about 1 mm. broad. Pistil at anthesis slightly shorter than
the stamens; ovary 3.5 - 4.0 mm. long, 1 - 2 mm. broad, short-stipitate, tomen-
tulose; style 0.5 - 1.0 mm. long; stigma truncate. Capsules subglobose to obovoid,
2-valved, obtuse to apiculate at apex, stipitate at base, 5 - 10 mm. long, 5 - 8 mm.
broad, yellow, slightly rugose, glabrous; valves convex to sulcate in transverse
section, less than 1 mm. thick, coriaceous, with a conspicuous placenta more or
less fused to form 2 chambers and bearing 1 - 2 minute funicles near the base;
seeds 1 - 3, black, irregular.
Occurs from northern Queensland to northern New South Wales, Australia.
jenes in November, December and January.
USTRALIA: QUEENSLAND: Canaan Valley, near digna: gs 973 jem
NSW); T ooloom Range, Maiden s.n. (NSW); Rockhampton, Thozet s.m. (MEL); sam
locality, Ou 87 Ze (MEL) ; in red, sandy loam, 12 ac east of Emerald, ere
District, Everist 2522 (BRI); in mixed soft-wood forest, Guluguba, €— dE rios.
White 1146 (BRI); Son een Brisbane and Dawson, Moeller a s.n. (K); Dawson Rive
Mueller s.n. (MEL); Gladstone, Bailey s.n. (NSW); Mt. Perry, Keys s.n. (BRI); Eidsvold,
d laville, 50 wed ; of Bundaberg, Bancroft s.m. (BRI); pari
of Ka Sape: — Seet ct? , Simon (BRI); Maryborough, Young s.n. (BRI,
NSW); Wide Be Bay, Bidwill 52 pe in he sub-xerophytic mixed scrub, Roma, White
[Vol. 43
176 ANNALS OF THE MISSOURI BOTANICAL GARDEN
9522 (A, BRI); Yalebone Creek via Rome, McKenzie s.n. (BRI); in "ringed" brigalow
scrub, Chinchilla, Beasley 27 (BRI); Blackbutt eta Shirley s.n. (A); Yarraman,
Clemens 5.5. (BRI); Crow’s Nest, Kenny s.n. (BRI); same locality, Clemens 43747 (A);
Crystal Brook, Bowenville, Fuller s.n. (NSW); Moreton Bay, A. Cunningham s.n. (GH);
Brisbane, Bailey s.n. (BRI, NSW); in shaded woods on the banks of the Brisbane River,
A. Cunningbam 29 (BM, BRI, K); same locality, A. Cunningbam s.n. (MEL); same
locality, White s.n. (NSW); Enoggera, near Brisbane, Boorman s.m. (NSW); Ipswich,
Nernst 34 (MEL); Beau-desert, Brass s.n. (A); Roberts Plateau, Lamington dinge
€ SE 6045 (A); Milford, Fassifern District, ety el 2024 (A dree"
, White 1903 (A, BRI); Acacia Creek, prés arney, Dunn. 136 (N Y)
Pubs ark, Shirley s.m. (NSW); in regrowt d? in-forest margin, ca. Wei
O'Reillys, Lamington National Park, Smitb t$ Webb 361 2 (BRI); common in a rain
forest, Unumgar, near Mt. Lindesay, NSW-Q border, White 12507 (BRI). NEW SOUTH
ALES: Tweed, Guilfoyle s.n. (MEL); Burringbar, Betche s.m.
NSW);
Z
f
id
ver,
Richmond River, Watts s.n. (NSW); Clarence River, vitiaat calicis sm. (NSW
WITHOUT LOCALITY: Walter s.n. (MEL).
The plant is useful as an ornamental tree, and I have seen cultivated speci-
mens from California, Florida, Queensland, New South Wales, and South Australia.
Vernacular names: White Myrtle, Diamond-leaf Laurel, Rhombus-leaved
Queensland Laurel, Burrawingee.
^U EH m 450 a
x 7 -
quem
d $
i 2
LS `
S e
AL Es d Ww RS
e - iee MA
= far "9 Es
i be rhe Th Zë
| AS CTS A e
+ De, een p t t WX S KR , e
e erg e he SE EE
dä Ki J pa £ ^ d AD Rn
> E 3 Ser Ce r5 +
yx
jijé A S e Ve
A SNS
é
` Aad D
MA AA
ó Ai E m p d
. lu 2 i $ d£
MOE | KN /
| Aly
^ L Se R
Ka GE
P SS a
Fig. 34. P. rbombifolium. Fig. 35. P. ferrugineum.
1956]
COOPER— AUSTRALIAN AND NEW ZEALAND PITTOSPORUM 177
30. PrrrOsPORUM FERRUGINEUM Aiton, Hort. Kew. ed.2. 2: 27. 1811. (T.: ex
bort. Kew. !).
deng "wë A. Cunn. in Ann. Nat. Hist. 4: 109. 1839, as tinifolium. (T.: A.
nning 2.1).
oia Qvatifolium F. Muell. Frag. Phyt. Austr. 2: 78. 1860. (T.: ary s.n. $
Pittosporum MUT Domin, in Fedde. Repert. Sp. Nov. 11: 201. 1912. (T.
R. Brown 5449 !).
Trees or shrubs 8-20 m. tall; branchlets grayish-brown, rusty-tomentose
when young, glabrate. Leaves alternate, frequently crowded at the tips of the
branchlets, elliptic to elliptic-ovate or obovate, acute to acuminate or occasionally
obtuse at apex, attenuate at base, entire, 4.8 - 11.5 cm. long, 2.0 - 5.5 cm. broad,
green above, paler beneath, rusty-tomentose when young, soon glabrate except on
costa, margins flat or recurved, costa sunken above, raised below, secondary nerves
4 - 8 per side, anastomosing, obscure above, raised beneath; petioles 1.2 - 2.2 cm.
long, rusty-tomentose when young, glabrate. Flowers terminal or axillary, many,
in subumbelliform cymes; peduncles and pedicels 1.5 - 4.0 cm. long, rusty-tomen-
tose; peduncles subtended by 1 to several leaves and a whorl of ciliolate bud scales
about 1 cm. long; pedicels subtended by 1 to several caducous bracts 2 - 4 mm.
long. Sepals free or coherent at the base, linear to linear-lanceolate, acuminate,
2.5 - 3.5 mm. long, 0.6 - 1.2 mm. broad, sparsely tomentulose and ciliolate; petals
linear to oblanceolate-linear, 7 - 9 mm. long, 1.2 - 1.5 mm. broad, coherent in a
cylindrical tube with recurved tips, yellow; stamens 4.0 - 6.5 mm. long, anthers
oblong, 0.5 - 2.0 mm. long, 0.3 - 0.6 mm. broad. Pistil at anthesis slightly shorter
or longer than the stamens; ovary 3 - 4 mm. long, 0.8 - 2.0 mm. broad, tomentu-
lose; style 1.2 - 2.0 mm. long; stigma capitate and 2-lobed to almost truncate.
Capsules globose, 2-valved, about 7 - 10 mm. in diameter, slightly rugose, glabrous;
valves convex in transverse section, less than 1 mm. thick, coriaceous, with a
placenta thickened at the base, bearing 2 rows of short, stout, peg-like funicles
from the base to just above the middle; seeds 15 - 16, black, irregular.
Occurs mainly on the coast, from Cape York to Rosedale, Queensland; also
found in Malaysia from the Solomon to the Nicobar Islands. Flowers in Australia
from fj to August.
ORTHERN TERRITORY: wier. River, Lea s.n. (BM); QUEENSLAND:
fiy and, Hill 12, T (K); same locality n dry ridges, Mueller s.n. (K); Cape
ity, Duis s.n. » MEL, E US); Vrilya Point, J. F. Bailey s.n. (RD on banks
of the Endeavour Rives: A A. Cunningham : 27, 117 (BM); Endeavour Ridge, at some dis-
ance from the sea, A. Cunniighem s.n. (K), 27 (MO), 117 Zen it) d e ; — Mp
ees 4 (MEL al in rain forest, 4 (BRI,
sane local y, Sayer s ^. (BM); bo botanic r e, Cairns, Ki 10560 BRE, MEL);
Port se owed M acGillivray 105 (BM) ; Rate Cree » Priétpiné, Moree) Zen mei, o"
(K); Pioneer River, Mackay, Griffiths s.m. (MEL, NSW); Port Mackay, Dietrich
[Vol. 43
178 ANNALS OF THE MISSOURI BOTANICAL GARDEN
1299 (MEL); scrub, Pilot Station, Mackay, Griffiths s.n. (BM, BRI); in rocky wa
gullies, and in thickets on the hills, Percy dom. A. Cunningbam s.n. (K); koala
nd, R.
Court s.n. (BRI); Keppel Bay, R. Brown 5449 (BM, K); Rockhampton, Fitzroy River,
Dallachy s.n. (U); in remnants of light rain forest along Archer Creek, near Rockhamp-
ton, Wbite 12216 (BRI); rare, pcs of d ranges, Gracemere, O’Shanesy 1824
(MEL); Crocodile Creek, Capricorn, Bowman 96 (MEL); Curtis Island, Mueller s.n.
(NSW); Gladstone, Hedley 13 GRD Bustard Ba. J. B. [Banks & Solander] s.n. (BM);
common on sandhills near beach, Rosedale, Dovey 140, 153, 1010 (BRI). WITHOUT
LOCALITY: New Holland, Banks & Solander s.n. (MO); East Coast, R. Brown s.n. (BM,
» MO); east coast of Queensland, Mueller s.n. (NSW). curTIvATED: ex hort. K
Se? (BM).
Flowers which may be female have capitate and 2-lobed stigmas, ovaries
about 2 mm. broad, and stamens about 4 mm. long with anthers only 0.5 mm.
long and 0.3 mm. broad. Flowers which may be male have weakly capitate to
truncate stigmas, ovaries about 1 mm. broad, and stamens about 6 mm. long wit
anthers about 2 mm. long and 0.5 - 0.6 mm. broad. Intermediate forms occur
with weakly capitate stigmas, and stamens 4 - 6 mm. long.
Vernacular name: Rust-leaved Pittosporum.
31. PrrrosPoRUM DALLI Cheesem. Man. N.Z. Fl. p. 1134. 1906. (T.: Dall
11).
Small trees 4 - 6 m. Se branchlets gray, the young parts brown and puberu-
lent. Leaves alternate, lanceolate-elliptic to oblong-elliptic, rarely obovate, obtuse,
acute Or acuminate at apex, acute to attenuate at base, coarsely serrate to entire,
5 - 10 cm. long, 2 - 4 cm. broad, dark green above, paler beneath, glabrous,
coriaceous, the margins thin, flat, and ciliolate when young, thickened, slightly
revolute and glabrate when mature, costa raised above and beneath, the secondary
veins 14 - 18 per side, anastomosing, distinct above and beneath; petioles sparsely
puberulent when young, glabrate, 3 - 18 mm. long, 1 - 3 mm. broad. Flowers
terminal, about 40, in condensed umbelliform cymes; peduncles subtended by an
approximate whorl of leaves and by caducous glabrous ciliate bud scales 15 - 21
mm. long; peduncles and pedicels 4-angular, up to 2 cm. long, accrescent in fruit,
white-tomentose, each peduncle bearing 1 to many pedicels subtended by caducous,
glabrous, linear bracts up to 1 cm. long. Sepals not imbricate, linear, 5 - 6 mm
long, about 0.5 - 1 mm. broad, glabrous; petals obovate, obtuse, 8 - 9 mm. Lag
3.0 - 3.5 mm. broad, free, spreading from the base, white with red veins; stamens
7 - 8 mm. long, anthers elliptic-oblong, reflexed, 2 - 3 mm. long, up to 1 mm.
broad. Pistil at anthesis longer than the stamens; ovary glabrous, about 3 mm
long, 1 mm. broad; style about 2 mm. long; stigma truncate. Capsules ellipsoid,
2-valved, about 15 mm. long, 9 mm. broad, green to black, glabrate; valves
convex in transverse section, less than 1 mm. t ick, coriaceous, with a slightly
thickened placenta bearing alternate flattened and peg-like funicles, up to 2 mm.
long, from the base to near the apex, dividing longitudinally into a woody meso-
carp, which is shed on dehiscense, and a persistent membranous endocarp covering
a cone-shaped mass of viscid seeds; seeds about 25, dark red, somewhat trigonal.
1956]
COOPER—AUSTRALIAN AND NEW ZEALAND PITTOSPORUM 179
New ZEALAND: NELSON: mountains, near Collingwood, Dall s.m. (AK); same
locality, 3500', Gibbs s.n. (A, AK, BM), 1854 (K).
Eleven trees have been found at Specimen Creek and Snow's Valley, near
Boulder Lake, northwest Nelson. Others are reputed to occur in the vicinity, but
as the locality is isolated and difficult to visit, the size of the population is not
known. I suspect from their appearance that the flowers of the material available
are male and that female flowers have not been collected.
Cheeseman!?% described the foliage as sharply and coarsely serrate from
the flowering and fruiting specimens available to him ( Dall s.n., Gibbs s.n.).
At Duncan & Davis Nursery, New Plymouth, cultivated plants have serrate
leaves on the lowermost branches and as the lower cataphylls on new shoots. At
the crown of the trees and at the apex of new shoots the leaves are entire. The
species is cultivated as a rarity in New Zealand and England.
Fig. 36. P. dallii and P. eugenioides.
32. PrrTOsPORUM EUGENIowEs A. Cunn. in Ann. Nat. Hist. 4: 106. 1839.
(T.: R. Cunningham s.n and 614 !).
Pittosporum microcarpum Putterl. Syn. Pittosp. p. 15. 1839, ex char. (T.: A. Cunning-
am s.n.).
P ittosporum enkianthoides R. Cunn. & Hueg. in Putterl. loc. cit. p. 7- 1839, nom. nud.
in :
Pittosporum umbellatum A. Cunn. in Putterl. loc. cif. 1839, nom. nud. in synon.
Pittosporum elegans Raoul, in Ann. Sci. Nat. III, 2:121. 1844. (T.: Raoul s.n. !).
Trees 6 - 13 m. tall; branchlets brown, glabrous. Leaves alternate, fre-
quently crowded at the tips of the branchlets, oblong-elliptic to elliptic, subacute
to acute at apex and base, entire, occasionally notched at base in juveniles, 5 - 15
cm. long, 2 - 4 cm. broad, glabrous, light green above, paler beneath, subcori-
aceous, margins often undulate, the costa raised, the secondary veins 20 - 30 per
side, anastomosing, obscure above, distinct beneath; petioles 0.8 - 1.8 cm. long, 1 -
2 mm. broad, glabrous. Flowers terminal or sometimes becoming lateral on
development of a leading shoot from an axillary bud, 30 - 70, in spreading um-
lliform cymes; peduncles subtended by an a
caducous, glabrous bud scales 1.0-1.7 cm. long; peduncles bearing branches
which are again divided to bear 1- 8 pedicels, branches and pedicels subtended
176 Man, N.Z. Fl. ed.2. p.496. 1925.
[Vol. 43
180 ANNALS OF THE MISSOURI BOTANICAL GARDEN
by caducous, glabrous, linear bracts about 1.5 mm. long; peduncles, branches, and
pedicels sparsely tomentose, accrescent in fruit. Sepals not imbricate, lanceolate,
1.5 - 3.0 mm. long, 0.5 - 1.0 mm. broad, glabrous; petals oblong, subacute to
obtuse, 5 - 7 mm. long, 1.5 - 2.0 mm. broad, free, spreading from the base, yellow;
stamens 3.0 - 5.5 mm. long; anthers sagittiform to elliptic-obovate, 1 - 2 mm. long,
up to 1 mm. broad. Pistil at anthesis slightly shorter or longer than the stamens;
ovary 1.5 - 3.0 mm. long, 1 - 2 mm. broad, tomentulose; style 1 - 3 mm. long;
stigma capitate and obscurely 2-lobed or truncate. Capsules ellipsoid, 2-, rarely
3-, valved, acuminate, 9 - 10 mm. long, 5 - 6 mm. broad, green to black, glabrate;
valves convex in transverse section, less than 1 mm. thick, coriaceous, with a
slightly thickened placenta bearing near the middle 2 - 4 flattened and peg-like
funicles up to 1 mm. long, dividing longitudinally into a coriaceous mesocarp,
which is shed on dehiscence, and a membranous endocarp covering the seeds; seeds
1 - 8, reddish-black to black, irregular.
Known from the North and South Islands of New Zealand. Flowers from
August to November.
New ZEALAND: NORTH AUCKLAND: Oruru, near Kaitaia, Matthews s.n. (AK);
Kaitaia, Matthews s.n. (AK, MO); Onawero Bay, Wangaroa, R. Cunningham s.n., 614
(K, as P. microcarpum R. Cunn.); in coastal forest remnants on Parua Bay Road, near
€
Whangarei, Carse s.n. (AK) — aroa, em Kirk s.n. (AK); Hender-
son, Cheeseman s.n. (AK, GH, US); Waitakere, Matthews s.m. (AK); same locality,
Mackie s.n. ( ; Anawhata, Ke north of Piha, Mackie s.m. (AK) subtropical rain
forest, Titirangi, Chapman s.n. (A); in second growth Agatbis australis KE forest,
Huia, Manukau Harbour, Wood s.n. (AK). THAMES: Kennedy Bay, Matthews s.n. (AK,
MO); Coromandel, Cheeseman s.n. (AK, BISH, GH, NSW); Table Mo eee Adams
s.n. (AK); Kaueranga, Adams s.n. (AK). VOLCANIC PLATEAU: : ixed forest,
Ohakune, MacDaniels p.599 (CU); Ruapehu, Attwood s.n. (AK); Ohakune River,
Crompton s.n. A. WELLINGTON: York Bay, Meebold € (BISH). NELSON:
Waimea woods, Monro 66 (K); Nelson, L. Travers s.n. (K); n Mountain, Mellor s.n.
eee MO); Foxhill, "Kirk Ag (BM). CANTERBURY: presqu ea de v prie Raoul s.n. (GH,
US); Akaroa, Raoul 83 (K); same locality, Belligny s.n. (GH); "Banks Peninsula,
Së 296 (GH, US); Lyttelton Hills, <i 4745 (BISH). WESTLAND: cd en Flat,
Grey City, R. H. [Helms] s.n. (BM, MO). oraco: Dunedin, Hector s.m. (K); near
Dunedin, Thomson s.n. (AK); bush, Dunedin Belt, Watt s.n. (K); Mokopeka, Meebold
5497 (BISH); in mixed forest, T dim Hills, Anderson 210 (A, F, K, MO, US). WITH-
OUT LOCALITY: A. Cun 2D s.n. e U); Bidwill s.n. (E); Brown Hr qM
G
Flowers which are probably female have 2-lobed capitate stigmas slightly
exserted, plump ovaries about 3 mm. long, 1.5 - 2.0 mm. broad, and sagittiform
anthers which appear to be abortive. Flowers which are probably male have
truncate stigmas slightly below the stamens, slender ovaries 1.5 - 2.0 mm. long,
about 1 mm. broad, and elliptic obovoid anthers which appear to be functional.
Godley has informed me (personal communication) that the species is diocious.
I am indebted to Mr. W. T. Stearn of the Bristish Museum of Natural History,
London, for the following note on the priority of Cunningham's names over those
of Putterlick.
1956]
COOPER—AUSTRALIAN AND NEW ZEALAND PITTOSPORUM 181
That part of Allan Cunningham's paper "Florae Insularum Novae Zelandiae pre-
cursor" which deals with Pittos sr a was published in Annals of Natural äm
Vol. 4, dated 1840, on the titlepage, vite Aloys Putterlick's eier bye
arum (Vienna) is dated AR and hence would appear to have prio
assumption J. do A. Franco in € x ` Broter Ao "s se (1951) m “replaced ge
enerally accepted name Pittospora enioides by e microcarpum Putterl.
This is not Ca only example of beans PB pore temas the two publications.
Hence it has seemed dude able to bees mp "ut rei Lag eet Cunning-
ham's paper (pages 106—111) was published i w.
dated ‘October 1839' both on Ay wrapper and on S 45a D ndon * publication it was
m
A
=
=
D
ER
a
received by the Linnean Society LB 14 Jan. 1840; “this suggests that it may
s end ds of the
other Viennese
source of information has been found. However, information abo ut Leem? pub-
1 i ig, t l-org erman
Thus the Fig Heit "x catalog of J. C. Hinr ichs IV ecidich. neuer
ished une
d December 1839. Mor recise evidence is give the weekly Allgemeine
w publication. Even allowing as much as six weeks for the announcement of its
Pinis to hs sent to Leipzig d Friederich Beck, Putterlick’s work would still be
antedated b ningham's. No evidence has been found, despite extensive search,
to Së Lë Putterlick's came um rlier. Hence it can be accepted as published
at the same time, in which event Cunningham" es 0 be retained, h
do y] Hooker T was the first to unite the species of Cunningham and
Putterlick (cf. Int. Code . Nom. 1952, art. 67), Cunningham's pa u
on oe 1839 can be accepted as having pron over Putterlick's Synopsis,
takin ember 1839 as the date of issue for this ccordance with the only
g N
definite etii ribi. ` Both procedures confirm the arabllehed nomenclature.
The type sheet at Kew bears three labels:
1. "Pomaderris microcarpa, Wangaroa, N. Zeal'd. R. Cunningham.”
2. “Pittosporum microcarpum. R.C. (non. Putt.). (P. eugenioides A.C.
Mss.). I gave it the Baron with this name. fl. non. vid. at Onewero Bay, Wanga-
roa, New Zealand, R. Cunningham. 1833.
3. "Taken from Herb. A. Cunningham, No. 614. P. umbellatum Gaert.
(Putt.). Pittosporum eugenioides A. Cunn. Fl. N. Z. 1-22. Ann. Nat. Hist.
V. iv. p.106. New Zealand. R. Cunningham 1833." The sheet is cited as "R.
Cunningham s.n. and 614" to identify it. Presumably "the Baron" is Baron von
Huegel whose collectings of Pittosporum were described by Putterlick.
Pittosporum eugenioides is commonly cultivated as a hedge-plant and orna-
mental tree in the United States, southern England, France and New Zealand, and
I have seen specimens from Portugal (Franco s.n. K). The plant distributed by
C. E Baker of California as P. undulatum, “one of the most valuable species of
the genus for garden and hedge planting” (Economic Plants of the World. No.
119.) is P. eugenioides. (A, CU, MASS, MO, NSW). A form with meen
leaves has been available in the trade for many years as var. variegata.
man??? recorded that the abundant fragrant flowers were formerly used by de
E
V Man, N.Z. Fl. ed.2. p.496. 1925.
[Vol. 43
182 ANNALS OF THE MISSOURI BOTANICAL GARDEN
Maoris who mixed them with fat and applied them to their bodies, and the essen-
tial oil present might be suitable for the perfume trade.
Vernacular names: Mapau, Tarata, Lemon Matipo, Lemonwood, Lemon Tree.
Tarata is the name preferred by modern writers.
EXCLUDED OR DOUBTFUL SPECIES
Pittosporum cect Dom. in Bibl. Bot. 22: 713. 1925. I have not seen
a specimen of this speci
Pittosporum nanum m Hook. in Comp. Bot. Mag. 1: 275. 1835 — Mariantbus
procumbens (Hook.) Benth. Fl. Austr. 1: 117. 1863.
Pittosporum procumbens Hook. in Comp. Bot. Mag. 1: 275. 1835. = Mari-
anthus procumbens (Hook.) Benth. Fl. Austr. 1: 117. 1863.
Pittosporum parviflorum Putterl in Lehm. Pl. Preiss. 1: 189. 1844-5. I
have not seen a specimen of this species but the drawings preserved at Vienna and
generously lent by the Director of the Naturhistorisches Museum do not represent
a species of Pittosporum.
INDEX TO ExsICCATAE
talicized numbers refer to the collectors’ numbers, s. n. (sine numero) to besseren
collections parenthetical numerals indicate the sb of the species or subspecies
the key.
Adams, J. s.n. (7); s. m. ipit n. (12); — F. M. & Simmonds, J. H. zg
s. n.
(18); s.m. (2); - (23);
(32). Beier y P s.n». (30
Allan, H. H s.m. (8a); s.n. (16); s.m. Baker, E RN (10); s . n. (26).
(22). Bancroft, T. L. s.n. (9); s. n. (25a); 5. f-
Allison, K. s.n. 0); a t. (6); s. m. (22): SO .n. (30).
Anderson, anks, J. & Solander, As C. s.m. (6); 5
Anderson, A. W. Ü THES 210 (32). jte s. n. (12); s.n. (22); s. n. (30).
Anderson, J. e£ al. 321 (26). Barnard, C. 109 24).
Andrews, C. 37 (9). Basedow, H. 84, 112, 159, 171 (9).
Archer, W. H. s. ». (10). Bate, M. 94 (26).
Armstrong, J. fy f n audin, e k a (24).
Aston, B. C. Seen s. n. (18 Bauer, F. s.7.
).
Atkinson, C. L "v. 24); s.m. (26 Baylis, G. T. S. 22707 (3); s. n. (12).
Atkinson, E. EAS HW OVI E Belales; RUD» :
Attwood, J. E. s.n. (8a); s n. (8b); s
= 11 5
a9; s.m. dies ». A085 de. (22); e^.
n. (23)3 s.n. (32). ee Dee E
B pS. w.». (23).
Backhouse, J. 669 (1); s.s. (26). d M
Baeunton, = = (t0); > (24) 5 SM. SE 9).
(26); s «tilo Beuzeville, W. A. a de. LI (10
Bailey, F. M. 182 (9); s.m. (24); s. Bidwill, d G 98 (5); 96 (20); =
I3, 20 (25a); s.m. (26); s. n. 25) (22); s 8 (24); 52 (29);
s. n. (30). (32).
1956]
Birch, GN:
y 53 (4).
——. 14982, 14996, 15242 (25a);
s: fi. (10).
de
5 3. (9)
014 (30).
Bligh, Wiles & Smith.
n. (10
S s. n. (24).
Ss (2); SER. (9) 525.8
30
iin L. ^ 2090, 2220 (252); s. n. (26);
Brass, L $ & White, C. T. 44 (9); 279
owl s. n. (26).
Brown, F t & E. D. W. e? (32);
Brown, R. 7 (9); 5448 (10); s
M raid (24); s (26); s. n., 5449 (30
wne, J. S. (9
SCH án, Fe?
Wa T s. n. (15).
Buddle, G. A. pe (3); 27204 (22).
Bullock, P:S. 3m O
Burbidge, N. T; 1962 (9); 3280, 3359 (10).
Calaby, Sa (9
).
Caley, G. s.m. (10); s.m. (24); 5.”
26).
Cambage, R. H. 3573, 4404 (9); 3821
Ca rafeld, LN s.m. (24); s. n. (26).
Campbell, W. D. s.n. (9).
rey, —. e».
Carne & Hudson.
e Fi. E ($2); "Si (8b); s
(12); s. m. (32).
Carse, H. & Matthews, H. B. s.n. (21).
Cecil, Lady. 2109 (10
Chapman; V. f LM (4); s. n. (32).
Cheel, E. s. s. (24).
Cheeseman, T. F. s.s. (3); s.n. (4); 5
(5): s (63: Ss ae s. n. (82);
sa (8b); s.m. (12); s (13); sn
(142); s.m. (14 (15); s.m.
(19); s (20); s Wie S-n., I2
(23); s. n. (32).
erus M. S. s.n. (24); s.m. 43747
29
Clinton, S. F. s.m. (10).
Cocks & E (18); 8551 (20);
421 (22); 6419 (23
Colclough, ^ iy 5 $
Colenso, W. s.m. (5); 206, 344 (6); $- Ms
COOPER—AUSTRALIAN AND NEW ZEALAND PITTOSPORUM 183
mber, H. F. 1550; 1833 (10).
(9).
Conservator of Forest. s.n.
Mo: (
Constable, E
ookson, I. s. n. (10).
Cooper, 6006, 36014 (8a); 24448,
C. oí
ut 35960 (1 p 36009 CAM ; 24318
Cranwell, n. (6);
$. fi. (2); 5. fi. wir
Crawford, G. A. s.m. ai
Crocker, R. L. s. 5. (9).
Crompton, W. s.n. (32).
Cunningham, A. s.n., 26, p e s. =
II, O15 (8a); s. n., 30, 3
143 (9); 33 (10); AL CH dg
618 (14a); s. n., ÓI, 616 (22); d 246,
Curtis, W. M.
Daemel, E. s.n. (30).
Daintree, . n. (24).
int
Dall, J. s. ». (31).
s.n. (24); s. m. (25b); s.m.
0).
(27)55. ". 3
D'Alton, St. E. 31 (9).
I son, 8 (24)
I5 :
e, H. s.n. (9); s. n. (10).
SE E. s.n. (8b).
$. H. (24) 5 Fed i 1299 (30).
Dietrich, A.
Doggrell, R. H. A.5
Dovey, L. G. 55 Pun 140, 153, IOIO
(30
Down OHNE s. n. (4).
Downing, R. Ehe
Drummond, la n., 31, 57, 76 (9 ).
Dunn, W. 129 Gei 136 (29).
Dwyer, J. s.n.
Eaton, H. G. s.n. (24)
aves, SCH, eie
Edgerly, . s.m (14a).
Edwards, E 2 m. (9); 5 10).
Everist, S. E ' 786, 3307 (9): pee (29).
Father, OR. (26).
Fawcett, H. C. s.n. (24)
Fenby, R. C. s.n. (24).
Field, H. C. s.m. (4).
Filtzalan, E. s. n., 77 (25a).
184
i hae oF e n. (9).
Flem
Ford, "N. E EX - Q0); s. n. (26).
Forest Officer. 20 ier
xO 35.1. A En
Fraser, R
Fraser, C. (9) s. Mss kg IIe (24).
Fraser, L. be Vickery, y: (24): 5. m.
26).
French, C Kë
French, C., Jr. sn (9); s. m. (10)
Froggatt, W. s Ké
Fullager, J. s.s. (2).
Fuller, R. H s. 2. (29).
€ F G k*-$);s5x:(10)) 65».
Gibbs Gë 6393 (10); 1296 (23).
Gilbert, e?
Godley, E. 2 ` 24963
, D. A. & Smith, ` ^ 142 (24).
Gray, Oo 3» UD,
n, W.H
Gregory Expedition. s.n. (9).
Gregson, I. s. 24
Griffiths, H. L. s.n.
Guilfoyle, W.R. Wa (24) s s.
Gunn, R. C. 5», 154 eer 650,
651, 651/1842 (10).
Haast, J. 69 (8
Hadle ey, H. $ we b (9); s. n. (24).
Seni 1591
n. (6).
+. s.n. (24).
H (27); 366 (30).
ëch? S.G. s.m. (10).
h
176 (5
6 (7); s.n.
n. infia ia s
pe ie 6 rt
Hectic; $ & Buchanan, J. 22 (8b).
Hedley, C. 13 (30).
Hedley, C
Hel
[Vol. 43
ANNALS OF THE MISSOURI BOTANICAL GARDEN
Holding, ios 82):
Holt, È 5
Hooker, J. D. 842, 844 (10); 350, 351
(12); 191, 352 (14a); s. n. (22).
Hort. Kew. s.n». (24); s. n. (30).
-n. (9).
Hubbard, C. E. 5024, 5572 (9); 3701,
Hm 4278, 4466, 4815 (24); 5225
Ps
ynes, P.
(13).
eweli, —.
66 (8b).
26567 ir s. n. (12); 26568
Irvine,
Jackson LS £9).
Tenis. T. Eo (23).
Jeban, LAS LM Ki s.n. (24);
. (25a) ; s. n., 106 (26
perg La AS & Conable: E.F £%
ae He (IR).
enny, 24);
erry, s. f. gt
Kershaw, J. A. s.n. (9)
eys, LL 29).
ripe. e $: n: m
King, >=" n. (2); s. n. (10).
Kirk, T. s.n., SC 623 (4); s. n., 25, 110,
123 (5); s.m, Ó56 (6); s. m., "86, 167,
194, 518 (7); 54, 83,85, s. n. (8a); 54
564, s.n. (8b); s. n., 6, 12); S. n.,
4^ 88, cado 521 dex S. N., BU 515, 538;
(14a); s. (15); ,7 (20); 5S. ts
130, 136 (22); $. f, e 362 s.n.
Koch, M. 7, 2983 (9).
La Bien A H H 1.9 (10).
` Laing, R. M ` 05); s.n. (19); s.
Gourlay, H. W. s”.
Lawren e n. (10
Tea tf, 3. L5» 6 s. n. (30)
Le Gra nat
i ern ,& Carse, H. s. n. (4)
nault, coh m. (9).
ec FE dw eu kx. (25),
immer,
s: Be m (91
Lloyd, R. C. & ANH J. 36309, 36310
(12).
Long, F. H. 355, 375, 1008 (10).
Longman, H. A. s.n. (9); s. n. (24).
1956]
COOPER—AUSTRALIAN AND NEW ZEALAND PITTOSPORUM 185
Longman, H. A. & White, C. T. s.n.
(24); s. n. (26).
Lord, E. E. ta (9); s. n. (10).
Lothian, N. sn. (9); s. n. (10).
Lowe, H. r4 (24
Lucas, ——. s.m. Qo; $5, (25a).
Lyall, ——. s.n. (8b).
Lyne, £f. is
MacArthur, W. s.n. A iim 6 (26).
MacDaniels, L. H. . (8a); 1225 (8b);
MacGillivray, J. I, 105 (30).
s. n.
Mackie, N (42); 5,16, (22); tet
2415.58 ui
Maclean, n. (29
MacMahon, J 205 (83); 35 (19); $. f.
(20); s. n. (22
Maiden, J. H E 3991 (10); s
(24): GG 7 Q9: Rs (2
"oen J.H. & rman, e) L cA dji
Maiden, rf H. & Cambage, R. H. s.n.
TA TEE oe
Matthews, H. B. 1822, s.n. (5); s
(2); SR (01: 5. e MEE I (8b); s. s.n.
. (14a); s. n., 1855
5.8, (13) ta: n. (16) r.a. (17).
Matthews, H. B., Leroy, E. & Carse, H.
s. n. (4).
Matthews, H. B. . pap D.
Matthews, H. J. (8b).
Matthews, R. H. s.n. en s.n. SEN
Matthews, R. H. & Carse, H. s.n. (143);
sn. (17).
McComish, J. D. 136, 136« (2).
McKenzie, C. s.n. (29)
s, n. (19);
McWilliams, .- Sci. (26).
Meebold, A. 5389 m ; perta 18280,
4823, 5 4730, 5410,
5564 (6); 4096,
5845, 18277, 18282, 18283 (8a); 4442;
18276 (n 5289 (22); 4745, 5497»
18278
Mellor, T. M 27212 v s. n. (32).
Menzel, O. E
Merrah, E. s.n. (10).
Michael, N. 9, 30, E (25a); 973, 2024
(29) ; s. n., 1484 (3
Michie, R. H. dn SC
Milligan, E : n. (10).
' Mitchell, s. n., 228, 229, 273, (9)-
Molesworth, B. 23320 (5); s.m.
(7); s. n. a»: AC (13).
Monro, D
Moore, C ed 23 (2); jud (24); s. 5
(26); s 29).
yer L. B. & Cranwell, L. M. s.n. 0
(6); s.m. (8a); s." (8b); s.
(12) s.n. (16)3 s. m. (18); s. m. eg
n. (22); s; m. (23).
00 pe, Spencer, e (9).
Moran, R. 2
orris, A. s.m. SCH 1441, (10).
orrison, A. T aie 14338 (9).
Morrison, EN 39 (20).
Mould iG. We 26).
Mueller, F. s.n. (1 , 16
); s.n. (5);
(9); s.n., ILIQ (10);1 Ew QA ^ n.
(26); s. n. (28); s. m. (29); s.m. (30).
Murphy, 9 (26).
eal, M. C. ro (8a).
SE J. 35 (är 34 (29).
Oldfield, A. s.n. (9).
Oliver, W. R. B. s. n. (6).
Olsen, IO (5); E (12)
Olsen, A. M. s.n. (10
Osborne, ——. s.n. (5).
O'Shanesy, P. A. 87/9 (29); 1824 (30).
A. s " ).
Paddison,
n 95. (1); s. 9. (10).
RS. (093 6417 (5); 6418 (7);
(8a); Era i R 4/4 (8b); s.n.
SCH s.n. (18); s. m. (20); s”. C22);
Petrie b. E Matthews, H. B. s.n. (8b);
eue; Y. En 120 (24).
Powell, H. 26437 (142).
Preis, L. 1297 (9).
Prime, H. s. ii (24).
. n. (10).
Rodway, F. A. 871 , 875, 912, 1204, 1954
(24); 361, Sap, 874,1 1690 (26).
od L: kw (10):
(25a)
R. . Inst. Hort.
Rupp, H. MR 5. #. "Qu y er,
186
Sainsbury, G. O. K. s. »., 164, 622, 623,
7);
624, 708, 709, 710, 711 (6); s.n. (1
s. n. (20); 15 (22).
St. John, P. R. H ve (25).
Sayer, W. s.n. (10); s.n., 186 (25a);
Sn(30):
Schneider, s £950) s:
Schomburgk, s. n. (9) ; s. n. (26)
chultz, 26, 758 (28)
Searcy, —— (9)
Shakespeare, 25
Shirley, J. s.n. (26); s. n. (29)
ieber, F. W. 2, 26).
Siemssen, T. s.n. SE 97 (26).
Simmonds, v (24); s.m. (26);
Ed M.
Simon, M. H. 37 (29)
Simpson, A. s.n. (10).
5 (5); 5. 8. TEE E 5
(1
Sledge, V. A. 266 (8b); 74 (22).
mith, C. s.n.
ict L. S. & Webb, E J. 3612 (29).
Staer, J. s. n. (10).
Steel, T. s $
Steenbohm, L. s.n. (24)
Stirling, J. sn (10).
uart s.n. (10); s.m. (24); s.m.
(26).
Sturt, n. (9).
Swain, E. H. S. s.n. (26).
Tanner, H. 8 (24); $. n. (29).
Taylor, T. W. s. 2. (9).
Thon. E. 16 (9
Thomson, G. M. s.n. (32)
Thozet, A. s.n. (29)
Toogood, B. 9 GEI
Ko ez Ké (6); 669, 670 (8a);
- (18); 7s (19).
Tieren: L
eN
5.
n (3
Travers, W. T. E n. (8b); SC (18).
[Vol. 43
ANNALS OF THE MISSOURI BOTANICAL GARDEN
Turbott, E.G. 23023 e n. (12).
Turbott, E. G. & Bell, L. C. 22925 (3).
159, 816 (8b); s.n. (12);
(16); 156, s.t. (18); Ej TOR
804 (22); 793 (23).
Turner, E
Vachell, F. H. s.n
er —
EA
1961 (10); 46, 44P, 406
(26
Vickers; E ose (9): S (24).
Walker, E. H. 4316 (8b); 5364 (12);
526
Wall, A. s.m. (19).
Walter, C. ee (9); 5.8. (1077 MB
(29).
Waterhouse, F. s.n. (9)
att, M. )
Watts; Wo. W: s.m. (29).
eber, (26).
Weindorfer, G. s.n. (10)
Whaite, T. H. 695 (24).
Whetter, : o (12)
White, C. A. s.m. (9).
White, C. T. va 9032 (9); 6406 (10);
pines (11); 1370, s.n., 11474 (24);
1255, 10618 (25a); 3570, 6179, 9026,
s. n. (26); 8142, 8974 (27); s. n., 1146,
1903, 6045, 9522, 12507 (29); 10560,
12216 (30).
White, C. T. & Brass, L. J. 227 (252).
Wien s.n. (5); s. n. (8a); s.m. (9)
; $. ft. (14a) ; s. n. (22); 5%
(2 6).
Williams, G. T.
Williamson, H. B. s.n. (9).
27210 (4); s" (82); 4
s. n. (4).
Wood, K.
(22): $056 (23) A OX.
Woolls, W. s.m. (9).
Young, J. E. s. #. (29)
Zotov, V. D. s.n. (86).
1956]
COOPER—AUSTRALIAN AND NEW ZEALAND PITTOSPORUM 187
INDEX TO VERNACULAR NAMES
Italicized words are aboriginal (Australian) or Maori (New Zealand) ; numerals refer
to the species or subspecies
Apricot Tree, 9
i426
KEE leaved roo. 22
e, 9
e chef Laurel, 29
n's Blood Pittosporum, 10
Engraver's Wood, 26
Genoa Pittosporum, 24
Haekaro, 12
Karo, 5
Kirk's Pittosporum, 23
obubu, 8a
Lemon Matipo, 32
Lemon Tree, 32
Lemonwood, 32
Locket Bush, 9
Lord dus Island's Hedge Laurel, 2
number recognized in this thesis.
Mapaurik.
Mountain pa 8b
mie.
New South Wales Mock OH. 26
Poison-berry Tree, 9
Quinine Tree, 9
Rhombus- leaved ege Laurel, 29
rg Wood, 1
wild Yellow Jasmine, 24
Willow
Lei Brisbane Laurel, 24
INDEX OF SCIENTIFIC NAMES
combinations are in bold face type; synonyms are in italics, and previously
New
published Geet accepted names are in ordinary
Marianthus procumbens e 182
Pittosboroides — cs 121
umbellata p P yd
meea Dor RI
acioi as E OU
angustifolium A rn AA
alum o EDO
bi ee EE _.. 144
BIVALVAE des ih n 1
bidwillian E ER
Beggen Ee EE
these a MN
collicarpen ii 10%
colensoi do PE
var. fasciculatum -.——————— 135
cornifolam o EE LT
crassifolium EE Y dí
var. uod oe qeu
var. $ a A ss MAS
ar. "variegatum" .— — —— — 130
crassifolium | ——————— 130
renulat A DI
allii — rs f dept uique 27 8
iscolor — EE MM
divaricat PR A e d
eege — e 133
elegans | ———— ——————————— i?
elliptic SE ee EE
ssp. ellipticum — ————————— 125
var. "` LL M 125
ssp. ovatum .————————7— 127
var. ovatum een 127
134
188
intermedium
lanceolatum
ligustrifolium ` —
matthewsii
mayi
y
melanospermum
microcarpum
icum
n
nigrescens
gei
. “Silver Queen”
sip
wire itane
var. kaitaiaensis
m
var. gilliesianum
ssp. major
var. major
J
ssp. pimeleoides
eflexum
var. reflexum
pimeleoides
Pom
Trichilia monopbylla _.
[Vol. 43
ANNALS OF THE MISSOURI BOTANICAL GARDEN
rocumbens —-
var. jomenioniss - o A
rhombifolium Adel n
var. E EE E
A A
transluc
TRIVALVAE d oh es BI LONGAS RUE
te 4 a sc iN Sabre
Seit Gen EE x
var. cordabim . — ——— ee
atum
"riens
waiforem LLL ee
Venulosum Loa ee uud
Werticilata. s S s sss SD
virgatum
var. crataegifolia
var. seratum
var. nnuadlum = —————————
viride _ Eua sats
giel
derris microcarpa een
A aa
THE CULTIVATED BEANS OF THE PREHISTORIC SOUTHWEST*
LAWRENCE KAPLAN**
Formerly Associate Curator of tbe Museum, Missouri Botanical Garden
Beans, corn, and squash were the basic cultivated plants of pre-Columbian
North America. There is no monograph on prehistoric beans, and the history of
their distribution and a description of their variations have not been published.
Although prehistoric collections are few as compared with those of corn, a study
of available material can contribute significantly to the history of agriculture in
North America.
Apart from horticultural, agricultural, and other practices which create or
modify ecological conditions so that domesticated plants and weeds might survive,
man exercises conscious and unconscious varietal selection. Patterns of selection
are set by culture-based criteria, and attitudes towards plants and selections are
made within the limits of plants available and primary factors such as geography
and climate.
The roles of plant geography, ecology, and of human culture in shaping the
species composition and distribution of beans are taken up in the first part of this
paper. The factors which determine the kinds of plant materials available for this
sort of study are also discussed. Descriptions of the materials and their distributions
in the Southwest are subsequently given, with discussions of the part they play in
the history of cultivation and variation in beans.
Acknowledgments:—The author wishes to express his gratitude to the Chicago
Natural History Museum and the Missouri Botanical Garden for the use of their
research facilities, and to Dr. Hugh C. Cutler, Associate Director of the Missouri
Botanical Garden, who proposed this study and contributed many useful sug-
gestions. "Travel in the Southwest was made possible by Dr. Cutler and the
University of Chicago where the author was enrolled as a graduate student.
Grants from the University of Chicago and the Chicago Natural History Museum
aided field work in Mexico. This study could not have been completed without
the cooperation of the institutions (listed in the explanation of Table VII) which
made their collections available.
TAXONOMY AND DISTRIBUTION OF CULTIVATED BEANS
For purposes of this study beans are defined as the seeds of cultivated plants
belonging to the genus Phaseolus. Bentham (1841) placed this genus with other
trifoliolate genera in the papilionaceous tribe, PHASEOLINAE, sub-tribe EUPHASE-
OLEAE, but separated it from other genera of the sub-tribe by the coiled keel
characteristic of Phaseolus flowers.
* An at the Wope? wi Chicago, pena Natural History Museum
and the Missouri Botanical Garden, and submitted as a thesis in partial fulfillment of the require
TOn for the degree of Doctor of Tlaky in = ae t of B ës University of Chicag
** Department of Botany, University of Chica (189)
investigation carried out
[Vol. 43
190 ANNALS OF THE MISSOURI BOTANICAL GARDEN
The genus Phaseolus contains approximately 180 species (Ditmer e£ al., 1937)
distributed throughout the tropics and warm temperate regions of the World
(Bailey, 1949). Distinctions between the Old and New World cultivated species
are several, indicating long separation and independent evolution as cultigens
(See Table I).
TABLE I
DISTINCTIONS BETWEEN OLD AND NEW WORLD CULTIVATED BEANS
Old World New World Authority
Flower color Usually yellow Red, rose, violet, white | Bailey, 1949,
(may be yellow-tinged) | Piper, 1926
Seed size Small Large (teparies small)
Testa pattern Little variation Highly varied
s Narrow r : Hedrick, 1931
Flower morphology Left keel petal with | Left keel petal without | Piper, 1926
horn-like process | horn-like process
Growth habit Annual Annual or perennial Piper, 1926
Pathology
Cercospora cruent Susceptible Resistant Piper, 1926
Gloeosporium lindemutbianum| Resistant Susceptible
Another distinguishing character which might be added is the geographical
distribution of species included in the genus. According to Ditmer (Ditmer ef al.,
1937) 126 species of Phaseolus occur in the Americas; 54 are natives of South
Asia and East Africa; only two species are indigenous to Australia, and but one
assigned to Europe.
As a result of the facile development of cool-temperature ecotypes and their
acceptability as food, the range of the cultivated beans of the New World has been
extended to the regions of Asia and Europe for which the Asiatic species have
proven too thermophilic (Ditmer eż al., 1937). Photoperiodic adaptability (Allard
and Zaumeyer, 1944) has apparently also contributed to the wide latitudes avail-
able to cultivation of the New World types. The Asiatic species, on the other
hand, have not proved as successful for dry edible beans in the Americas as the
highly productive and familiar common and lima beans. The mung bean, Phase-
olus aureus, used in the sprouted form in oriental cookery, is the only Old World
bean to have been grown in quantity in the United States. These beans were
cultivated in Oklahoma to supply domestic needs for sprouting beans during the
1940's when Asiatic supplies were cut off.
Phaseolus vulgaris, the common garden bean, or kidney bean of Anglo-America,
is frequently confused with the "cow pea” or “black-eyed bean," Vigna sinensis
of Asiatic origin, which also provides green immature pods and dry seeds used for
human food. The snap beans, both green and wax-podded, and field beans also
1956]
KAPLAN—PREHISTORIC SOUTHWESTERN BEANS 191
may usually be referred to P. vulgaris. In Mexico they are known as the frijol, in
Colombia as the frisol, the names of Spanish origin having largely replaced the
indigenous terms in these countries and elsewhere in Latin America. The numerous
Spanish or English horticultural varieties are usually multinominal and are based
upon seed characters such as color, color pattern, shape, supposed derivation of
original seed (as Mexican Red, Hidatsa Red), use (as Navy Bean), productivity,
growth habit, precocity, and the like.
CENTERS OF DOMESTICATION OF AMERICAN BEANS
The cultivated beans which were domesticated in the New World are repre-
sented by four species. A brief history of their origins is given below.
Phaseolus vulgaris L. (Sp. Pl. 723. 1753), Kidney or Common Bean: This
highly variable species was considered by Linnaeus to be Asiatic in origin. De
Candolle (1886) was convinced that the common beans are native to the New World
but, because he was uncertain of the validity of archaeological evidence, placed
them among plants of doubtful origin. The use of archaeological material in the
demonstration of the New World origin of common beans is indicated below.
Ivanov, according to Bukasov (1930), assigned the common beans to the Mexican-
Guatemalan highlands region on the basis of varietal diversity.
McBryde (1945) and Burkart (1943) reported Phaseolus species which they
considered were non-domesticated forms of the cultivated common bean. Collec-
tions of similar kinds from the Mexican plateau and Guatemala also exist, the
descriptions of which have not as yet been published. The only such material
Which has been investigated experimentally is that of Burkart and Brücher (1953).
They found beans from Honduras and Argentina to be sexually compatible with
cultivated varieties of P. vulgaris, but not with those of P. coccineus or P. lunatus.
The floral characters were similar to those of P. vulgaris. The Argentinian collec-
tions were from uninhabited (at the present time) high mountain valleys and
were not thought to be escaped forms. The chromosome number in root tips of
both the Central and South American material was 22, as in other members of
the tribe. Burkart proposed the following nomenclature for Phaseolus vulgaris as
now underst
P. vulgaris L. sensu amplissimo
P. vulgaris subsp. aborigineus Burkart
P. vulgaris subsp. cultigenus Burkart (all cultivated varieties)
It would seem likely that a number of the so-called "wild P. vulgaris” types might
have had an influence on what are known as the cultivated common beans by way
of hybridization.
There is little doubt that the common beans were domesticated in the New
World from a form or forms not yet specifically known, and it is probable that
the highlands of Mexico and Central America which show the greatest diversity
of cultivated forms (Ditmer et al., 1937) were the center from which domesti-
; [Vol. 43
192 ANNALS OF THE MISSOURI BOTANICAL GARDEN
cated varieties spread first throughout agricultural Indian-America, and later to
the other continents.
Phaseolus lunatus L. (Sp. PI. 724. 1753), Lima Bean:—Ames (1939) cited
Bentham as having referred the origin of the lima bean to Brazil but stated that
it is not known there in the wild state. Mackie (1943) concluded that Guatemala,
as the center of varietal diversity and the region from which a wild bean (con-
sidered by him to be P. lunatus) has been collected, is the place in which lima beans
were first domesticated.
Phaseolus coccineus L. (Sp. Pl. 724. 1753), Runner Bean or Scarlet Runner
Bean:—Runner beans are not definitely known from any reliably dated archae-
ological site of pre-Hispanic America. The scarlet runners which have been found
in the Southwest are cultivated on a small scale by the Hopi or have been found
in caves and reported as viable. Whiting (1950) made reference to the oral
traditions concerning these supposedly ancient beans; when planted, they were
supposed to have sprouted and produced crops of the edible dry seeds. The author
has heard from individuals in the Southwest that progeny of seeds taken from old
cave habitations are of the large white type (R 1, Tables IV and V) and are said,
when cooked, “to taste like mashed potatoes.” It is more than probable that the
limits of duration of viability would preclude any considerable age being assigned
to these collections.
The reports of Ivanov (Bukasov, 1930) and of Ditmer (1937) of distribution
of P. coccineus limited to southern Mexico and Guatemala, and other reports (cit
below) of the occurrence of P. coccineus as a weed in fields of cultivated common
beans in the same region indicate that this region is a center of domestication for
this species.
Phaseolus acutifolius A. Gray (Pl. Wright. 1:43. 1852), Tepary Bean:—The
cultivated forms of this species, described as P. acutifolius var. latifolius (Freeman,
1912), are known mainly from two centers: one is that of the Indian populations
of southern Arizona (Freeman, op. cit.); the other, found by Russian botanists, is
the Pacific Coast of Chiapas in southern Mexico (Bukasov, of. cit.). Although
Freeman proposed the Sonoran Desert region as the probable center of domestica-
tion of the tepary bean, Bukasov stated only that Mexico is the center of origin.
The question of origin of the tepary bean will be discussed again in this paper.
The greatest diversity among the cultivated beans was found by Russian in-
vestigators (Bukasov, op. cit., p. 510) in southern Mexico and Guatemala. They
thought that the highlands of these regions saw the earliest domestication among
the Maya, with subsequent adoption by lowland peoples, along with corn and other
cultigens. Bukasov, according to Ditmer (1937), maintained that contact be-
tween Mexico and Peru did not come until a relatively late date and that on the
arid coast of Peru a second center of origin for large-seeded, cultivated P. vulgaris
was established. Prehistoric common beans of the Ica valley (Table VII, collec-
/
1956]
KAPLAN—PREHISTORIC SOUTHWESTERN BEANS 193
tion numbers 225-231) are larger than those found in archaeological sites of other,
non-Peruvian cultures; but so are the lima beans, and Cuzco maize (collections of
Hugh C. Cutler) is larger-kerneled than any other race. It would seem at least
possible that ancient Peruvians had a predilection for large-seeded crops and,
through selection, developed large-seededness to the extent of the genetic poten-
tiality of each variety.
Wild species of Phaseolus are abundant throughout the warmer regions of the
Americas, especially North America (Piper, 1926). Coincident with this distri-
bution may be found not only the greatest number of forms of the common bean,
but also the ranges of species which have likely had influence in building the
polymorphic character of P. vulgaris.
Bukasov (1930) and others reported the common occurrence of Phaseolus
coccineus as a tolerated, useful weed in plantings of the common bean in Chiapas
—a situation which may be expected to persist in Guatemala. This species,
although observed by the author while in Mexico in 1954 in abundance in the
markets of towns situated at higher elevations in Chiapas, is not seen in the lowland
tierra caliente, The “tropical black complex,” indicated by Freytag (unpublished
thesis) to be the source of variability of the common bean in Central America,
certainly extends its influence at least into the tropical lowlands of Mexico. In
other regions of ancient bean cultivation, such as the arid Southwest, other hybrid-
izing species probably have had little influence on the common bean because of
the low rate of outbreeding in P. vulgaris under arid conditions.
For a review of the systematics and synonymy of the cultivated beans, as well
as their recorded history and appearance in the herbals, the reader is referred to
Van Eseltine's discussion in “The Vegetables of New York” (Hedrick, 1931, pp.
3-16) and to Burkart’s (1943) diagrams of seed types of all cultivated bean
species,
GEOGRAPHICAL LIMITS OF THE SOUTHWEST
Although of importance to the present problem, the precise geographical and
ethnic limits of the Southwest region are not easily determined. The question of
how to define the Southwest, whether in terms of distribution of subsistence
patterns, geography, ecology, or other criteria remains a matter for open discussion
among anthropologists and other specialists. This is evidenced by Kirchhoff's
recent paper (1954) and the comments of Kroeber, Beals, and Sauer. Kirchhoff
considers the '* . . . Greater Southwest areally to include Central, Southern and Baja
California, the Great Basin, Arizona, New Mexico, Southern Coastal Texas and
Northern Mexico south to the Sinaloa and Panuco rivers.” Ecologically and
culturally he distinguishes arid America with a gathering population from coexist-
ing Oasis America and farming peoples.
The difficulties in presenting a definitive boundary scheme for this region are
discussed by C. Sauer following Kirchhoff's article. He reviews the vast areas of
the Greater Southwest for which the archaeological record remains virtually un-
examined and for which there is still little information on migrations of peoples,
[Vol. 43
194 ANNALS OF THE MISSOURI BOTANICAL GARDEN
diffusions of agricultural techniques and crops, and cultural data, including
descriptive and analytic linguistic materials. He suggests that Kirchhoff's classi-
fication is essentially an ecological one, based upon geographic considerations, and
that only more information will enable classification in cultural terms and reveal
the relationships underlying the culture history of the region.
For purposes of this paper, the Southwest region includes the present states of
Arizona and New Mexico, and parts of Utah, Colorado, Texas, Nevada, California,
and northern Mexico.
SOUTHWESTERN CULTURE HISTORY AND ENVIRONMENT
Geology and archaeology give evidence of the presence of non-sedentary human
cultures in the Southwest at least 15,000 years ago. Some time before 300 B.C.
differentiations appear in the archaeological record which show that these hunting
and gathering peoples had begun to develop along somewhat divergent cultural
lines.
In southern Arizona, the most arid portion of the Southwest, the prehistoric
cultural development has been termed the "Hohokam." It is not clear at what
time agriculture became established in this region, nor is it known whether it was
preceded or followed by pottery. However, agriculture was well established by
300 B. C., and irrigation reached a high development among the Hohokam. The
contemporary Pima and Papago tribes live in the area once inhabited by the
Hohokam.
Peoples living to the east and northeast of the Hohokam, in what is now New
Mexico, have been called the “Mogollon,” and their region included the best-
watered and most mountainous sections of the Southwest. Corn and squash culti-
vation was established among the Mogollon as early as 2,000 B. C., and pottery
did not appear until considerably later. The Indians now occupying the Mogollon
area are of the Apache group.
In northern Arizona and the Four-Corners region of Arizona, Colorado, New
Mexico, and Utah emerged the “Anasazi” culture. The Anasazi of pre-pottery
times have been called the “Basketmakers,” and their successors, who made
pottery, have had their culture history divided into four "Pueblo" periods. Pottery
came to the Anasazi later than to the other groups; corn and squash agriculture
were pre-ceramic among the Basketmakers and appear not to have been derived
directly from the Mogollon or Hohokam. The Hopi, the Zuni, and Rio Grande
Pueblos continue the Anasazi Pueblo tradition. The Navajo, who, like the related
Apaches, were late arrivals to the Southwest, also occupy much of the Anasazi area.
Entering the Pueblo Southwest in the 16th century, the Spaniards found a few
concentrations of farming peoples inhabiting a large area which formerly supported
a more disperse population. These concentrations probably began with the droughts
of the 13th century and the destruction of agricultural land as a result of arroyo
cutting when more reliable water sources, such as those of the seeps and springs of
the Hopi mesas, were sought. Religious and political domination by the Spaniards
1956]
KAPLAN—PREHISTORIC SOUTHWESTERN BEANS 195
brought about rebellions and migrations of refugees which augmented the normal
contact among native peoples and certainly led to the introduction and exchange
of crop varieties.
Contemporary Southwest Indian peoples, who are to varying degrees the heirs
and continuers of the cultures of pre-Hispanic and pre-Anglo-American times,
continue the cultivation of many ancient crops in addition to relatively recent ones.
Use and Preparation of Beans:—It has been pointed out that, with the possible
exception of the tepary, the centers of origin of American beans must be sought
outside the area of prehistoric occupation with which this paper is concerned. It
follows that beans entered this region as cultigens by diffusion and with the plants
came modes of use and cultivation. i
Carter has observed (1945) that boiling of dry beans without preliminary
soaking is the common practice among native southwestern peoples in modern
times. This practice has been similarly observed in many parts of Mexico by the
author and is probably a very early form of preparation. Laborious preparation
of foods is common in modern non-industrialized societies; witness the stone-
grinding of corn in tortilla-making and the grating and leaching of mandioca.
But these are basic starch crops. Were beans important or favored enough to
warrant the protracted stone-boiling in woven baskets and high fuel consumption
necessary in pre-pottery times?
Carter (1945, p. 75) thought it likely that, if bean culture did begin in pre-
pottery times, the use was probably as a green vegetable. It is worth while here
to consider the possible uses of beans by prehistoric Indian groups.
The green pods of common beans have been used among native peoples of
Indian America; in Mexico they are called ejote, a word of Nahuatl origin (Santa-
maria, 1942). Pod beans as used by ancient Americans, however, were certainly
not like the string or snap bean of today. The earliest string beans of low fiber
century (Hedrick, 1931). No stringless bean is known to have been contributed
by Indian agriculture.
The use of the immature fruits of the common bean is described by Waugh
(1906) for the Iroquois of Canada, in which the boiled pods were chewed and the
fibrous remains ejected. That this pattern of consumption, or one like it, might
have been expected among the southwestern peoples is evident from the finds o
masticated vegetable fiber masses in many archaeological sites of the region. Most
of these so-called quids have been described as agave or mescal quids, but for the
most part they have been reported by persons unfamiliar with the identification
of fragmentary plant remains. However, remains of this type have been exam-
ined by Cutler (Martin e£ al., 1952) for Tularosa Cave and by Kaplan (unpub-
lished) for Cordova Cave, two Mogollon sites with pre-pottery remains, and in
Tularosa Cave, with beans associated with the pre-pottery remains. Only one of
the hundreds of fibrous wads proved to have been a bean pod. There is no reason
[Vol. 43
196 ANNALS OF THE MISSOURI BOTANICAL GARDEN
to believe that masticated indigestible fibers from bean pods would not have been
expectorated on the cave floor as were the wads of agave, yucca, cactus stems, and
certain unidentified fruits, probably asclepiads. The conclusion that bean pods in
the early levels of these sites were not extensively used is inescapable. The possi-
bility of removal of the "strings" before cooking, however, cannot be excluded.
Shelling and cooking of immature seeds would not be more difficult than the
preparation gf the immature pods in a pre-pottery technology.
That peoples of Indian America relied upon beans rather than animal. foods as
a source of protein is well known. Linton (1940) discussed the role of a balanced
diet in culture history and pointed out that in Indian America starch and protein
crops (beans) have been used, in contrast to other parts of the world where starch
crops and animal food, or other kinds of legumes, have provided the balance. D. B.
Jones e£ al. (1938) indicated the complementary nature of bean and corn amino
acids in providing dietary protein needs.
The adequacy of beans as a protein source depends ultimately upon their
utilization and preparation. The protein value of immature beans as given by
Chatfield and Adams (1940) is highest for (almost mature) green-shell lima
beans, 7.5 per cent of the total weight, while in green snap beans there is less than
3 per cent protein. For several commercial North American mature dry-shell
varieties tested, protein was 22 per cent of the total. In addition, the green
products are quite low in fuel value while the mature seeds are high. When mature
and dried, beans are at the most useful stage, for then they can be employed by an
otherwise protein-poor economy and also can be relatively easily stored.
Soils.—Tolerance of a wide variety of soils, if they are well-drained, is char-
acteristic of cultivated American beans in general. The species differ, however, in
response to alkali, acid, and saline soils, and these different responses have probably
influenced the distribution of beans in the Southwest. Acid soils are not encoun-
tered in the arable Southwestern lands, while alkaline soils are more common in
poorly drained alluvial lands of the southern and southwestern parts of the region
(U. S. Dept. Agr. Yearbook, 1938, p. 110). The irrigation practices of the
ancient Hohokam may well have contributed to alkalinity in these soils. Hendry
(1918) indicated the low tolerance of commercial P. vulgaris varieties to alkalinity
and considered P. acutifolius varieties as moderately tolerant.
Saline soils (.02 per cent NaCl) affect bean growth (Wadleigh e£ al., 1943) by
increasing the severity of drought conditions. The superior drought resistance of
tepary beans is well known, while common beans are severely affected by a rel-
atively few days of wilting. The small-seeded limas or sieva beans are character-
istically more resistant to aridity and high-temperatures than are the large-seeded
limas (Mackie, 1943). That salinity has been a factor in Indian agriculture, at
least in the area of the Hohokam, is attested to by the Pima practice of tasting
soil preparatory to planting (Heintzelman in Emory, 1859, p. 112). If salt were
detected, the soil was considered unfit for agriculture. It is probable that salinity
1956]
KAPLAN—PREHISTORIC SOUTHWESTERN BEANS 197
below the threshhold of detection by human taste is sufficient to influence plant
growth by altering water relations under drought. Many of the sites selected as
acceptable could have been salty enough to intensify the effect of arid conditions.
Irrigation.—Emory (1859), in his survey of the Mexican-U. S. boundary,
noted that little agriculture could be engaged in without water supplied in addition
to the seasonal precipitation. To meet the water requirements the Pima and
Papago (Castetter and Bell, 1942) have employed irrigation systems, drawing
water from the Gila River as did the earlier Hohokam peoples. Halseth (1936),
by aerial survey, found about 125 miles of ancient irrigation canals in the Salt
River Valley and other canals about half that extent in the Gila Valley. Some of
the canals are more than ten miles long, and all ruins associated with them were
Hohokam in culture. The very high temperatures and high evaporation rates
characteristic of southern Arizona may be supposed to have encouraged the cultiva-
tion of teparies rather than other beans despite irrigation and flood-water farming.
Extensive irrigation by canals has not been shown to be characteristic of farm-
ing among the Basketmaker and Pueblo peoples to the north, and the author is
unaware of any irrigation in use in prehistoric Mogollon cultures. Some irrigation
is practiced by the Hopi of Moencopi according to Whiting (1950, p. 10), who
implies that this was learned from the Mormons. The terraced gardens observed
by the author at the Hopi village of Hotevilla were irrigated by hand, and Whiting
indicates that this is the general practice among the Hopi. According to him,
neither corn nor bush-type beans planted in the same or separate fields in sandy
washes and alluvial soils by the Hopi and the Navajo are irrigated except by
occasional flood water.
Effect of Daylengtb.—Carter (1945) has stated that the majority of native
Southwest beans are bush types, and the observations of Whiting (1950) and
others that field-grown beans are planted apart from corn would substantiate this.
Allard and Zaumeyer (1944) have studied the photoperiodic responses of many
strains of cultivated species of Phaseolus and found that the bush types of P. vul-
garis are mostly day-neutral in their flowering and vegetative responses. Day-
lengths used in their study were comparable in range to those of the latitudes of
the Southwest. The semi-determinate or semi-pole types studied by them exhibited
more positive reactions to photoperiod than the bush types. The twining habit of
the semi-pole varieties was emphasized by long days and accompanied by a delay
_in flowering, while shorter-day treatment was correlated with earliness and the
determinate habit. Thus varieties which were determinate at lower latitudes might
well have taken on vining characters when introduced to the Southwest. It is
apparently precisely this factor which prevents the semi-pole California Pink and
Pinto varieties from entering the northern part of the western range where the
delay in flowering would mean injury because of early frosts.
While many of the pole types were shown by Allard and Zaumeyer (1944)
to be short-day, they proved to be mostly day-neutral for flowering. This type
[Vol. 43
198 ANNALS OF THE MISSOURI BOTANICAL GARDEN
remained twining at all daylengths so that the varieties would be expected to be
twining even under conditions of short days when the semi-pole would be de-
terminate. The lima beans tested were all day-neutral while runner beans were
long-day. Allard and Zaumeyer did not include tepary beans in their work.
Planting dates to take advantage of the limited precipitation and of flood
waters, or to avoid frost periods, might have been adjusted without difficulties
arising from new photoperiodic relations. The diffusion of some bean varieties
within the Southwest may have been impeded by photoperiodic reactions, but there
is strong evidence from experimental and field studies that this environmental
factor does not restrict bean distribution within this area.
Vegetal Remains of the Prehistoric Southwest.—Much of the Southwest region
is arid, and many of the ancient peoples lived in caves, shelters, or houses which
have remained relatively dry. For this reason perishable materials have been better
preserved than in more humid areas such as in the Mississippi Valley and other parts
of the eastern United States. Even in less-protected Southwestern ruins fires often
occurred at or soon after the time of abandonment, frequently charring and
preserving vegetal materials.
Collections of prehistoric vegetal materials have been studied and reported upon
by various authors. Some of these studies have only provided descriptions of the
materials found at particular sites; others have related their descriptions to other
collections and other aspects of Southwestern culture. Among the latter, Carter's
(1945) studies have been most comprehensive for remains of cultivated plants.
He theorized that the division between Hohokam and Anasazi corn and a change
in corn types is a result of Mexican and later Eastern (United States) influences.
Some question exists as to the extent of the Eastern influences in the corn of the
prehistoric Southwest. The steady reduction of the percentage of maize cobs
with high row-numbers reported by Cutler (Martin e£ al, 1952) for the more
recent periods (especially 200—400 A. D.) of Tularosa Cave appears to be a pattern
for the entire Southwest. Nickerson has recently (1954) provided evidence on
the basis of anatomical characters to show the similarity of maize from widely
distributed Southwestern sites.
Beans recovered from archaeological sites have received the attention of bot-
anists and students of human culture. Farlier studies were directed toward
demonstrating their place of origin by their presence in undisputed pre-Columbian
ruins. Jones (1952) has adequately discussed the historical phase of prehistoric
beans. He has shown that a mixed collection of charred seeds, later described as |
both Pbaseolus vulgaris and P. acutifolius, found in Canyon de los Muertos, Ari-
zona, was used by Ludwig Wittmack in 1888 to demonstrate conclusively the
American origin of the garden or common bean, P. vulgaris. More recently beans
dating from prehistoric times have been described from sites in the Southwestern
United States in relation to migrations and diffusion of culture traits and other
aspects of culture history. Most of this discussion has been of a highly general
nature, and little attempt to classify bean types on a subspecific level has been made.
1956]
KAPLAN—PREHISTORIC SOUTHWESTERN BEANS 199
It is hypothesized that the distribution of beans in the prehistoric Southwest
will show a relationship to the cultural subdivisions which have been recognized
by archaeological studies, The distribution patterns can be expected to reveal
something of the antiquity and history of beans as cultivated plants in this region.
MATERIALS AND METHODS
SOURCES AND COLLECTION OF MATERIALS
The data presented in this paper were collected from materials studied at or
obtained on loan from museums and universities where they had been deposited
by archaeologists and ethnologists active in the field of Southwestern prehistory.
A trip! to the Southwest during the summer of 1953 enabled the author to study
materials in museums of that region; the institutions visited are given in Table
vm. In the course of this field work, observations were made of Hopi and Navajo
cultivated lands. Further observations during field work? in rural Mexico in 1953
and 1954 contributed to the author’s understanding of comparative native
American farming and food habits.
Modern archaeological excavations are carried out with careful regard for the
position of artifacts recovered and their relation to dates of occupation and culture
change. Vegetal materials obtained from excavations are to a greater or lesser
degree artifacts and can sometimes be dated reliably with respect to their associa-
tions with other material for which dates have been established. Cutler (Martin
et al., 1952) has pointed out the value of large samples of plant materials, such as
corn cobs, in describing a specific class of vegetal remains occurring in a particular
time level of a prehistoric site. One advantage of a large sample is that it mini-
mizes the influence of extraneous material which may have been brought in from
other levels by rodents or by human disturbance such as burials in the fill. Beans,
unlike corn, seldom have a waste or by-product which can accumulate in the debris
of a cave or dwelling over a period of years. Threshing of dry beans, as has been
described in historic times among Indians in the Southwest ( Whiting, 1950;
Castetter and Bell, 1942), is carried out in flat cleared areas well removed from
habitation. The bean seeds arrive at the habitation ready for the pot or storage
with nothing to be discarded except, perhaps, the culls. The practice of threshing
beans in the field from the dry picked pods or from piles of the harvested whole
plants is widespread in Indian America. It is reported from coastal Peru (Gillin,
1945) as well as from the southwestern United States, and for representative climatic
regions of Mexico (Brand and Nunez, 1951; Foster, 1946; Kelly and Palerm,
1950), where it was observed by the author in 1954. A clue as to why pods are
occasionally encountered in some abundance in prehistoric occupied sites is pro-
1Supported in part by a Wychwood Fellowship in the Department of Botany, University of
Chicago. ui
2 Supported in part by a Chicago Natural History Museum Fellowship in the Department of
Botany, University of Chicago.
[Vol. 43
200 ANNALS OF THE MISSOURI BOTANICAL GARDEN
vided by Kelly and Palerm (1950) referring to the hand shelling of beans during
rainy weather when the pods cannot be rendered fragile by drying. Also, small
amounts of beans may be brought in and shelled by hand for daily use before t
main harvest or threshing. These practices, if carried on in the shelter of the
dwelling, could account for the large numbers of pods described by V. H. Jones
(Steen and Jones, 1941) and materials recently excavated from caves in Tamauli-
pas, Mexico, by R. S. MacNeish.
e uncharred bean seeds, fortunately for the archaeologist and botanist, have
escaped being eaten or have been left after being used in connection with burials
or ceremonial practices. The beans of Tularosa Cave, for example, were dispersed
throughout the debris and were probably seeds which had been dropped acciden-
tally. Some of these seeds may have been moved from their original positions by
rodent or human activity, as mentioned above, but the collection as a whole can
be said to characterize the period of occupancy of the site. Notwithstanding the
tact that there are sites (see Table VII) where time sequence of bean collections
has been fixed with some certainty, the emphasis in this paper is on the larger time
and cultural units and their relation to the regional history of agriculture. Thus
the individual collections which we have before us obtain greater significance as
they are considered with related collections in the Southwest. Dates or cultural
affiliation for each collection are taken from published or otherwise indicated data
based on standard methods and nomenclature used in Southwestern archaeology.
Samples obtained have been generally small, or at least of smaller numbers than
desirable. Samples always suffer in some degree from subjectivity in treatment.
The vegetal materials were collected by persons who, through lack of specialized
training in the recognition of plant materials and working under trying field
conditions, often may have failed to recognize and obtain all the material available.
Of the various classes of prehistoric cultivated plant materials, corn cobs, because
of their size and durability, are most easily recognized, while squash and bean seeds
are less noticeable. This is particularly true with scattered charred materials:
squash seeds become frustratingly fragile, and beans, losing their seed coats, occur
as elusive, single cotyledons. Beans are most readily collected when observed in
conjunction with larger artifacts, such as ceramic vessels or fragments of more
immediate interest to the archaeologist or his hired diggers. Laborious screening
of the fine debris which usually fills long-abandoned ruins is the most effective
technique for the recovery of seeds. Flotation also is useful in the separation of
charred materials from less buoyant debris. :
CLASSIFICATION OF MATERIALS
As each collection was received it was first examined for state of preservation;
that is, was it desiccated and in good condition, partially decayed, charred? Then
all bean material was separated from contaminants. The state of preservation
determined the number of characters which could be used for diagnosis, and
accordingly a classification was made using as many as possible of the characters
1956]
KAPLAN—PREHISTORIC SOUTHWESTERN BEANS 201
discussed below. The divisions were made first on the basis of species and then
into types of the species. The latter were the smallest divisions, and each putative
type was treated as a population which was described in its entirety or from a
random sample of its best-preserved units.
The term “type” is used here in a sense which is coordinate with “variety” as
used in such publications on bean classification as "Garden Beans” (Irish, 1901),
“American Varieties of Garden Beans” (Tracy, 1907) or “The Vegetables of New
York: Beans” (Hedrick, 1931). “Variety” in these works, although not specif-
ically defined, is taken to mean sexually propagated generations of plants which
retain characteristics enabling the separation of one such continuum from another
of the same species. Thus the characteristics used would be subspecific and, in
a cultivated plant, are those which affect its usefulness or recognition. An
example of the first kind of character might be precocity or disease resistance, and
of the second, seed-coat pattern. Since only a relatively few of the cultivated
beans are recorded to have been developed under recent cultivation, the term
“horticultural variety” is not much used. In agricultural practice bean-seed
characters, along with agronomic characteristics, are much used in classifying the
varieties. The nature of the present study precludes the use of agronomic char-
acteristics; seed characteristics are relied upon. This makes it uncertain, at best,
to include the materials treated here with the named varieties used in the above-
mentioned works or as listed in seed catalogues. For this reason the series of types `
described for Southwestern beans cannot have formal taxonomic standing but are
Presented as entities with which new materials may be compared.
Cardenas (unpublished thesis), Freytag (unpublished thesis), and others have
presented certain correlations between seed and vegetative characters, but these
are largely of a general nature not always useful in determining the nature of the
parent plant if only the seed is available. Pending further studies which will
enable more certainty in correlations of this sort, one can state similarities between
the seed types and the named varieties.
Diagnostic Characters.—Because of their large size and frequently distinctive
colors and markings, the seeds have been extensively used in the classification of
bean species and varieties. Hedrick (1931) and Bukasov (1930) have dealt
sufficiently with the synonymy and history of bean classification.
Diagnostic characters of bean seeds have been studied individually from the
standpoints of their genetics, physiology, and anatomy. The genetics of seed-coat
coloration and variegation has been shown by Lamprecht (1939b), Smith ( 1939),
and others to be quite complex. Smith found that six color genes plus modifiers
were responsible for the red testa color of the common bean variety Red Kidney
and that in other varieties no linkage was found among ground color, variegation
color, and extent of eye color. Glossiness of seed coat was reported by Prakken
(1937) as appearing to be genetically independent of factors for color and varie-
gation of seed coat and pod and vegetative characters in crosses of P. vulgaris
[Vol. 43
202 ANNALS OF THE MISSOURI BOTANICAL GARDEN
varieties. However, as far as I am aware, no one has studied the dull testas of P.
acutifolius seeds which is a diagnostic character of that species.
Shaw and Norton, according to Kooiman (1931), distinguished two classes of
color in bean-seed coats, a red-and-purple, and a yellow-black. The pigments of
the second were only slightly soluble in alcohol and alkali.
Use of the character of color quality introduces the problem of dealing with
color change over time. Hedrick (1931) recognized this problem and chose to
describe seed-coat color as it appeared in the first year subsequent to harvest. It is
common to find dark brown specimens in old collections labeled “yellow beans,”
or to find familiar varieties of quite a different (dark) color than the fresh seed
of the same variety. Skalinska (cited by Kooiman, 1931) showed that yellow and
yellowish pigments oxidize to brown in vitro.
It is possible to divide color change into two physiological periods, that of
ripening and during dry dormancy. Very young seeds are green and may remain
so or may be tinged with green through maturity, as in French horticultural and
certain greenish-white tepary beans. Loss of the chlorophyll may be accompanied
by whiteness or lack of color which may persist or be quickly replaced by light
tints of what would probably be the color at maturity. Coloring is first noticeable
in the eye ring and then appears in the other parts of the testa, but darkening
earlier with proximity to the placenta. Where there is patterning, it occurs as a
sort of "developing out" process, reminding one of the development of a latent
image on photographic paper. Streaks, spots, or other forms of variegation appear
as islands of coloration on a light ground which itself may become colored subse-
quently. The color change which occurs during this period is largely quantitative,
as indicated previously, but may proceed to a point where it appears to be quali-
tative. In the tropical black P. vulgaris the young rose-colored testa changes by
the time of maturity to a purple so deep as to give the effect of black.
Subsequent to ripening, the rate of color change in the dry bean is slower but
may be accelerated under certain conditions. The darkening of the light pink
ground color of the seed coats of “Mexican pinto” (Type C13) beans has been
observed to occur at a differential rate when the seeds were exposed to strong
sunlight for several months. The sides of the seeds exposed to sunlight were much
darker than the sides not so exposed. Samples of seeds which had been stored at
room temperature for fifty to sixty-five years failed to show any color change
when maintained at a temperature of 60° C. for thirty days, suggesting that
limits of normal change had been reached at some time during this period.
It is sometimes difficult to distinguish what the color of the seed coat might
be were it not for opacity. Thus, among black beans it is difficult to determine the
nature of the blackness, The Hopi “Blue Dye” beans (Type C29), for example,
are very dark blue while other “blacks” may be dark purples. In this study the
use of color in the classification of seed types has been made with a realization of
the changes in color which have undoubtedly occurred; nevertheless, all designa-
tions are based upon color quality at the time of examination. The larger color
1956]
KAPLAN— PREHISTORIC SOUTHWESTERN BEANS 203
TABLE II
DISTRIBUTION OF COMMON BEANS ON THE BASIS OF SEED-COAT COLOR PATTERNS
Indian Southwest Mexico* Peru* U. S. Commercial* *
-
KEE
3 o a * à
SG m E of
9 E 3 [29
"ol uu RER
CES, |
< [3 5 ola
Self
Number of types 19 6 7 162 43 117
Per cent 65 55 70 65 67 56 50
Variegated
Number of types 11 5 3 3 84 34 118
Per cent 37 5 45 1.30. | 35 33 44 50
* Ditmer, e£ al. (1937).
** Hedrick (1931).
categories have been used rather than an exact description which would be
subjective and not too meaningful.
Seed size in beans has been estimated (Kooiman, 1931) to be the result of the
action of many genes which apparently affect the various dimensions equally. Thus
within a population of seeds which have been selected for uniformity, the ratios
between the dimensions of the larger seeds and those of the smaller seeds may be
expected to vary little, providing all have reached normal maturity at the time of
harvest and were grown under similar conditions. Within a given pod the largest
seeds usually occupy a medial position and the smaller seeds are at either extremity.
It would be well, then, to obtain seed samples for measurement from the entire
contents of fully matured pods. Seed measurements, considered as reflections of
the size and shape of seeds, represent the most useful set of criteria for the quanti-
fication of variation in these organs. But it should be recognized that seed
dimensions may be differently affected by environmental conditions. Bean growers
in humid eastern lands who plant disease-free rom western arid lands note
that the beans that they harvest are shorter and less flat than those which they
planted (Hardenburg, 1942).
Prominence of the radicle and plumule located on the ventral (adaxial) margin
of the seed, just anterior to the hilum, lends an apiculate appearance which is more
frequent in some varieties than others. The extremes or ends of the seed are
rounded in the lateral view, or one or both extremes of the seed may be truncate
or appear to be flattened. If only one end is truncate, it is the posterior end and
the apiculation may still be seen at the other extreme. In the pod, seeds are borne
on the placenta along the ventral surface, with the growing parts of the embryo
toward the apex. The scar, formed by abscission of the placenta and seed, is
oriented longitudinally on the ventral margin of the seed, with the micropyle at
[Vol. 43
204 ANNALS OF THE MISSOURI BOTANICAL GARDEN
L gre o characters of seeds: A, reniform; B, apiculate anterior end;
aie ends; D, entire; E, testa venation radiating from hilum of lima bean; F, e A
G, truncate ends; H, flecked pattern; I, eyed; J, bsstciliatlly striped.
the anterior end closest to the growing parts, and the paired elevated peaks
of the testa, the caruncle, at the posterior end. It has been suggested that
truncate seeds in Phaseolus are the results of: (a) interference by intralocular
parenchyma tissue; (b) abutment of the seeds themselves. The first explanation
assumes that mechanical pressure of the parenchyma is sufficient to limit the
growth of a maturing seed and that it exerts its influence in some varieties but
not others. In the second, truncate seeds are characteristic of some of the tepary
beans the seeds of which are not at all appressed in the pod
Technique.—All measurements of seed dimensions were taken with a vernier
caliper and are given in the metric system. Each dimension was measured to give
the maximum result so as to standardize the method and make repetition easier.
In the actual measuring of length, for example, the apical anterior end was placed
against the stationary jaw of the caliper and the sliding jaw closed until the
posterior end was contacted. The jaws were then tightened to the extent that,
when rotated on the apical end of the seed, the basal end would just slip by,
neither sticking nor passing freely the surface of the movable jaw. An adequate
degree of precision is obtained in this manner. Each seed was measured in e
length from anterior to posterior extreme, in width from ventral to dorsal margin,
and in thickness from one lateral surface to the other.
TABLE III
EXTERNAL SEED CHARACTERS OF FOUR SPECIES OF PHASEOLUS
WITH PARTICULAR REFERENCE TO SOUTHWESTERN VARIETIES
P. vulgaris P. acutifolius P. lunatus P. coccineus
Dimensions (cm.)
Length 1.85—.74 1.20—.66 1.80—1.14 2.39—1.46
Width 1.08—.49 78—44 1.29— .85 1.48-1.13
Thickness .85-.34 56-.27 0- 35 1.21- 72
Form Various but seldom cuboid Semi-spherical or flat, often | Reniform, especially in larger- — usu ally semi-round
cuboid with sharply trun- seeded grou ms sma m cross-section, ends round
owes nds ed pi w with
or and md
cate, anterior end equo
in dorso-ventral plan
Seed coat
Venation Reticulate or not apparent Not apparent Parallel and branching Not apparent
radially from hilum
Glossiness Very glossy to moderately Dull Glossy to dull Highly glossy
after polishing dull
Hilum
Caruncle* Two prominences separate Same as P. vulgaris Prominences joined Same as P. vulgaris
anteriorly
Ring** Elevated Little elevated Elevated Elevated
Size Not proportional t to seed size, | Same as P. vulgaris
Roughly proportional to
seed size
remains small in large-
pes
seeded ty
Same as P. vulgaris
* See fig. 2.
** Margin of testa
encircling hilum.
SNVH8 NYALSAMHLNOS OIHOLSIHdHd—NV'IdVMX
soz
[9561
[Vol. 43
206 ANNALS OF THE MISSOURI BOTANICAL GARDEN
In many cases among the archaeological collections it was possible, owing to
the fragmentary or charred nature of the material, to measure only a few of the
seeds in a collection.
It is possible to obtain a surface impression of the bean seed using any one of
several acetone or ethyl acetate soluble plastics or Duco cement. Microscopic
examination of a number of such impressions from seed coats of varieties of the
four species dealt with here failed to show consistent differences which could be
used in the classification of see
DESCRIPTION OF BEAN TYPES
The use of scatter diagrams was adopted for graphic presentation of the data,
as this method (Anderson, 1949, pp. 81-101) facilitates the comparison of sev-
eral classes of data simultaneously. The source and size of each collection and its
measurements are given in Table VII, as well as the number with reference to the
typing system discussed previously. In Table IV the qualitative data—color,
variegation, and form—are listed for each type. Table V includes the maximal,
minimal, and median dimensions for each and the numbers of all collections repre-
sented by each type. Tables IV and V, along with figs. 1 and 2, provide all
the characteristics of each type. A summation of species characteristics for
Southwestern beans is found in Table III.
Association of Seed Characters.—The scatter diagrams (figs. 3-13) show that
in the collections studied greater size in either seed length or width is associated
with larger size in the other dimension. The degree of seed thickness has a less
regular association with either of the other two dimensions than they have with
one another. It may also be noted that variegated seed-coat patterns in the
common bean also tend to accompany large seeds.
Seeds of the two principal species of Phaseolus dealt with here, Phaseolus vul-
garis and Phaseolus acutifolius var. latifolius, form a discontinuous distribution
readily seen in the scatter diagrams. Although this discontinuity is a reflection of
seed dimensions, except for the patternless, or phenotypically patternless, white
seeds common to both species, the color patterns are discontinuous also. It is the
symbols mainly of the small white (Type C 6) seeds of P. vulgaris which cluster
below the dimensional modes of this species and are found within the range of P.
acutifolius, in the lower left-hand portion of the diagrams.
In addition to the small white type, the two curious small red (Type C20)
and small flecked red (Type C12) P. vulgaris types overlap the teparies in size
range. These were classed with Phaseolus vulgaris with some misgivings. How-
ever, should further study prove them to be variants of P. acutifolius rather than
P. vulgaris, two interesting aspects will remain, First, the Murder House (Collec-
tion Number 195d) beans are from an area of tepary importance and common beans
as well as lima occurrence; the other (Collection Number 206) are from the “North-
ern Periphery,” an area of which the cultural and agricultural affinities are not
yet well defined. Second, the seed characters show similarities to those of both
1956]
KAPLAN—PREHISTORIC SOUTHWESTERN BEANS 207
x i4 JN
2
3 2
3
3
4
? 4
A B C
Phaseolus vulgaris P. acutifolius P. lunatus
(or P. coccineus) var. latifolius
2. Variation in hilum form: 1, micropyle; 2, placental scar; 3, hilum ring;
i lunatu
4, A cid Note incomplete separation of caruncle ridges in P. lunatus
species to a greater extent than is usual It is true that among white-seeded
varieties it may be quite difficult to distinguish between P. vulgaris and P. acuti-
folius seeds, but this is not the case with the darker-colored and patterned seeds.
It would be most interesting if viable seed of these types could be obtained and
sown, to examine the vegetative characters of the plant.
The Earliest Bean Remains.—The earliest well-dated beans reported from
Southwestern sites are those of the Mogollon in the pre-pottery or prior to 1 A. D.
occupation level of Tularosa Cave (Martin e£ al., 1952). This singular discovery
offers sharp contrast to pre-pottery cultures of the Anasazi and Hohokam. In the
Anasazi and Hohokam areas beans have not been found in cultures reliably dated
as prior to Basketmaker III (ca. 500 A.D.) and from early Hohokam sites. In
Tsegi Canyon in Northern Arizona, a prepottery Basketmaker site (Kidder and
Guernsey, 1919), investigators failed to find beans, but they reported corn and
other vegetal remains which evidence good preservation of artifacts of this sort.
Beans, as well as squash and corn, were found in Tsegi Canyon sites subsequent
to the introduction of pottery. Jones and Fonner reported fully on the vegetal
remains of a number of Basketmaker II sites near Durango, Colorado (Morris,
1954), but they did not describe any beans. Gladwin’s (1937) excavations at
Snaketown, a Hohokam ruin now the site of a Pima Indian settlement, uncovered
carbonized corn in houses dated at about 500 A. D. at the earliest. The only beans
from this site were charred tepary and common beans dated at about 1000 A. D.,
which would correspond to Pueblo II in the Anasazi. The identification of these
remains was made by Volney Jones (Castetter and Bell, 1942).
As has been indicated previously, in the quest for bean remains we are looking
primarily for the very materials which the prehistoric Indians would have least
wanted to leave behind, In times of food scarcity, single beans dropped on the
floor of the dwelling may have been carefully retrieved. The most that can be
said then of the occurrence of beans in the early periods is that, if present, they
were not plentiful.
TABLE IV
MORPHOLOGY OF TYPES
Type Color Ground Gloss* * Eye ring Anterior Posterior Three- Varietal
pattern color end end dimensional similarity
shape
SCH Self Orange-red +++] Dark Truncate Truncate Cylindrical,
to dark entire
orange-red
C2 Self White +- Indistinct Round Round Subreniform,
semi-round
C3 Self Dark +4 Indistinct Subapiculate Round Reniform, Large Red
red-brown o round flat Kidney
C5 Faint mottle Dark ++ Dark Apiculate Round Entire
red-brown
Ce Self White +-+ Indistinct Subapiculate Round Entire
C Self Cream to +++] Indistinct Subapiculate Round Entire, flat
yellow
C8 Self Dark red d Indistinct Subapiculate Round Entire, flat
C9 Self White -+ Indistinct Subapiculate Round Entire,
semi-round
C10 Self Dark ++ Indistinct Subapiculate Round Subreniform,
red-brown cylindrical
* C refers to common bea
a dint.
n (P. Kam mp vulgaris) types.
80c
NHGWV9 IVOINVLOG INNAOSSIN AHL AO SIVNNV
EF 1A]
Cila
Faint
longitudinal
stripe
Distinct
longitudinal
stripe
Fleck
Brown fleck
and stripe
Red or tan
mottle
Red or tan
mottle
Self
Red-brown
eye
Self
Self
Self or
dark fleck
Dark fleck
Self
Pale violet
to violet-
rown
Brown-violet
red-brown
Pink
White to
cream
White to
cream
Dark red
White
Yellow-
brown
Violet-
brown
Dark red
Dark red-
brown
Red-brown
-++
Dark
Dark
Dark
Dark
Indistinct
Indistinct
Dark
Indistinct
Dark
Indistinct
Indistinct
Indistinct
Dark
Subapiculate
Subapiculate
Round,
tapered
Subapiculate
to apiculate
Round
Round to
apiculate
Subapiculate
to apiculate
Round
Subapiculate
Round
Round
Round to
truncate
Round
Round
Round
Round to
flattened
Round
Round
Round
Round
Round
Round
Round
truncate
Round
Entire,
semi-round
Entire,
semi-round
Entire,
semi-round
Reniform,
flat
Entire,
semi-round
Reniform,
flat
Entire, flat
ntire,
semi-round
Subreniform
to reniform
Subreniform
Reniform
Entire, semi-
cylindrical
Garrapata or
into
eg s cattle
; Vaquita
Red Mexican
China Red Eye;
a
Golden Wax
Vayo (Bayo)
SNV3IH NYIALSIAHINOS ODHOISIHHWHUd—NV'IdVM
60c
Loser
TABLE IV (Continued)
Type lor Ground Gloss* * Eye ring Anterior Posterior Three- Varietal
pattern color end end dimensional similarity
shape
C21 Self Dark red MR Indistinct Round Round ntire,
semi-round
C22 Self Yellow, +++] Dark Round Round Entire, Bolitas
brown, semi-round
brown-red
C21 Self Brown +++] Dark Round Round Entire,
y semi-round
C24 Brown eye- White +++] Indistinct Subapiculate Round Entire, Dwarf Golden
pattern semi-round Wax
superimposed
C25 Violet Pale violet +++] Indistinct Subapiculate Round ire,
stripes semi-round
C26 Dark red Red qe Indistinct Round Round Entire, flat
stripe
C27 Self White +++] Indistinct Round Round Entire, semi-
cylindrical
C28 Self Pink +++] Dark Subapiculate Round Entire, flat
C29 Self Black (dark 44 Indistinct Round Round Entire, flat
blue) to flat
C30 Self Black +++ | Indistinct Apiculate Round to Entire,
truncate cylindrical
"Li Self Dark purple +H Indistinct Round to Truncate Entire
subapiculate
L2 Self White ++ Indistinct Round Round Entire
*L refers to lima bean (Phaseolus lunatus) types.
Oc
NüGWVO TVOINVLOd@ INNOSSIN AHL 40 SIVNNV
EF 'I0A]
Dark fleck
Dark fleck
and stripe
Light tan eye
pattern
suffused
Self
Black fleck
Self
Self or
brown fleck
Self
Self
Black flecks
almost com-
pletely cover
ground
Dark fleck
Self
Purple mottle
and fleck
Brown
Dark red
Dark brown
Red-brown
Tan
White
Orange-brown |
Brown
White
Tan
Red-brown
White
White, violet
+++ +
+ +
Indistinct
Indistinct
Indistinct
Indistinct
Black
Indistinct
Indistinct
Dark
Indistinct
Indistinct
Indistinct
Indistinct
Indistinct
Subapiculate
to round
Round
Subapiculate
Subapiculate
Round
Round
Truncate
Truncate
Round
Truncate
Round
Round
Round
Round
Truncate
Round
Truncate
Round
Truncate
to roun
Truncate
Round
Truncate
Round
Round
Round
Entire
Entire
Entire
Entire,
semi-round
Entire, flat
Semi-round
Entire, flat
Entire
Entire,
semi-round
Entire, flat,
c
uboid
Semi-round
Semi-round
Semi-round
Jackson's
Wonder
*T refers to tepary bean (Phaseolus acutifolius) =
to runner bean (Phaseolus coccineus
*R refers
SNVdIH8 NYIALSIAHINOS ODIHIOLSIHdHUd—NV'IdVM
[9561
[Vol. 43
212 ANNALS OF THE MISSOURI BOTANICAL GARDEN
The Mogollon furnishes a situation which may help to explain both the spotty
occurrence of early beans in other areas and the clear regional differentiation of
varieties in later times. The peoples of Cordova and Tularosa Caves (Martin ef al.,
1952) were contemporaries living in the Reserve, New Mexico, region. They
shared many basic cultural traits, but the people of the Cordova Cave site relied
more upon hunting than did those of Tularosa Cave. Bean remains were relatively
abundant in the latter site, but only a single bean and one pod fragment were found
in Cordova Cave, and vegetal materials from other pre-1200 Mogollon sites indi-
cate non-importance of bean cultivation. If the same kind of distribution—few
bean-growing and many non-bean growing communities contemporaneous in the
same region—occurred in the Basketmaker Anasazi region, it may be said that
early Basketmaker beans have not yet been found but that they do exist. With
greater attention on the part of persons trained to recognize and identify charred
seeds and other remains, more Basketmaker beans and those of other pre-pottery
peoples might be recovered.
Varietal Change.—On the basis of seed characters the earliest beans of Tula-
rosa Cave do not differ in any way which could be interpreted as part of an
evolutionary sequence from the most recent beans of Tularosa Cave, despite the
difference of over 1,000 years. This is generally true of Southwestern bean remains.
Changes in cultivated prehistoric beans of the Southwest which are correlated with
time changes are those which show the introduction of new species or varieties
from an older to a newer region or culture. The question of evolutionary change
in beans under domestication may be reopened with the anatomical study of bean-
pod remains. However, it can be said that the apparent lack of change of beans
under domestication in the Southwest is further indication that development from
the wild kinds occured elsewhere or outside of the Southwest, and at an earlier date.
Bean Preparation.—Among the beans collected from contemporary Indian
peoples there is in the Museum of Anthropology of the University of California at
Berkeley a sample (Collection No. 117a) of parched beans labeled “parched small
white P. vulgaris" obtained from Yuma Indians? A collection of more than one
quart of common beans from Kiet Siel, a Pueblo III ruin in Tsegi Canyon, northern
Arizona, was examined at the Museum of Northern Arizona and found to have
been lightly toasted. This sample, as well as the preceding one, emits a character-
istic odor; seeds which obviously were originally white are tanned irregularly, and
cotyledons of the white seeds which have the most tanned testas are browner than
those of less tanned lighter seeds. Seeds of this collection are described in Table
VII Nos. 188, 189, 190, 191, 192, 193, 194a, 194b.
3 These, however, are teparies (Type T 3) rather than common beans.
1956] `
KAPLAN—PREHISTORIC SOUTHWESTERN BEANS 213
TABLE V. DIMENSIONS OF SEED TYPES
Range of Measurements (cm.)
Median of Measurements
Maximum Minimum
> S. ae Fe Seen SLU äi SE E E
d TR) Ron eh ETT > "
Cl 1.69 94 67 1.05 61 48 1.30 71 57
C2 1.49 .87 Set .64 A4 1.25 73 55
C3 1.67 92 66 | 1.07 .68 42 1.43 80 57
C5 1 95 203 11.15 76 1.35 85 57
Ce 1.28 75 55 79 50 37 9 60 48
1.35 90 .68 | 1.15 73 50 1.29 87 51
C8 1.61 | 1.03 56 | 1.43 91 49 1.49 99 54
C9 1.50 68 1.05 66 6 1.3 3
C10 1.69 80 70 | 1.61 79 .65 | 2 seeds | 2 seeds seeds
sep. coll. | sep. coll. | sep. coll.
Cll 1.73 97 SCT .66 45 8
Clia 16 78 en | 1 71 44 12 seeds
sep. coll. | sep. coll. | sep. coll
C12 89 63 .50 52 34 .84 6 48
C13 1.71 89 .68 1.08 67 41 1.26 78 .54
C14 1.76 95 .68 | 1.04 65 39 1.32 .81 56
C14a 1.26 92 .74 | 1.03 66 55 1.16 28 64
C15 1.38 84 59 | 1.07 65 42 1.08 .65 43
Cie 1.22 70 59 .00 68 .57 | 2 seeds | 2 seeds | 2 seeds
C17 1.65 92 1.11 64 43 1.41 81
C18 1.70 Az. .67 36 68 47 1 76 54
C19 1.65 90 59 | 1.33 70 44 1.50 72 51
C19a 1.85 94 .67 | 1.39 63 49 1.58 85 58
C20 94 62 45 .87 56 40 93 59 43
C21 1.25 86 26 1 1.19 80 .62 1.20 82 69
C22 1.38 .83 .66 | 1.03 63 1.22 24 55
C23 1.38 78 57 95 .60 42 1.0 8
C24 151 | 1.08 .85 | 1.26 75 63 1.44 93 73
C25 1 seed 1.14 86 68
C26 1.54 .84 4t 1148 74 E 1.58 80 48
C27 58 42 77 36 9 53 39
C28 1.41 86 78 | 1.00 ee A3 1.18 74 50
C29 1.54 87 .60 | 1.10 70 E 1.26 76 52
C30 1.34 62 69 1.34 .60 seeds | 2
Li 1.64 | 1.16 .60 | 1.29 .90 40 1.47 1.03 53
L2 1.58 | 1.11 58 | 1.18 85 41 37 d 52
L3 1.80 | 1.20 i. 1.37 95 42 1.58 1.08 55
L4 1.76 | 1.29 35 LESE 1.60 49 1.62 1.14 52
Ls 1.48 | 1.04 gh ae hE gh Gaui | 35 1 91 44
T1 .67 552 37
T2 As 70 54 27
T3 1.19 | .70 | ap 87 | .57 37 | 100 e 43
T4 1.20 78 45 .81 50 31 1.03 67 Al
T5 1.05 | .69 At a AT SM 8 ep 39
Te 95.1 «ue bos bib ws 30 .84 55 Al
H iil bol 4 Bia 1.02 ei A2
T3 .82 55 36
R1 2 S 120 | 14€ | 120 |. 94 | 1.90 1.28 1.13
R2 Mer v & 1114 isi 72| 1 1.32 98
hus
4 [Vol. 43
214 ANNALS OF THE MISSOURI BOTANICAL GARDEN
West (1947) reports an interesting practice of certain Tarascan Indians which
may be related to this problem. A variety of P. coccineus, grown only by the
more conservative people and tended mainly by the women in their house gardens,
is used in special dishes, among which is a pinole made with a special "black maize.”
The corn is said to be toasted, but the preparation of the beans prior to grinding
is not indicated. "This practice was not found among non-Indian (Ladinoized)
residents of the area.
Whiting (1950) reported that the Hopi parch white and mottled teparies
before cooking. This treatment of the white tepary (or in its absence, "other
white beans") is noted in connection with the breaking of a ritual fast by priests.
Although the material from Kiet Siel, which, according to tree-rings dates back
to as early as 1106 A. D. and as late as 1221 A. D. (McGregor, 1934), contains
the oldest sample of parched beans yet seen, parching may be supposed to have been
practiced earlier and may well have formed the basis of dry-bean use for protein in
pre-pottery times.
Parching of dry mature beans for use in such foods as pinole may have once
been important and widespread, but now parching has been all but superseded by
boiling. Ritual food preparations, such as those referred to above, may be the
most likely conditions under which an ancient manner of food use might be
retained although reinterpreted.
DISTRIBUTION OF BEAN TYPES
Tepary Beans.—The greater importance of tepary bean cultivation in the most
arid region of the Southwest, that of the Pima and Papago (Hohokam), has been
shown among the contemporary Indians by Freeman (1912), and Castetter and
Bell (1942). Fewer tepary varieties than the forty described by Freeman were
encountered in the collections available for this study. However, collections from
the Pima and Papago and the Colorado River tribes (Table VII and figs 3-13)
substantiate the extensive use of teparies by these tribes in recent historic times.
Efforts by the author to obtain teparies in 1953 and 1954 on the Pima and
Papago Reservation at Sells and Ajo, Arizona, and from trading posts on the
Cocopa Reservation were unsuccessful. Local residents said that droughts of recent
years and availability of the popular pink beans (Type C26) may explain the
general decline of old bean varieties among the Indians.
As the agronomic, botanic, and ethnologic writings on the tepary invariably
refer to the drought and heat resistance of this species, some of the problems re-
lating to the history of its domestication should be discussed.
Freeman (1912) first reported the superiority of yield of teparies over common
bean varieties under dry-land and irrigated conditions in Arizona. Good yields
under dry-land conditions have been obtained as far north as Colorado, where the
Redfield tepary out-yielded the high-producing Pinto common bean (Brandon,
1943). The paucity of prehistoric tepary bean remains outside of the southern
1956]
KAPLAN—PREHISTORIC SOUTHWESTERN BEANS 215
Arizona-New Mexico region is not to be explained by any unsuitability of varieties
of this species to the northern, Anasazi, region.
If the tepary bean is a domesticate of the Hohokam or other peoples of the
Sonoran Desert region, then its involvement with the history of these people may
help to account for its importance among them, while the restricted occurrence in
the Anasazi and Mogollon may be attributed to partial unacceptability of teparies
to peoples long accustomed to common beans. If, on the other hand, tepary beans
were domesticated far in the south of Mexico, they must have found acceptance
among many peoples who were familiar with common beans, and probably lima
and runner beans as well, before reaching the Southwest. Although early Hoho-
kam archaeological sites are not good ones for the preservation of vegetal materials,
distribution patterns of teparies (Table VI) suggest their late introduction and
diffusion in the Southwest.
The question of origin of the tepary bean will be solved neither by De Can-
dolle’s criterion of the location of related forms in the wild state nor by Vavilov’s
“center of varietal diversity” hypothesis nor will studies of archaeological
materials supply all the needed information to elucidate the role of the tepary bean
in Southwestern prehistory. It is necessary to know the source of variability and
the history of domestication of this plant.
Freeman (1912) supposed that the tepary bean was domesticated from very
small, angular-seeded, broad-leafed indigenous forms (P. acutifolius var. latifolius)
growing in the watered canyons of the Sonoran Desert and that selections from
these types yielded the many varieties which he recognized among contemporary
cultigens. Freeman made no estimate of the time period necessary for the great
increase in seed size and change in form from the wild to the domesticated forms
nor were the mechanics of the change indicated. Carter (1945) accepted Free-
man’s hypothesis, and used the element of time as an argument for greater antiq-
uity of man in the Southwest. Castetter and Bell (1942) rejected Freeman's
conclusion on the basis of the great difference in size and form between the
cultivated and the non-cultivated P. acutifolius var. latifolius, and on the fact
that the earliest prehistoric beans which they had seen in no way resemble any of
the non-cultivated beans but can be equated with modern cultivated varieties.
The latter view concerning the identity of the prehistoric and modern materials is
supported by data presented in this paper (Table VII). It would be strange
indeed if beans of the already-high agricultural cultures which supply the earliest
material of this sort were to be of a primitive type. Cultivation under irrigation
of the small, angular-seeded wild teparies (Castetter and Bell, 1942) did not change
the seed characters. :
Selection and cultivation without accompanying or preceding changes in the
germ-plasm could hardly account for the differences between Southwestern culti-
vated and indigenous teparies. A source of change might be mutation, but seed
Size is not a single-gene character; rather it appears, as indicated previously, to be
the result of interaction among several genes directly, while dd other characters
are involved indirectly. The rates of mutation at these loci are not known, ee
extremely careful selection and breeding, as well as a very long time, wo
[Vol. 43
216 ANNALS OF THE MISSOURI BOTANICAL GARDEN
TABLE VI. OCCURRENCE OF TYPES
Prehistoric Contemporary
San Juan Anasazi
Mesa Verde
Northern Arizona
Anasazi
Verde Valley
Hohokam-Anasazi
Colorado River
Yuman
Northern Periphery
Arizona Piman
Zuni
Navajo
X | Rio Grande Pueblos
x
XX
X X
XXX |Hopi
o
XXXXXXXXX | Mogollon
XXX
X
X
G29
AA u
»
Xx
X XX XX
2)
au
X
NK UXKX
XXX
x
S
D
Xx
x
d Q
x
XX
XXXXXX XXX XX
x
d
XXX
XXX
XXX
X
Té x
T8 x
q
E MR XX
* C refers to common bean (Pbaseolus vulgaris) $ o li :
types; L, to lima bean (Phaseolus lunatus) types;
T, to tepary bean (Phaseolus acutifolius) types; R, to runner bean (Pbaseolus coccineus) types.
1956]
KAPLAN—PREHISTORIC SOUTHWESTERN BEANS 217
probably be necessary for the effect of any such mutations to be expressed pheno-
typically. A more likely source of variability in tepary beans is hybridization with
some other related species. This process is certainly suggested by Freytag’s demon-
stration (unpublished thesis) of the effect of introgressive hybridization in common
beans.
Unfortunately, very little genetic or even breeding work has been done on
teparies, and no interspecific crosses have been reported. Teparies in the Southwest,
and probably common and lima beans as well under arid conditions, are almost
entirely self-fertilizing owing to very early pollination in the unopened flower bud.
A thorough study of cultivated and non-cultivated P. acutifolius and P. acutifolius
var. latifolius and suspected hybridizing species is necessary to indicate what the
source or sources of variability in this species might be. Also of importance would
be the determination of the geography and ecology of hybridization in this species.
Soutbwestern and Otber Bean Assemblages Compared.—The grouping of bean
types presented in Table II suggested that large geographic areas have character-
istic bean assemblages. At present, it is not possible to give more significance to
the distributional patterns other than to indicate that they exist. Whether the
cultural or natural agencies shaping the distribution are selective or non-selective
is likewise obscure; however, some of the selective factors which might be worthy
of further investigation are discussed below. Data are drawn from Southwestern
materials examined by the author, from the reports of the Russian investigators
(Ditmer e£ al., 1937), and from Hedrick's study (1931) of the United States
commercial beans.
It will be noted that never does the percentage of variegated-seeded forms
exceed that of the corresponding self-colored forms. The highest percentage of
variegated forms, 50 per cent in the United States, reaches that figure largely
because of the great diversity and number of variegated forms among the bush
snap beans. Many of these are introductions from Europe, and the needs of the
canning industry and available year-round markets have stimulated the use of
many varieties with differing horticultural characteristics. Diversity in seed
characteristics here is incidental. Among the bush snap beans described by Hedrick
(about 127 named varieties), 55 per cent have variegated seed coats. In all other
varieties there are considerably fewer variegated forms than there are self-colored
ones. The percentage of strictly prehistoric types of the Southwest most closely
approximates Peruvian distribution, while the totals for the Southwest, including
prehistoric, contemporary, or both periods combined, are most like the Mexican dis-
tribution and very little like the 1:1 ratio for the United States. The high percent-
age of self-colored beans, which occurs only in contemporary Southwestern Indian
cultures, and the fact that few of these are found as field beans in other more humid
parts of the United States, indicate an influx during historic times of self-colored
beans. The trend in the Southwest thus has been toward absolute and percentage
reductions in the numbers of variegated bean types and a corresponding increase
[Vol. 43
218 ANNALS OF THE MISSOURI BOTANICAL GARDEN
in self-colored types which are dry shell beans, except for the Hopi “Black” string
bean (Type C30). The importance of the Pinto or Garrapata bean (Type C13),
a variegated dry shell bean, is also recent.
Freytag (unpublished thesis) states that Latin American beans used in the
dry-shell stage over a wide area are likely to be self-colored and not of an unusual
shape, while variegated or odd-shaped dry-shell types are of restricted distribution.
These selective factors are apparently not operative in snap beans as such beans
are consumed before seed maturity. This seems to be true in the United States,
as variegated dry beans, with the exception of the pinto varieties which are used
for Mexican or Texas style dishes, are of quite restricted distribution.
Interpretations of Bean Distribution: —
Three fairly distinct regions can be outlined in the prehistoric Southwest for
the cultivation of beans. So far as is possible, the source of beans, the dates of
their introduction, cultural affinities, and movements within the Southwest will
be indicated.
It was hypothesized that the distribution of cultivated beans in the prehistoric
Southwest would show a diversity resulting from differing climatic and ecological
conditions and from differences in culture history. The influence of the environ-
ment and some ethnobotanical factors such as food preparation have already been
discussed. A division of the Southwest following three main cultural groupings
was adopted for the organization of bean distributions into regions. These regions
can be compared with those of other plant materials and culture traits.
I. Mogollon:—
Although beans are known only from a few sites in a limited area of the
Mogollon, it is evident that the cultivation of the common bean in this area
preceded the introduction of pottery. The number of pre-pottery beans from
Tularosa Cave is small but the prepottery context is clear, and it may be stated
that by 300 B. C. common beans, along with maize and pepo squash, were being
cultivated by Mogollon Peoples. 'The number of communities or settlements
cultivating beans at this time cannot be estimated, but it is probable that it was
not large, as none of the Mimbres River sites nor Cordova Cave, which is even
closer to Tularosa Cave, show any evidence of important bean cultivation. The
open sites of Higgins Flat and Hinkle Park, which appear to be immediately sub-
sequent to and in the same cultural tradition as the Tularosa Cave, indicate that
bean culture continued with common types similar to those of Tularosa Cave and
may indicate a greater extent of bean growing. Since tepary beans do not occur
in the Mogollon before 1100 A. D. (Higgins Flat), they may be regarded as intro-
ductions from the Hohokam; there seems to be no prior association of P. acuti-
folius in the Mogollon with Cucurbita pepo of which abundant remains were found
in Tularosa Cave.
1956]
KAPLAN—PREHISTORIC SOUTHWESTERN BEANS 219
2. Hobokam:—
As noted previously, vegetal materials are absent in most early Hohokam sites.
While this lack may be due to poor conditions of preservation, beans first appear
in the Sacaton phase, or about 1,000 A. D. at Snaketown. The charred tepary and
common beans identified by Volney Jones are said by Castetter and Bell (1942,
P. 32) to constitute these remains, and may be the earliest record of tepary beans
in the whole Southwest as well as in the Hohokam. Not enough evidence is avail-
able at present to warrant conclusions as to approximate dates for the introduction
of cultivated beans to the Hohokam, nor is it possible to say whether beans were
utilized in pre-pottery times in this region. Since the distinguishing feature in
Hohokam beans seems to be that they are tepary varieties, which may repre-
sent a response to climatic rather than cultural factors, a long history of selection
for varietal preference need not be hypothesized. The climatic conditions of
southern and southwestern Arizona would quickly encourage the cultivation of
tepary beans after their introduction. If domestication of the tepary occurred in
the region of the Hohokam, it is not now possible to estimate when the cultivated
varieties emerged as different enough from the indigenous forms to be differentiated
from them. The Hohokam ruins which have provided the best bean material are
later than 1,000 A. D., and usually they show the influence of the Anasazi in
several aspects of culture. Montezuma's Castle (1300 A. D., Collection Nos. 201—
205a), located in the Verde Valley, central Arizona, is listed by Carter (1945,
P. 24) as a Pueblo III, Little Colorado River site, among those sites demonstrating
the advance of Curcurbita moscbata from the Mesa Verde region to the south and
west into the area of the Hohokam. In this site only have been found uncharred,
definitely identifiable bean remains representing the three species known to occur
in the prehistoric Southwest. A mixture of bean assemblages of the Hohokam and
Anasazi are evident in the teparies, and limas from the Hohokam and the charac-
teristic striped violet common bean (Type C11) of the Anasazi. Other sites of
the Verde Valley evidence the influence of the Anasazi bean types rather than
those of Mogollon.
The lima bean appears to be characteristic of the Hohokam. Mackie (1943)
has the theory that lima beans entered the Southwest by way of western Mexico
and the Colorado River tribes into the Hohokam area. He discounts the likelihood
of the eastern Mexico-Sierra Madre Oriental corridor as unsuitable because of
excessively humid conditions. Mackie also finds that the Hopi lima beans and the
ans of eastern United States Indian groups are identical and suggests that the
eastern limas are an extension across the Mississippi Valley of the Hopi varieties.
Carter considers that the absence of native archaeological lima beans east of the
Hopi mesas until Pine Bluffs, Arkansas, necessitates an eastern Mexico- Texas
corridor to the east. I have found, in undated excavated material from Sunny
Glen Cave, Texas (Anonymous, 1932), and in sites from Tamaulipas, Mexico,
(specimens and personal correspondence from McNeish, in 1955), extensive re-
mains of common bean pods and a few seeds. The lima bean pods found among
[Vol. 43
220 ANNALS OF THE MISSOURI BOTANICAL GARDEN
the vegetal remains from Tamaulipas strengthen the eastern Mexico corridor
hypothesis.
Lima beans are known from relatively few prehistoric Southwestern sites; these
are discussed by Steen and Jones (1941) and mapped by Carter (1945).* The
sites from which these beans are known are in the Verde Valley which shows
an admixture of Hohokam with other cultures. However, since the lima beans
are found neither in the Mogollon nor northern Anasazi sites, their relationship to
the Hohokam seems clear.
3. Anasazi:—
In the scatter diagrams the bean remains of the Mesa Verde pueblos are sep-
arated from those of the northern Arizona Anasazi to emphasize varietal diversity.
In this study only two types are recognized as present among the Mesa Verde
pueblos, and the larger number of types found in the ruins of northern Arizona.
The frequency of occurrence of the striped, violet common bean, type C11, in
both the northern Arizona and Mesa Verde Anasazi, and its rarity elsewhere
unifies these two regions with regard to bean distribution.
Collections of beans from Zion National Monument in southeastern Utah ex-
amined at the Museum of Anthropology, University of Michigan, provide evidence
which helps to clarify the relationship of Northern Periphery beans to those of the
Anasazi. Although the stratigraphy has been disturbed, habitation of the site
probably extended from Basketmaker II times until about 1150 A. D. (correspond-
ence from A. H. Schroeder to Volney H. Jones). Among these well-preserved
beans are to be found: types C11, C15, C12, C19a, and possibly C1—collection
numbers 232—236 respectively.
Jones" description (Brew, 1946) of charred beans (limas, common beans, and
others whose dimensions fell within those for Southwestern teparies) from the
Alkali Ridge site in teint Utah may indicate early agricultural influence
from the south. e the beans of Mesa Verde in southwestern Colorado—an
occupied area ee many cultural affinities with Alkali Ridge (Brew, 1946)—
are all common beans, none of which even in the charred condition could be
mistaken for lima beans or teparies, Northern Periphery and Anasazi beans do not
correspond completely. The presence of the small red, flecked bean, type C12 in
the Zion material (the small charred seeds found by Jones in the Alkali Ridge
collection may be of this type), links the Verde Valley (collection no. 162a) with
southeastern Nevada Virgin River sites (collection nos. 206, 206a), said by Reed
(1954) to be Pueblo II outliers of Anasazi culture, and with southeastern Utah.
The Verde Valley and Anasazi, then, appear to have had beans in common with
those of the Northern Periphery. Although Northern Periphery materials dis-
cussed in this paper are probably earlier than the Anasazi beans, which are mostly
4 The charred seeds cited by Carter as those of lima beans he Hodges site in south-central
Arizona are actually Canavalia ensiformis, as paipeir by the scr bilem which are about half the
length of the entire seed.
1956]
KAPLAN—PREHISTORIC SOUTHWESTERN BEANS 221
from Pueblo III, there is little to indicate a direct movement of agricultural
complexes to the Anasazi from the south by way of Nevada and Utah. However,
it does seem evident that the characteristic bean—C11—of the Anasazi did derive
from the Northern Periphery or, at least, although rare in the Anasazi, was
common to both regions in pre-pottery times.
In Basketmaker III times (Wormington, 1955) and later culture periods, many
traits point to a close relationship between the Northern Periphery and the San Juan
Anasazi. Among such evidence is the Central Mexican type of corn found in
prehistoric sites of southeastern Utah (Wormington, op. cit.). This type of corn
was described from the vegetal remains of Alkali Ridge by Anderson (1944) and,
with other artifacts, strongly suggest Mexican influence. However, neither
Nickerson (Wormington, 1955) nor Carter (1945) believed the corn to have
come directly through the Arizona Pueblo, Mogollon, or Hohokam areas, but both
authors consider a plains route as a likely alternative. The high row-number corn
of many early Southwestern sites, however, may indicate an early widespread
distribution of the Central Mexican corn. Evidence provided by bean remains
supports early southern rather than eastern agricultural influence.
The Existence of Bean Areas.—T wo theories may be advanced for the existence
of bean areas and their coincidence with culture areas in the Southwest.
One theory would involve an early introduction of beans to the Mogollon and
a later introduction to the Anasazi of varieties distinct from those found in t
Mogollon and unlike those found in the Hohokam, Thus the late introduction
to the Anasazi would not have come by way of either of these two regions and
therefore hardly could have come from Mexico directly except by way of eastern
Mexico and Texas. There are no dated remains to support this. An indirect route
from the Mexican-Central American center by way of the Caribbean, the Gulf
Coast, and eastern United States cannot yet be properly evaluated, but in the
opinion of the author this remains an unlikely route for reasons of cultural affin-
ities and plant geography.
A more likely theory would assume the early entrance and widespread distribu-
tion of beans throughout the Southwest but their adoption and culture by a
relatively small number of communities. This might explain both their presence
and their rarity in Basketmaker sites. The early beans thus could have been both
widespread and varietally more uniform than indicated by the later materials.
These later materials, as yet the only ones available to us, would have come from
a time postdating an intensification of bean culture, a time more likely to have
left bean remains simply because more people were growing them. During the
period between the introduction or introductions of beans to the Southwest and
the beginning of their more general cultivation, differentiation of types on the
basis of local or regional cultural preferences and regional climatic and edaphic
conditions would have occurred.
[Vol. 43
222 ANNALS OF THE MISSOURI BOTANICAL GARDEN
CONTEMPORARY BEANS
Early Spanish and later Anglo-American contacts brought new plants to the
Indian farmers, some of which became important and sometimes basic crops.
During the historic period contact among Pueblo peoples has been of more than a
casual or even trading nature, and introductions of Peruvian beans by way of
California and United States commercial varieties have occurred.
I. Hopi Beans:—
The assemblage of Hopi bean types (Table VI and fig. 9) appears to be most
closely related to that of the Verde Valley among the prehistoric groups. Beans
of the Verde Valley are the most diverse of the prehistoric groups and seem to be
composed of types which had come from the northern Anasazi, the Hohokam, and
the Reserve, New Mexico, Mogollon area. The mixture of types and species found in
the Verde Valley is not to be found in any one of the three great culture groupings
discussed in this paper.
In addition to the “old” beans which appear to have come down to the Hopi
from Verde Valley peoples, newer varieties which do not appear in the archae-
ological record have been adopted by the Hopi. These new varieties have had
several sources. The Pinto beans (Type C13) as known in the United States are
only a few variants of the Garrapata or tick bean group, which is highly diverse
and well distributed in northern Mexico (Collections of Oficina de Estudios
Especiales, Secretaria de Agricultura y Ganaderia, Chapingo, Mexico). The occur-
rence in the Verde Valley (Collection number 164) of the only prehistoric col-
lection of Pinto beans indicates a distribution limited to a single variety of the
Garrapata group as well as a rather restricted range of cultivation. The highly
important and widespread Pinto bean varieties now in use among most Southwest
Indians probably represent historic introduction of Garrapata varieties from
Mexico as well as a possible expansion of cultivation of the Verde Valley type
The arrival of these beans among the Hopi and other Indians may have been early,
with the Spanish missionaries, or with later Mexican contacts.
Two other types which have become important in the contemporary Southwest
but which are absent in prehistoric collections are the Pink and Bayo types (Types
C28 and C17, respectively) both of which are found among the Hopi. Hendry
(1918) states that the Pink bean is Chilean in origin, and that a translation of its
northern Mexican name is "White man's bean." The date of introduction of the
Pink bean is not known, and whether it arrived overland by way of Mexico, or
sailing vessel by way of California, is equally obscure. The Bayo types which are
found among the Zuni and the Hopi are said by Hendry to have come to California
prior to 1850 in sailing vessels from Chile and to have become well established in
the Southwest. These also have become widespread in western Mexico.
2. Zuni Beans: —
It is probable that the collections of Zuni beans studied are less complete than
those of the Hopi, and the same may be said for other contemporary Southwestern
1956]
KAPLAN—PREHISTORIC SOUTHWESTERN BEANS 223
groups. The Zuni types, without important exception, may be the same as the
Hopi types (See Table VI) of common beans. The fact that only one tepary
type and no lima beans occurred among the Zuni indicates less influence of the
Verde Valley, and ultimately Hohokam bean assemblage, upon the Zuni than on
the Hopi.
3. Rio Grande Pueblo Beans:—
From the scatter-diagrams it may be seen that the common beans of the Rio
Grande Pueblos are, as a group, smaller and less-variegated than the other common
bean groups (figs. 3-13). There is no information concerning prehistoric beans
of the Rio Grande peoples, but it may be that during the long Spanish rule of the
Rio Grande Pueblos these beans were introduced from Mexico.
4. Navajo Beans:—
It has been often said that Navajo agriculture was derived from the Pueblos,
particularly from the Hopi. There is nothing in the Navajo bean assemblage to
indicate that Navajo beans were not derived from the Hopi or, in the case of the
newer beans (e.g. the Pinto), from the same source as that of the Hopi.
5. Piman and Yuman Beans:—
Beans of the Arizona Piman and Colorado River Yuman peoples are most like
those of that part of the Verde Valley which was probably most influenced by the
Hohokam. There can be little doubt that the tepary beans of the Piman and
Yuman groups are directly related to beans of the ancient Hohokam. If lima
beans were a prehistoric Hohokam crop, as is believed by the author, one would
expect them to have persisted at least among the Pima. However, little remains
of the traditional crop varieties. It is extremely difficult to obtain corn of older
non-commercial types from the Pima, and beans other than the commercial types
are even harder to find. In addition to the difficult farming conditions of recent
drought years, it is probable that the importance of cash-cropping among these
people since the mid-nineteenth century has operated against retention of the older
crops, especially the poorly marketable tepary bean and dark-colored lima beans.
CONCLUSIONS
A complex assortment of varieties of Phaseolus vulgaris began to enter the
Southwest at least 2,000 years ago. Their appearance in the archaeological record
of the Mogollon in west-central New Mexico was prior to the introduction of
pottery, while in the Anasazi area of northern Arizona the bean record begins at
about the same time as the introduction of pottery, and in the Hohokam of south-
ern Arizona, subsequent to pottery. The existence of well-defined prehistoric bean
areas and the probability that beans were prepared for eating by parching and
grinding in early non-pottery times suggest that beans were cultivated in the
Southwest for a longer time than indicated by the now available archaeological
: [Vol. 43
224 ANNALS OF THE MISSOURI BOTANICAL GARDEN
materials. The parching and grinding of mature bean seeds may have been prac-
ticed by relatively few communities in a given area for a long time before culture
changes which are not now precisely known stimulated the general adoption of
beans. The early but restricted use would have afforded ample time for the
differentiation and selection of varieties characteristic of particular areas.
Lima beans probably entered the Southwest by way of both the Sonoran and
eastern Mexican corridors but at a later date than the common beans; their earliest
record is found in 13th or 14th century ruins. Lima beans remained restricted to
the central and north-central parts of Arizona and nowhere became as important
as common beans.
Phaseolus acutifolius appeared as an important bean only in extremely arid
southern Arizona, having come into use by the Indians of that region sometime
prior to 900 A. D. There is insufficient evidence as yet to locate the center of
domestication of the tepary bean; it may have been a domesticate of Sonoran
Desert Indians or of peoples in the southern part of Mexico.
The presence of P. coccineus in prehistoric Southwest cultures has not been
established, and its contemporary cultivation, limited to the Hopi, does not indicate
a position of importance for these beans.
The absence of change indicative of evolution under domestication in prehistoric
beans is attributed in part to the fact that the center of origin of the cultigens
represented is far removed from the Southwest, and in part that beans were culti-
vated previous to the period for which archaeological material is available.
Among the prehistoric cultures, large, variegated common beans with a high
frequency of a violet bean (Types C11 and C11a, which is distinctly or indistinctly
longitudinally striped) are characteristic of the prehistoric Anasazi cultural divi-
sion. e tepary bean is of greatest importance in the Hohokam region, while
the Mogollon culture division is characterized by large-seeded non-variegated
common beans and the absence of the striped violet bean of the Anasazi. Central
Arizona, including the Verde Valley, shows evidence of the bean influence of the
Hohokam and Anasazi and has the most varied assortment of beans, with common,
lima, and tepary beans present.
he prehistoric source of the contemporary Hopi beans appears to have been
the Verde Valley while the beans of the Pima and Papago tribes came from southern
Arizona. In addition to the prehistoric bean types, contemporary Southwestern
Indians have acquired beans from Mexico, Peru, and from commercial seed sources
in historic times.
SUMMARY
1. Collections of cultivated beans from contemporary Southwest Indian
peoples and bean remains from prehistoric sites in the same area were studied.
2. On the basis of external seed morphology and anatomy a series of cultivated
bean types is described in tabular and graphic forms. The types have about the
same taxonomic significance as named horticultural varieties. Thirty-two common
1956]
KAPLAN—PREHISTORIC SOUTHWESTERN BEANS 225
bean (P. vulgaris) types are recognized and described for the Southwest. Of these,
thirteen have been found only among prehistoric remains, ten occur only in con-
temporary collections and nine have been found among both prehistoric and con-
temporary collections. Eight tepary bean (P. acutifolius var. latifolius) types are
recognized from prehistoric and contemporary collections, and five lima bean (P.
lunatus), and two runner bean (P. coccineus) types are recognized and described
from contemporary collections only.
3. Using the criteria for classification of bean types as presented in this paper,
new materials may be compared to the existing Southwestern types. With this
basis for comparison bean collections from different cultural and temporal con-
texts can be related and the history of bean domestication, variation, and use
patterns may come to be better understood.
4. Bean areas are defined on the basis of distribution of the types; prehistoric
and contemporary divisions are made according to temporal distribution. The
areas outlined correspond to Indian cultural divisions and to climatic conditions.
5. It is suggested that parching and grinding of mature bean seeds was an
early method of preparation for food and that the role of beans in the diet was as
à source of protein to supplement animal and complement maize protein.
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gation of saline soils r. Soc ' Hort. Sci. Proc. 43:201—209.
Waugh, F e 6). Iroquois foals & nd food gege Canada Dept. Mines, Geol. Surv.
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; (1955). Mogollon culture siot to A. D. 1000. Amer. Anthrop. Assoc. Mem. 82.
Wie 21-242.
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KEY TO SCATTER DIAGRAMS, FIGS. 3-13
PHASEOLUS SPECIES SEED THICENESS COLOR VARIEGATION
P. =. 55 CM. WHITE EYE PATTERN
P. ACUTIFOLIUS st ad RED
k ei te / Oo, —U PEES
Ga | ) ( )
P, COCCINEUS S, 9 .76 = .85 BLACK FLECKED
FLECKS TENDING
BROWN TOWARD STRIPES
ABCISSA - SEED WIDTH sel ) AG)
NUMBER
26 YELLOW
o
ORDINATE - SEED LENGTA e
—
SHORTER LINE INDICATES
LIGHTER COLOR
EXPLANATION OF SCATTER DIAGRAM COLLECTION NUMBERS, TABLE VII
Fig. 3, Mogollon
5, b, 219; c, 224; d, 208; A 209; f, 212; g, 210; h, 218; i, 214; j, 220; k, 222; 1, 216:
m, I *« 225; o, 221; p, 217; q,
Fig. 4, Verde Valley, Arizona (Anasazi-Hohokam)
» 205a; b, 195a; c, 195, 196; à hei e, 195b; f, 185; g, 162; h, 201; i, 164; j, 195;
k, 162d; E 186; m, 204; n, 203; o,
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234 ANNALS OF THE MISSOURI BOTANICAL GARDEN
Fig. 9. Hopi
—43b; b—17a; c—42; d—16, 24, 29; e—31a; f—35; g—35a; h—31, 32, 34; ds oe
46, 55; k—49c; l—17b; m—19; n—57; o—14, 56, 58, 59; P ; q—26; r—27, s—33;
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KAPLAN—PREHISTORIC SOUTHWESTERN BEANS 235
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[Vol. 43
238 ANNALS OF THE MISSOURI BOTANICAL GARDEN
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Fig. 13. Colorado River Yunan
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240 ANNALS OF THE MISSOURI BOTANICAL GARDEN
EXPLANATION OF TABLE VII
The funt: giten are used:
rind — rind Foundation, Dragoon,
rm — Miss "pe Anthropology, ndo California, Berkeley, Calif.
C.N.H.M. — Chicago N Sage? History Museum, Chicago, Ill.
Denver — Denver Museum of Natural History, Zoe SE
Lab. of Anth. — Laboratory of deest Ab Santa N. M.
Mesa Vadis Mesa Verde Museum, Mesa e d Pak, Colo.
Mic Ethnobotanical eme ee of Anthropology, University of Michigan, Ann
Auber, Mich.
M.N.A. — Museum of A neis — 1 Ariz.
Tuc son — Arizona State Museum, Tucso:
Univ. — Department Ss Anthropology, University e kag Tucson, Ariz.
Commonly Soe abbreviations are used f of s F. B.M. refer to Pueblo
and Sie ker periods of the Pecos EH Zoe CR "Site, culture, and date
Collection numbers followed by an asterisk are collections inclu oq = pn scatter distant.
fis number eeds measured, if ed than the entire collection, esis under
"Number id s een of seeds." Pertinent notes on museum labels are SA Cad pre collection.
TABLE VI
LIST OF COLLECTIONS
Medial
imensio:
of seed (cm.)
i
ection a :
3 source and Site, culture and date SS Kë
E number GË Sá i
a AU re) E -á
Z E 3 HERI
Zo a EIE
3 M.N.A Havasupai. 1940 6 i| 78} 57] CH
M 1047/4559 fees vasu Canyon, | 3 51.581 44] TS
6a Berke. 3—7867 Prin Lower Colorado 1 J8| .55| .43| P. acutif olius
River. Con
6b »" 3-2976 Same as 6a 1 94| .56| .40 v
6c " 3-2982 "mom 1 90| eil .25 si
éd e 3-2983 eek E 1 82| .60| .38 is
6e * 3~2984 A 1 91 | .61| .37 d
éf xi 1~62242 Cave near Jacumba, San (1) $24.58 | 35 T
Diego Co., Calif.
Date unknown
6g > 1-62241 Same as 6f (1) Atl 69) 37 z
6h e 1~62240 EE (1) Dif .61 | .37 2
éi a 1-62243 NUN (1) .80| .55| 40 “i
S » 2~17461 Sia Binatone tas M. Contemp. | (5) 70) 353} 271 Li?
ped from cult.
8 M 2-17537 Dem Pueblo, N.M 2 1.06| .69| .43| T 4
tem
9 T 2-17419 Acomita Pueblo, N. M. (5) 147 | 78) 50; Cl
Contemp.
10* gs 2-17281 Pima, Santan, Ariz. (5) 75| .52| 39| Té
temp.
n" AS 2-17330 Papago, Ariz. Contemp. (5) 122| .77| .55| C 28
11a* * 2~17288 Pima. Ariz. Contemp. (5) Al 64) A5| TS Co
1956]
KAPLAN—PREHISTORIC SOUTHWESTERN BEANS 241
12* Berke. 2-17765 Gourd Cave, Nitair, L4] 0211641 C14
bie = Canyon, n. Ariz.
id
13 " 1776 Sam 4 1.44 | .74| .54| P. vulgaris
14* M.N.A. 771/1744 Hopi SC Surv. 1935 (10) 138) SEL x CH
15a* = Havasupai. Contemp. 4 1.56|1.10| .62| L 4
display cas
16* NA 771/1730 Hopi Crop Surv. 1935 (10) L42| 59811 6E]- C 11
17 " 771/1751 M a " " 10) 1.14| 861468 | -C 25
17a Berke. 2-17694 Hopi 4 1471 94 Ww €4
17b* de -17577 5 1,38] 791 46] C 26
18* M.N.A. 1046/3935 Contemp (10) 4,26 | 4761252 | 029
19* » 1/1810B Hopi Crop Surv (10) 1.38] .80| .60| C 3
20 T 2744 -— See Ariz. 1.44| .66| .48| P. vulgaris
21 3 725/2745 idi Crop Surv. Moencopi,| 3 1.60} .82] .75 d
Ari
241 S 771/1806 Hopi Crop Surv. 1935. (10) HE 339] 64
“Similar to robust”
23* 2 771/1809a Hopi Crop Surv. (10) 1.16] .81| .65| C 14a
** Jacob's cattle"
24* > 756/745 Hopi Crop Su 8 1.37| .80| .63| C 11
First Mesa, Ariz
25 2 771/1741 Hopi Crop S 4 n23| 77] 22| P vulgaris
"string bean"
26* d 771/1809B | Hopi Crop Surv. 1935 9 1371344 36) C 14
** Jacob's cattle"
27* x 771/1771 Hopi Crop 5 1221 72] 351 C2
28* E 771/1823B id e (5) 1.28| .80| .55| C 2
29* 1 1046/3939 Zeg Pres Ariz. (10) 148| .79| .59| C 11
30 x 771/1827 Hopi Crop S (10) 1.38| .74| .54| P. vulgaris
"ze aapa bean”
31* ? 771/1781 Hopi Crop $ (10) 1.60|1.08| .54| L 3
pes dion. lima"
3la*| Berke. 2-17591 Hopi 3 51 9| MILS
32* M.N.A. 756/B.729 | Hopi Crop Surv. (10) 1.58) 1.08] .55| L 3
33* ? 771/1835 malnm (10) 1.61|1.14| .52| L 4
d li
E " 771/1768A | Hopi Crop Surv. 10 1.57} 1.02] .56| L 3
“brown lima”
35* * 771/1825 Hopi Crop Surv (10) 1.42 | 1.00 | .53| L 2
“white lima”
35a Berke. 2-17592 Hopi Crop Surv 4 147|1.05| .54] L 1
36° N.A. 771/1727 | Hopi Cr (10) 1.03 | .68| .39| T 4
“mottled tepary”
e? e 771/1808 Crop Surv. (10) Olt 351 3581 T7
“black tepary'
38* A 756/B727 Hopi Crop p Se (10) 1.04] .68] 39| T 7
39* a 771/1807 M A (10) ie S61 Abi T 3
40° x 771/1758 wool E (10) 1.00| .66| .42| T 3
41* » 756/728 DI » 2 (10) J .64 42 T 3
42° * 771/1742 wo LM (5) 1.49 | .83| .47| C 29
“dark blue bean”
EA: * 771/1838 Hopi Crop Surv. (10) 1221 262 .$4| C 29
“Hopi blue dye bean”
43b*| Berke. 2-17635 Hopi Crop Surv. 2 £34) Sit 33} C50
44 M.N.A. 771/1769 "moo (10) 1.30| .76| .50| C 29
45% e 1046/3940 DE Td (5) 1.90|1.33|1.13| R 1
46* y 906/E250A Rech NR A 1.87|1.18| .94| R 1
46a a 771/1772 mou MOM 2 1.62| .80| .59| C 18
49 5 725/B2740 e Lu. * 3 197|121| 77| R 2
49b 3 854/2845 n Ne 4 2.20 | 1.40 | 1.07] R 2
49c* ad 906/E250B » » » 5 1.99|1.32| .98| R 2
[Vol. 43
242 ANNALS OF THE MISSOURI BOTANICAL GARDEN
TABLE VII (Continued)
12
dimensions
q of seed (cm.)
o
Collection ER tos
u source and Site, culture and date $'8 2 tp
A La >» 3 or type
E Sa 13815] 4
339 & ... E
e LARA
50* M.N.A. 771/1801A Hopi Crop Surv. (10) 1404 824 5601 Gir
yellow beans”
51 = 1046/3936 a n (10) 1.34] .74| .56 28
52 S 1046/4670 wo HW M (10) 1.42| .82| .50 17
53 A 771/1809D Honko e (10) 1.41 | .82| .54| P. vulgaris
54* x 771/1770 CO, X (10) 1.41| .80| .51| C 17
55* a 906/E250A DEE 3 05|1.28|1.14 | R 1
56* e 1046/3934 Won e 8 1581.78] S41 C ID
57* > 771/B1730 » M 13 138] 6L 56] C M
“purple string bean”
58* 771/B1731 Hopi Crop Surv. (15) 1351 76.571 013
59* = 771/1731 E ae (15) 1.45 | .82| .64| C 13
60 " 771/1837 Hg 9 n (15) 145| .82| .58| C 17
61” Berke. 2-17407 Zuni. Ojo Caliente, N. M. | 2 1.10| .69| .46| C 28
Conte oo
62* js 2-173588 "sb: N M. Cont (5) 1.04 7| 38| T4
63* Mesa Verde 417/345 cig Ve Mesa eng, 5 2| 74| .58| C 11
64* x 3324/275 cis Canyon, Mesa Verde. | 2 15-27 | 1 G2
64a y 3441/298 Square Se House, 3 127] 761581 C]
S i L a Verde. P. III
= Hi Tees as 64a 5d. 78 63| C 11
66 2848/171 Step House, Mesa Verde. 10 (8) [155 7 371 Cl
67" " 2488/86 Same ss 66 Y pint [1.42 | .82| .59| C 11
250
68a Amerind 992K Kukendahl, s.e. Ariz. : 0 e 1.20| .64| — | P. vulgaris,
near Nogales. 1200- 4 char reniform
1450 A. D. rag. subapiculate.
single cotyle-
Lë,
destroyed by
charring
68b 5 ap Same as 68a 4 frag. 89| .62| .43| P. vulgaris,
possibly C12
20 or 27,
tas
affected by
€.
69 » 1049 » » » 1 frag. 1.31 721]: -— Similar to 681
» charred
70a P/501B(2) | S. Ariz. 16th cent. 2 1.56| .97 | .53 | P. lunatus
charred
70b » » sk oe an DI 5 .80 .50 .38 Probably a
charred small-seeded
: P. vulgaris
70c » » » » » 1 55 .87 .55 P. vulgaris,
charred entire, ends
1956]
KAPLAN—PREHISTORIC SOUTHWESTERN BEANS
243
M.N.A. N.A.862
Berke.
DI
508/NA1764B.2
2-17417
2—17764
2-17291
2-17384
2-17731
2-17347
2-17290
2-17766
2-17720
2-17408
2-17347
2-17480
2-17480
2-217771 441A
2-173435
2-17709
Site unknown
Dead Man's Fort, Ariz.
P. II
gri dt M. Contemp.
ourd e Nitair
usi nyon, n. Asie.
P.
Pima. Santan, Ariz.
Contem
Zuni, N. M. Contem
Cave, S. Francisco ET
N. M. Undated
eed E Ariz.
Geen
Sam
Pima. ede Ariz.
Contem
Ge: Ss Nitair, Bes
on, n. Ariz II
Near ar en N N. M.
Contem
Zuni. Ojo Geng N. M.
temp.
Navajo. Jeddito, Ariz.
Contemp.
temp.
Turkey House, Casa
Grande, Ariz. Hoh
kam-Anasazi
Same as 99
Acomita Pueblo, N. M.
temp.
Navajo. Jeddito, Ariz.
Con: `
Truchas, N. M. Contemp.
io
>
1,14
.63
b
D
v
oo
Probably C 6
ike C 19a,
but smaller
ntire,
rounded ends
P. vulgaris
Ends flat
[Vol. 43
244 ANNALS OF THE MISSOURI BOTANICAL GARDEN
TABLE VII (Continued)
al
imensions
ER of seed (cm.)
$
i ak : DE Species
3 source and Site, culture and date EK á
a 5 > E or type
3 A "5 = E)
Z 23 |$ 3/3
Ze |i || P
106* Berke. 2-17434 Acomita Pueblo, N. M. (5) 1.32 | .81| .62| C 14
ntemp.
107* d: 2-17701 Chimayo, N. M. E (5) 1.22] .78| .60| C 22
108* Je 2-17326 Papago, ies nea (5) 80] 355] 9| E
Mex. bord. EE
109* i 2-174435 TEEN E N. M. (5) 1.00| .65| .46| T 3
ntemp.
110* ei 2-17327 Papago. Big Fields, Ariz. (5) 88| .65| .3| T 4
Contemp.
111* = 2-17389 Zuni, N. M. Contemp. (5) 1.12] .78| .64| C 142
112* " 2-17479 Cochi Pueblo, N. M. (5) 1.08| .69| .51| C 23
: Contemp.
114* : 2-17703 Cordola, N. M. Contemp. | (5) 1.28] .80| .60| C 13.
115 x " " ^ » (5) 1.11 | .72| .53| P. vulgaris
116* 3 2-17332 Papago, Ariz. Contemp. (5) 8 54| 41| T 6
117% Y 2-17230 Yuma, Ariz. Contem 4 Zei 452] 42| € 27
117a " 2-17728 " ags. T 6
"parched small Med P. | parched
ne ari is eaten thus o
118* = 2—17344 etus Jeddito, Ariz. (5) 34.83] 237 | <2
S weg.
eg " 2-17341 SEH as (5) 120473] 50[|[. C2
120 2-17390 A. OK Clm. N. M. (5) 1:27 82 801 C23
poe
121* og 2-17262 Cocopa, Somerton, Ariz. (5) 86| .60| .46| T 6
Cont
122" e 2-17249 Mohave, P » Parker, Ariz. (5) 84| .61| .47| T 6
123* o 2-17345 Maisch Jeddito, Ariz. (5) 1.24) 731 Sf] GH
ntem
124* 2-17778 Navajo, Twó =. Hills, (5) 116/74) 531 622
N.
125* : 2-17387 Zuni, Ojo Deeg NM | (5) 1.44] 93| .73| C 24
Contemp.
e = 2-17272 Havasupai, Ariz. Soop 3 1.20]..68 | .531| G6
ane $ 2-17232 tem 6) 1.41| .72| .53| C 18
28 2-17346 Navajo, — Ariz. (5) 88| .60| .45| T 6
Ws oen
au a ica Sam (5) 1.34| .77| .58| P. vulgaris
130 2-17310 Piss rise Ariz. (5) 76| .56| .41| Té
131 E 2-17754 Kiet Siel, Tsesi SCH 1.43 | .97| .62| P. vulgaris o
n. Ariz. charred frags. of
testa
131a M 2i Same as 131 85| .45| .43| P. acutifolius
132* ” charred ?
2-17248 Mohave, Parker, Ariz. (10) 89| .60| .46| T 6
133* " Kante
2-17235 Yuma Indians, (10) A61 127] A1 L8
134* 2: Ariz. Contemp.
2-17261 Cocopa, Somerton, Ariz. (5) 89| .62| M| T 4
Contemp. IE
1956]
KAPLAN—PREHISTORIC SOUTHWESTERN BEANS 245
135* Berke. 2-17261 Same as 134 (5) 10331 21] -AL F
136* = " (5) 261.651 381 1.4
137* g 2-17337 Papago, Ariz. Contem (10) $9] .60] AE. TS
138* 3 2-17756 Tusigoot, near Clarksdale, 5 0| .45| .32| P. acutifolius
., Verde Valley charred
view den ens
1392 T 2-17414 Acomita Pueblo, N. M. (5) 1.45| .90| .56| C 14
ontemp.
140* E 2-17342 Navajo, Jeddito, Ariz. (5) 131] 057-531 C 43
Contemp.
141 ” » Same as 140 (5) La dl. © 22 2
142* y 2-17318 Papago, Ariz., near (5) 1321::801:.491 C 13
Mex. bord. Conte
143* 3 2-17271 Havasupai, Ariz. Contenp (10) 95051 60] . MI T5
144" ES 2-17247 gcc veia Ariz. (5) 7| SU 35] T 9
145* y 2-17250 eem (5) 961 «601 0l T3
146* " 2-17401 Zuni, Ojo Caliente, N.M. | (5) 1.43] .76| .54| C 18
147* d 2-17267 Walapai, Pes Peach onam (5) a4] 59] 451 T 6
148* xi ," a (5) SV]. TS
149 xc 2-17233 uma Indians Ariz. (5) 1211:.97| 45 |. L 2
Con
150* x 2-17260 Cocopa, Somerton, Ariz. (5) 1. A TS
Cont
151* * 2-17405 Zuni, Ojo Celiente, N. M. (5) 1.32| .81| .60| C 11
iis : Contemp.
2-17409 Same as 152 (5) 1.23 | .82| .54| C 14
153* 1 2-17406 (5) 10521 861 Est
154* 3 2-17410 (5) 1.48 | .83 | .57| C 17
155 eg 2-17704 Cordel, N N. M. Contemp. | (9) 1.10] .71| .55| C 22
156* uf 2-17324 Papaso, B Big Fields, Ariz. (5) 1.26| .78| .54| C 13
157* z 2-17418 Acomita Stier N. M. (5) 1.23| .71| eil C 22
Con
158* a 2-17231 T" Tadians, Ariz. (5) .82 | .56| .41| T 6
159 M.N.A. 695/NA2519 ae Siel Tesi Canyon, 1 1.51 | .85 | .64| P. vulgaris
P. Il charred
160* y 621/405 ach) ‘Pueblo Little 1 1.14 | .72| .61| C 14
M18 "es 1000—1200
161 " 712/NA2520.32 Turkey pen. Tsegi Canyon | 6(3 im- |1.42 | .82| .61| ClorC ll
mature)
162* jx 692/ A86b eg? ouse, Verde 1 1.16| .78| .60| C 11a
Valley, pod 1100-
162a* » » Sam 1 BA 62 .48 C 12
164* A 514/1814A. pa Kee n.e. of H 91] .58| 34| C 13
Ré Flagstaff, Ariz. 1000—
1200 A. D.
165 " 561/NA739.18| Walnut Canyon, Cliff 3 39] 82) 58) C3?
Dwellings,
Sinagua, P. III
166 ^ — 583/117-U- ; 2 2.67 | 1.66 | 1.09 slis
3/117-U-26| No inf bere
167a | Lab. Anth. 30/1343 | Cave on S. Francisco R., 8 (5 .87| el .43| T6
adi arched)
167b » ” Same as 167a 1 .98 | .60 43 638!
parched
RA: oe
[Vol. 43
246 ANNALS OF THE MISSOURI BOTANICAL GARDEN
TABLE VII (Continued)
12
dimensions
$ 2
E of seed (cm.)
Collection 2 i :
9 source and Site, culture and date KK a. mes
2 a > 40 or type
E number 32 A gl? E
z EE W 1312
EI Ki i=] wen ud
Ze EE ele
168 365131. Vial 2 Sunny Glen Cave, 12 1.359) 317.61) C9
Big Bend, Tex
oe prehist
169 e AT : Sam 20 (10) | 1.38 sro] 31] C3
170 1 .07 0| .58| C 14a
171 Tucson Rm er Ari. Pt. of Pines. Ca.1qt. | 1.05| .64| .54| P. vulgaris
Y. Ca. 1280 A.D. (15)
charred
172 » d S Same as 171 30 |1.50| .80| en T
frag.
charred
(10)
173 » » » » » » 3 47 72 68 Lid
charred
174 " G.P.11071 Gila Pueblo excavations 3 .28 | .74| .58 $
175 a S 74 4 LITT EE es
176 sid G.P.16962 Bon 2 120} 22] 9741] —
177 ” Ll ” E » 2 E Fa .61 48
177a » » » c» 1 1.37 .90 56 Py Iunatus, Li?
178 » DI Te Ee 1 id -TE A8 Igaris
179 T G.P.11084 nas 4 1.46 | 78| .67| ”
181 T G.P.11055 — Se 2 1.54| .82| .64| ”
182 » sis» $e 1 1.14 75 47 de
183 z G.P.11032 s e ad 2 1511021 EL...
184 > G.P.11046 wis der ded 1 tati Cr ar
185* ve G.P.19691 "n MM 1 1.23 80| .47| C9
186* DI » » D DI 2 .8 .60 45 T 6
187 4 H.899 Hodges #793, s.e. Ariz. 6 -|1.70|1.10| .71| Canavalia
H.959 Pre 1300. Hohokam charred iformis
H.111 (Called “lima
H.1115 ” by
Carter, 1945)
188* M.N.A. 695/NA2519| Kiet Siel, Tsegi Canyon, H 1.38| .88| .62 11
n. Ariz. P. III, 1274- parched
1286 A. D.
189* " E Same as 188 (10) 1.47| .84| .57| C 19 or 198
parched
190* » » » 5 5 (11) £55 81 61 C 11
di d parched
191* » » wio (15) 1.25 71 Si Cz
ched
192* 2 » ur Obr. vg 1.35 83 55 C 14
parched
193* » » noc d. roy» 58 | 1.02 .54 CGS
parched
194a xd F "aon 16 1.58| .85| .58| C 19a
h
194b* » » nm nm » ee 1.40 .81 57 Gi
parched
195a* > 692/A85 Murder House, Lay 1.46 | .84| .58| C 18
ge Verde Valley, Fe?
P. IM
1956]
KAPLAN—PREHISTORIC SOUTHWESTERN BEANS 247
195b* M.N.A. 692/A85 Same as 195a 1 134] 22] 58] C 14à
195c* A x T te i 1 127] J1 | OU
195d E a qu 1 951 641.391 C 20 \
196 2 A E 2 145] 2] «MIL
197 " 712/NA2730.4| Dry cave, Tsegi-ot- 1 L24| .671 54] C 5 ?
: ES sosi. B.M.I. frag.
198 909/NA2520.13| "Turkey Cave, Tsegi 1 1491 28] 655] C 11
Canyon, Ariz.
P. I-
199* " 712/NA2519M97| Kiet Siel, Tsegi Canyon. 3612610351 Ci
z parched
200 z " M97b| Same as 199 1.48 | .82 | .55| P. vulgaris
charred
200a* R » M97c 2 133] 2215971 Ge
01 Tucson mae s ciae Verde 4 .94| .59| .42| C 20
Valley,
ca. 13
éi * Same as 201 1 RD EDAD Ti
Sé, = i : a 1 a2] IN T 3
sar E ee 3 .85| .54| 40| T 6
fos * E 8 144] oh [eos Ld
ance : : 1 Al] 29] .e1| C.H
6 Univ. Ariz. B.P.6004 Rock Shelter, Overton, ca. 100 .$1] 4535] 391] G I2
4590 Nev. P. II (15
i : parched)
206a Univ. Ariz. B.P.5990 € — Virgin R., .09| .71| .44| C lla
207* Denver 7146 nie se. Utah. B.M| 1 1.51| .75| .49| C 3
208* C.N.H.M. Eu pem w. cent. 3 15641] 741 311 C 19
ogollon
x = oes as as 208 2 155] .871 .359| C3
eed A 2 1.48 | .80| .56| C 5
n H noH 1 »1341 561.47] GE
2 Higgins Flat, nea 6 frags. | 1.50] .59| — | C 12
e, N. M. After charred
e > 00 Fa D. SE
See as 212 60 85| .44| — | P. acutifolius
(frags.
charred)
214* » (4)
Tularosa Cave, p Re- 2 1461 .98] .521 C 8
serve, N. M. M lon.
ini A 300 B. C.-1100 A. D.
Se 2 Same as 214 2 tél 711 .4] C 19
ns E » » » 4 1.29 | .8 51| C7
mat ££. Ow UM 4 99 .60 .48 [o 6
18 » » m» ^ 1 1.50| .88| .60| C 9
219* » Pe In a 2 1.635) 661 781 C 10
220* ” wo a 1 13501 701,581 C1
221* » ».- » 2 £111 91] .58 C 16
222* se SE d 19 1491 4 soi 05
223* » an ^» nm 11 1.19 .80 55 C 15
224* ” e i | 1 1.65 9 ,59 C3
225 Berke. — 4—4821 Ica Valley, Peru. Early 20 lbs. |2.47| 1.37] .61| P. lunatus
Nazca. 200 A. D. (5) Yellow-tan to
red-brown.
Black-eyed,
p
or
226 ^^ 4-7255B San Nicolas, near Supe, IN o6] 39] Ee
Peru. Middle Supe. E Dark purple
227 » » erën (5) 1.8] .92| .76| P. vulgaris
' Dark
——
[Vol. 43
248 ANNALS OF THE MISSOURI BOTANICAL GARDEN
TABLE VII (Continued)
nsions
3 of d (cm.)
Collection d Speci
9 source and Site, culture and date DS 2 pe
a be ka o» o or type
E number EXE le |as
z Eg ERCE
EI x] & . KZ
GEI a ee
228 Berke. ^ 4—7255B pee E (5) 1.50 | .80 | .62 | P. vulgaris
Red-brown
with blac eye
extending ant.
229 ig e EE (5) 1.51| .95| .64 | P. vulgaris
Red ground, ca.
cov: T
s black and
ant. mottle
230 js E fuse c Sev- 2.83|1.53| .75| P. lunatus
eral Yellow-brown
un- self, some with
dred black ant.
(5) mottle
231 E: 4-7269 EET 1 1.80| 1.26 | .93| P. lunatus
Black
232 Mich. 4445 c-3, c-4 Zion Natl. Mon., s.e. Utah | (19) 1235] 22] 56 | e E
4452 c-1, c-2
c-6
233 ^" sd eN 6 Same as 232 (14) BAG 1771530] Col
234 "o 4445 b uod ue (12) .88| .57| .38| C 12
"n : b
235 c-2, 4 deis ox 5(3) |139| .63| .63| C 19a
236 a ana 5, 6 Buren (2) 1.34| .70| en
1.49| 72 Giz
EXPLANATION OF PLATE
PLATE 1
Common and runner bean types. Slightly less than natural size.
ANN. Mo. Bor. GarD., Vor. 43, 1956 PLATE 1
S I9 0 196
KAPLAN—PREHISTORIC SOUTHWESTERN BEANS
ANN. Mo. Bor. GarD., Vor. 43, 1956 PLATE 2
KAPLAN—PREHISTORIC SOUTHWESTERN BEANS
[Vol. 43, 1956]
KAPLAN—PREHISTORIC SOUTHWESTERN BEANS 251
EXPLANATION OF PLATE
PLATE 2
T 1-T 8. Tepary types; L 1-L 5. lima bean types. Seeds slightly less than natural
size.
A. Dried pods and beans, type C11, Mesa Verde, Colorado. Pueblo III.
B. Charred beans, in situ. Point of Pines, Arizona.
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(0 Missouri Botanical
Annals
of the
Missouri Botanical Garden
Vol. 43 SEPTEMBER, 1956 No. 3
A REVISION OF RAUVOLFIA WITH PARTICULAR REFERENCE TO
THE AMERICAN SPECIES*
ARAGULA SATHYANARAYANA RAO**
Rauvolfia is one of the apocynaceous genera that long has awaited a taxonomic
revision. Since the inception of the genus by Plumier! in 1703 with two species,
the literature has accumulated an abundance of novel specific epithets, based fre-
quently on insufficient material and often due to misconceived synonymy. The
‘Kew Index’, including all the supplements, lists about 175 names for the world,
_ While the Kew and the Gray Indices together account for about 90 names for the
New World alone. An inadequate comprehension of the generic characteristics
also has led to the proposal of new genera, such as Opbioxylon, Cyrtosiphonia,
Dissolaena and Heurckia, to include the Asian and New Caledonian Rauvolfias.
Bentham and Hooker correctly recognized the synonymy of these genera with
Rauvolfia. Plants belonging to other genera, and even other families, in the past
have been described as Rauvolfias. Thus Ruiz and Pavon? described several species
of Citharexylum, a genus of Verbenaceae, as species of Rauvolfia.
Alphonse de Candolle,3 in his treatment of Apocynaceae for the ‘Prodromus’,
included 23 species of Rauvolfia and, for the first time, provided lists of dubious
and excluded species, even though he maintained Ophioxylon as a distinct genus.
umann’s* much later account for the ‘Pflanzenfamilien’ recognized about 45
species for the world, but the taxonomic treatment necessarily was brief and
sketchy. Pichon® estimated about 110 species for the world in a brief synopsis of
P mier, C. Plant. Am. Gen., p. 19. 1703.
Ruiz & Pavon, Fl. Peruv. et Chil. 2:26. 1799.
DC. in D
SE Schum. in Engl. & Prantl, Nat. Pflanzenfam. 41-2:153-154. 1897.
Pichon, M., in Bull. Soc. Bot. Fr. 94:32-39. 1947.
——
* An investigation carried out in the graduate laboratory of the Henry Shaw School of Botany of
Washington University, and submitted as a thesis in partial fulfillment of the requirements for the
ree of doctor of philosophy.
** Department of Botany, University of Toronto, Toronto, Canada.
Issued October 31, 1956.
(253)
[Vol. 43
254 ANNALS OF THE MISSOURI BOTANICAL GARDEN
the genus, and included an analytical key and brief Latin diagnosis of the fourteen
sections. Pichon's treatment has badly suffered from the fact that he had not
examined more than half the species which he listed. Apart from these, no com-
prehensive monograph of the genus for the world exists.
As far as the New World species are concerned, the most outstanding account
is that of Mueller-Argoviensis.? This, however, was limited to about 11 species
of Brazil. Markgraf's” essay on the relationships of the American Rauvolfias was
more comprehensive and included a discussion of the geography and taxonomy of
the tropical American species and provided a E to distinguish the species.
He recognized 34 species, including 7 new of his o
Like several other related members of the BE Plumeroideae, several
species of Rauvolfia contain alkaloids of considerable pharmacological importance.
The Asian Rauvolfia serpentina recently has attained phenomenal publicity in this
regard. This naturally has resulted in the extension of interest in these plants to
a far wider informed public
The present revision of the genus in the New World was begun in September
of 1954. It is based on a critical study of more than 2,000 herbarium specimens,
and also a handful of plants grown in the greenhouse of the Missouri Botanical
Garden. Even though the living plants did not yield any statistically significant
data, they played a very important role in suggesting the growth and variation
pattern of the species concerned.
Fully realizing that the American Rauvolfias are only a small cluster of the
world species and cannot therefore be studied entirely by themselves I have given
some time to a critical evaluation of Pichon’s fourteen sections. I have examin
all the specimens which he had for study, and more, although it must be confessed
that I too have not been able to examine material for all the species. Any authori-
tative and detailed presentation of the taxonomy of the extra-American Rauvolfias
will have to await more extensive study.
I have reduced the fourteen sections of Pichon to nine, the American Rauvolfias
comprising two of these sections and including 34 species, grouped under four
series and three subseries.
HISTORY OF THE GENUS
The Minim monk and pioneer Caribbean botanist, Charles Plumier, founded the
genus Rauvolfia in 1703 to commemorate Leonhart Rauwolf, a German drug-plant
collector who had widely traveled in the eastern countries. Plumier briefly diag-
nosed two species: Rauvolfia tetraphylla angustifolia and Rauvolfia tetraphylla
latifolia.
Linnaeus,? in adopting the genus, described one species and held another simply
as a variety. Thus originated a confusion concerning the identity of the two
ë Muell.-Arg., J., in Mart. Fl. Bras. 61:30—34. 1860.
ER F., in Fedde, Rep. Spec. Nov. 20:111—122. 1924.
ort. Cliff. 75 1737.
1956]
RAO—REVISION OF RAUVOLFIA 255
original species of Plumier. Rendle,? who was fortunate in having all the relevant
original material at the British Museum, has related the entire story and has ably
untangled the complex question of synonymy. The description of Rauvolfia tetra-
phylla in the ‘Hortus Cliffortianus’ was based on Ehret’s drawing of a plant that
Linnaeus had seen growing in the Chelsea Garden (pl. 3, fig. 3). Linnaeus wrongly
equated this with Rauvolfia tetraphylla angustifolia of Plumier and considered
Plumier’s second species merely a variety. He repeated this error in the first edition
of the ‘Species Plantarum’. This must have been due evidently to the very brief
diagnosis of Plumier on which Linnaeus had to depend in deciding synonymy of
his Rauvolfia tetraphylla. In 1775 Burmann's edition of Plumier’s ‘Plantarum
Americanarum Genera’, containing illustrations of both of Plumier’s species, was
published (pl. 3, fig. 1). Jacquin, apparently unaware of Burmann’s work, pub-
lished in 1760 three names: R. nitida, R. hirsuta, and R. tomentosa, and later, in
1763, gave descriptions for them with an illustration for R. tomentosa (pl. 3,
fig. 2). His R. nitida was synonymous with Rauvolfia tetraphylla angustifolia of
Plumier, and R. hirsuta and R. tomentosa were conspecific and identical with Rau-
volfia tetrapbylla latifolia of Plumier and Rauvolfia tetraphylla of Linnaeus.
Apart from Jacquin’s illustration of R. tomentosa, I have studied a specimen from
Jacquin’s herbarium at Vienna, annotated in his own handwriting as Rauvolfia
tomentosa, which proves the correctness of the above dispositions.
The fact that Linnaeus had become aware of his earlier mistake and that he
had also seen the descriptions and illustrations of the two species in Burmann’s
edition of Plumier is indicated in the second edition of the ‘Species Plantarum’. In
this Linnaeus described £wo species: Rauvolfia nitida and Rauvolfia canescens.
Linnaeus was correct in citing Rauvolfia tetraphylla angustifolia of Plumier in
synonymy with Rauvolfia nitida, and Rauvolfia tetraphylla latifolia of Plumier
with Rauvolfia canescens. In establishing this new name did not realize that
it was conspecific with his earlier Rauvolfia tetraphylla.
Jacquin’s Rauvolfia hirsuta was allegedly based on the reference to the same
specimen of Patrick Browne on which Linnaeus based his R. canescens. Hence it
is a just cause for surprise as to why Linnaeus should have ignored Jacquin’s prior
name, even if he had considered the Patrick Browne specimen (pl. 3, fig. 4) dis-
tinct from the one of Ehret's drawing on which his R. tetraphylla was founded.
The Linnaean name R. tetraphylla has priority over Linnaeus later name of R.
canescens, and is the equivalent of Plumier’s second species Rauvolfia tetraphylla
latifolia. R. nitida of Jacquin is equivalent to Plumier’s first species Rauvolfia
tetraphylla angustifolia.
Another question of some concern is that of orthography. Plumier used the
Latin version of Rauwolf’s name and called the genus Rauvolfia. The fact that
Rauwolf himself was inconsistent about the spelling of his name, once spelling it
Leonhardt Rauwolff and at another time Leonhard Rauwolf, should not concern us
? Rendle, A. B., in Proc. Linn. Soc. Lond. 149:106. 1937. This contains references to all the
pertinent literature.
[Vol. 43
256 ANNALS OF THE MISSOURI BOTANICAL GARDEN
very much here. Linnaeus himself was faithful to the original spelling in all his
editions of the ‘Genera’ and the ‘Species Plantarum’. However, Burmann”? ha
used three different spellings in quick succession, even in the same page, thus:
Rawolfia, Rauwolffia, and Rauwolfia. In the seventh edition of Linnaeus’ “Genera
Plantarum’ under Reichard, the name was spelled Ravwolfia in the text and Rau-
wolfia in the index. The eighth edition under Schreber used Rauwolfia and there-
after Willdenow, Alphonse de Candolle, Bentham and Hooker, and Schumann have
all used the altered spellin
Woodson,! who at first used the original spelling and later came to prefer the
corrected” spelling of Rauwolfia, recently has recounted the story of inconsistency
in the spelling.!? In that account, however, he has clearly indicated that though
the spelling Rawwolfia is contrary to the requirement of the International Code of
Nomenclature, yet he preferred the altered spelling in view of popular usage in
that form and particularly because the publication was meant for non-botanical
readers.
As mentioned earlier, the spelling Rauwolfia transgresses Article 82 of the
International Code of Botanical Nomenclature.13 In this work the original spelling
of Plumier, Rauvolfia has been consistently used.
MonPHOLOGY AND ANATOMY
Habit: Plants of Rauvolfia, except R. purpurascens of Panama, which Stand-
ley has described rather incredibly as a vine, are undershrubs, shrubs, or trees.
Most of them are moderately good-sized shrubs, only a few being either puny under-
shrubs or big trees. The smallest one, appropriately named R. nana, of northern
Rhodesia in Africa, attains a height of only 15 cm.
Amongst the American species, almost all species belonging to section RAU-
VOLFIA are shrubs scarcely exceeding a height of about 2 m. The majority of species
of section MACROVOLFIA, on the other hand, are sturdier and more robust shrubs
or trees. R. linearifolia is the smallest, hardly reaching a height of 5 dm. Amongst
the arboreal species, R. praecox is the tallest, reaching a height of about d m.
Most of the other species have an intermediate habit that can be described at times
as shrubs, and at other times as trees.
All the plants are lactescent, particularly in the aerial parts. In R. tetrapbylla,
R. littoralis, and R. serpentina, roots at any stage of their growth have failed, in
my experience, to show latex exudation, when cut. But all the other parts above
the hypocotyl exude a milky latex on wounding.
Brancbes: Branching is usually whorled, often rather densely so. The branches
are usually terete, rarely angular or even winged to some extent, as in the African
1, Burmann, ys DN Plant. em Geng er 252.1755) $
11 Woodson, R. E., in Ann. a 1930; in N. Amer. Flora 29%:134-138. 122%
"Serien et T goo e Lë Chemistry id tiere Little, Brown & s
Boston. In
"inge Code E Bot. Nomencl. p. 43. 1952.
1956]
RAO—REVISION OF RAUVOLFIA ESF
R. macrophylla. Except in several species of section RAUVoLFIA, where the young
twigs are variously pubescent, the branches are glabrous, and in most species
lenticellate to varying degrees.
Nodes: The nodes are emphatically marked by the pectinate glands that usu-
ally are present in the leaf axils, as well as by leaf scars. Except in members of
series LATIFOLIAE of section MACROVOLFIA, where the terminal nodes are very
short and condensed, unlike the more distantly spaced nodes further down, the
nodes of all the species are fairly evenly and distantly spaced. However, a con-
densation of the terminal nodes is evident also in R. sellowii, a member of series
ANGUSTIFOLIAE. A similar situation exists in the Hawaiian species and in the
African R. macrophylla and R. caffra. This condensation of the terminal nodes,
often accompanied by foliage only at the tips of twigs, gives a characteristic
appearance to these species.
Leaves: ‘The leaves are whorled, the number in a whorl being variable in dif-
ferent species and sometimes even in the same species. There is also an inconsistency
in the shape, size, and indument of the leaves, even in the same species. In spite
of this, leaves offer some fairly reliable characteristics for the taxonomic diagnosis
of the different species.
The leaves of a given whorl are more or less anisophyllous. They are petiolate
or sessile, simple, entire, and ovate, elliptic, lanceolate, obovate, or oblanceolate in
outline, The venation is pinnately reticulate, the secondary veins and the vein
network extremely variable in clarity on the two surfaces. The secondary veins
are mostly arcuate and often unite at the margins to form a marginal vein. How-
ever, in species such as R. nitida, R. sarapiquensis, and R. pentaphylla the secondary
veins are transverse or almost so.
The leaves are either membranaceous or coriaceous. All the species included in
section RAuvoLriA have membranaceous leaves, whereas the species in the other
sections have leaves of variable texture, from membranaceous to coriaceous. The
two surfaces usually are opaque, but in species such as R. cubana, R. salicifolia, R.
braecox, and R. nitida, the upper surface is lustrous and the lower opaque. Though
there is a certain amount of difference in the emphasis of the green color on the
two surfaces, the upper being usually dark and the lower varying shades of light
green, and although Markgraf!* has used this as a key character, my experience
with fresh and dry leaves of identical species has shown that no reliance can be
placed on this character. The leaves are glabrous except for some species of section
RAuvoLrFia, where one or both surfaces exhibit pubescence to varying degree.
The leaves are variable in size. Like certain species of Asia and New Caledonia,
the three Cuban species, R. cubana, R. salicifolia, and R. linearifolia, have rather
narrow leaves, the last having the narrowest leaves of all the species. With the
exception of a few species which may possess rather narrow leaves at times, most
of the species have comparatively broad leaves.
——— \
14 Markgraf, F., in Fedde, Rep. Spec. Nov. 20:113-118. 1924.
[Vol. 43
258 ANNALS OF THE MISSOURI BOTANICAL GARDEN
The petiole varies considerably in length. In most of the species it is short and
becomes obscured by the gradual decurrence of the lamina. In R. fraecox and R.
sellowii, however, the leaves have fairly long and quite distinct petioles. The
petiole usually is entirely glabrous, but in species of section RAUVOLFIA it has also
either a sparse or dense glandular covering.
Stipules and Glands: As in other members of Apocynaceae, the nodal region
in Rauvolfia is marked by a distinctive ring, and the leaf axil is provided with a
number of pectinate glands. ere has been some difference of interpretation of
the nature of these glands. The two observers holding diverse opinions on t
score, Woodson and Pichon, have based their conclusions on an observation of more
extensive material than was possible for me. Woodson!? has held, following
Gluck's original interpretation, that these appendages are "doubtless in the cate-
gory of stipules”. Pichon,!? in a lengthy discussion of the stipules and glands of
Fig. A. Rauvolfia littoralis. Twig showing stipules and glands.
APocyneceie, has concluded that the glands are in the nature of foliar appendages
"sui generis" and are distinct from the stipules. There is an obvious bc in our
knowledge of the morphology and physiology of these glandular appe
I have studied the nodal region in living plants of R. viale K ere,
and R. serpentina with special interest. In the first two species, in addition to the
numerous pectinate glands in the leaf axils as well as on the petioles, there are
distinct stipules at each node. These stipules are interpetiolar in position, subulate
or linear, and are about 2 mm. long. They are deciduous and leave a minute scar
on the node (text-fig. A). In R. serpentina, the leaf axil does not exhibit many
glands, but lateral to the petiole minute subulate structures are clearly observable
which are obviously stipules. I have noted similar interpetiolar stipular structures,
zx 15 Woodso & Moore, i in Bull. Torr. Bot. Club 65:148. 1938.
Pichon, M, in Bull. Mus. Nat. Hist. Nat. II Ser. 1:467-473. 1949.
+
1956]
RAO—REVISION OF RAUVOLFIA 259
in addition to the ubiquitous gs in young shoots of R. cubana preserved in
alcohol. Furthermore, in all members of series LATIFOLIAE, there are prominent,
deciduous bud-scales which keng are stipular structures. In Rauvolfia at
least, examination of both the glands and the stipules leads to the conclusion that
the two appendages, although associated, are distinct.
Inflorescence: The inflorescence is terminal. There is no recent discussion in
the literature on the position of the inflorescence either in Rauvolfia or any of its
close relatives in the family. However, some observations and interpretations have
n made in certain genera of the allied family Asclepiadaceae. Bugnon,!' in a
study of the inflorescence in Gomphocarpus fruticosus, summarized the three prin-
cipal views on the position of the inflorescence. The three views are: (1) The
inflorescence is a sympodium. (2) The inflorescence is an axillary branch. (3)
Rauvolfia ioi mie Inflorescence as it appears
di ud as it is interpreted (right).
The inflorescence is one branch of a dichotomy. On the basis of his studies, Bugnon
has rejected all three views and has put forward a rather ambiguous interpretation
of his own. Holm,}8 on the other hand, in his studies on the inflorescence of species
of Sarcostemma, has preferred the classical interpretation of Payer and others, and
has held the inflorescence to be a sympodium.
have studied carefully the development of the inflorescence in living plants
of R. tetraphylla. The vegetative shoot sooner or later ends in a simple dichasium.
By the time the terminal flower-bud is ready to blossom, one of the axillary buds
in the subjacent leaf whorl becomes active and soon develops a vigorous shoot. In
this process the originally terminal cyme is gradually pushed to one side and the
axillary shoot increasingly veers to assume a terminal position (text-fig. B). The
inflorescence thus becomes interpetiolar in position after this spatial transposition.
Holm has pointed out that the lateral sliding of the inflorescence of Sarcostemma
Je la Bugnon, F., in Bull. Soc. Bot. Fr. 102:105-114. 1955.
18 Holm, R., in Ann. Mo. Bot. Gard. 37:485—486. 1950.
[Vol. 43
260 ANNALS OF THE MISSOURI BOTANICAL GARDEN
occurs alternately at successive nodes and as the result the inflorescences appear
to be in two rows. In R. tetraphylla, however, the situation is further complicated
by the leaves being in whorls of four. There is no constancy or visible pattern in
the order in which axillary buds at any given node develop, or the relationship of
the inflorescence to any one axillary bud in the whorl. Commonly, however, only
one of the axillary buds in the whorl is developed, be it associated with a smaller
or a larger leaf. Yet occasionally two, three, or even all four axillary buds develop,
in which case the terminal nature of the inflorescence is left in no doubt whatever.
Although these observations have been made in only the living plants of one species,
a careful study even of herbarium material of the rest of the species confirms the
view that the inflorescence is primarily terminal.
The most generalized type of inflorescence in Rauvolfia is the dichasium.
Woodson,!? in a discussion of the inflorescences of Apocynaceae and their phylo-
genetic significance, concluded that a reduction has taken place in the originally
complex inflorescence. This trend is also evident in the inflorescence of Rauvolfia.
The most complex type of inflorescence is the "aggregate dichasium". A wide
range of variation is exhibited within this type alone, in numerous species, by differ-
ences in the degree of branching of the peduncle and the relative proportions of
the branches. A further contributing factor is the nature of the ultimate cymule,
which may be umbelliform or corymbiform.
The reduction trend is perhaps best exemplified by the species of section Rau-
VOLFIA. A species such as R. littoralis has much-branched, multiflorous inflores-
cences. On the other hand, a closely related species, R. tetraphylla, possesses few-
flowered, mostly unbranched inflorescences. The fact that even within a single
species such as R. tetraphylla or R. ligustrina the two extremes from a much-
branched, multiflorous, to an unbranched, pauciflorous condition is not uncommon
contributes further to the idea of reduction in the evolution of the inflorescence.
Incidentally, much value was formerly attached to the inflorescence patterns as
key characters, but the variation noticed in this study renders them valueless. A
pauciflorous inflorescence is also present in species such as R. macrantha and
R. paucifolia, both belonging to the essentially pluriflorous section MACROVOLFIA.
The peduncle in different species is variable in length and in degree of branch-
ing. It is usually glabrous, but in the species of section RAUVOLFIA, it may be
variously pubescent.
The pedicel is usually distinct and of varied length. In some of the African
and Asian species, however, it is much reduced or often even wanting.
Flower: As in the other genera of Apocynaceae, the flower in Rauvolfia is
regular, bisexual, tetracyclic, and pentamerous, usually pedicellate, rarely sessile as
in some African and Asian species. It is relatively small, except in several species
of sections MACROVOLFIA, OPHIOXYLON, and OPHIOXYLANTHUS. In several
species the flowers often exude a pleasant odor.
19 Woodson, R. E., in Ann. Mo. Bot. Gard. 22:1-42. 1935.
»
1956]
RAO—REVISION OF RAUVOLFIA 261
Calyx: The calyx is usually green, synsepalous, and mostly campanulate with
the lobes usually imbricate in aestivation. The lobes vary considerably in size,
shape and texture. In all the American species they are considerably shorter than
the corolla. They are ovate to lanceolate in shape, with the apex varying from
acute to obtuse or rounded. They are eglandular within and usually glabrous
without, except in section RAUVOLFIA where the calyx is variously pubescent with-
out. In some species of section MACROVOLFIA, they are either ciliate or glandular-
dentate at the margin.
Corolla: The corolla is sympetalous and of varied shapes: urceolate, campanu-
late, salverform, or infundibuliform. It is usually white, but some of the species
show either lilac or rosy hues or are even variously spotted.
The corolla-tube is generally cylindrical, sometimes constricted at the throat,
and sometimes dilated. It is usually glabrous without and variously pubescent
within, The extent of pubescence in the tube, the relative proportion of the tube
to the corolla-lobes, on the one hand, and to the calyx-lobes, on the other, and the
region of constriction and dilatation, all offer fairly reliable taxonomic criteria.
The corolla-lobes are sinistrorsely contorted in aestivation. They are ovate,
obovate, elliptic or oblong in outline with acute or obtuse apices. In species of
section ENDoLoBus, the apices are inflexed in the bud.
Stamens: The stamens are epipetalous, five in number and fairly uniform in
all the species. The anthers are ovate, acute or apiculate, and slightly varying in
size in different species. They are free from each other and the stigma. The
thecae are fully fertile and enclose free spherical or subspherical, triculpate pollen
grains. The connective in species of sections MACROVOLFIA, OPHIOXYLON and
OPHIOXYLANTHUS appears to be projecting in the form of a conical hump on the
dorsal surface of the anther, just at the point of attachment with the filament.
There is a distinct filament in all the species, though it may vary in length from
species to species.
Pistil: The pistil is superior, bicarpellary, the ovary being variously syncarpous
or apocarpous. The ovary is of various shapes: globose, cylindrical, ovoid or obo-
void. Each locule encloses one or two anatropous ovules, on an axile or ventral
placenta.
The style is simple, linear or columnar, glabrous except in the African R.
vomitoria, where it is villous at the base.
The stigma-head is prominent and variously shaped: depressed-capitate, calyp-
triform, tympaniform or sub-mitriform. Miers?? was the first to use the term
“clavuncle” to describe the stigma-head of Apocynaceae. The stigma-head possesses
a basal ring or collar, a contracted or expanded median region, and a distal indusium
encircling the biapiculate apex. Pichon has used the term “strophium” to indicate
the median region of the stigma-head. Much of the variation in the shape of the
stigma-head in the several species is brought about by a difference in the relative de-
?? Miers, J. Apoc. S. Am. p. 2. 1878.
[Vol. 43
262 ANNALS OF THE MISSOURI BOTANICAL GARDEN
velopment of the three regions and the presence or absence of hairy induments on
them. I have noticed in R. tetraphylla a frequent clustering of masses of pollen
grains around the base of the apiculate tip and in the depression formed by the distal
indusium, an observation in consonance with that of Miers. However, it is not
possible to conclude from this observation whether the depression itself is truly
receptive or if it is the apiculate tip that is so. As there is some confusion in
the precise use of the term “clavuncle”, I have preferred to use the simpler term
stigma-head.
The shape of the stigma-head differs with the age of the flower and becomes
distorted in the pressed herbarium specimens. This renders the use of the stigma-
head valueless as 3 taxonomic character.
The disc is annular or cupuliform, usually shorter than the ovary which it
encircles. It is usually entire.
Woodson,” in an elaborate study of the floral anatomy of 39 genera and 60
species of Apocynaceae, commented on the inconsistency in the vascular pattern,
particularly in the calyx of R. nitida (referred to as R. tetraphylla in that paper).
He also pointed out the carpelloid nature of the disc or nectary.
In a taxonomic treatment it is not my intention to digress in an account of the
floral anatomy. However, I was able to make a few observations concerning
flower-buds of R. tetraphylla and R. cubana which Dr. Woodson had kindly pre-
served for me. After a study of whole cleared mounts, hand and microtome
sections of flower-buds of these two species, no inconsistency in the vascular pattern
of the calyx was observed. In both there is a ring of five sepal traces, each of
which gives off two lateral traces just prior to entering the calyx-lobe. The corolla
and the stamens each have a typical and consistent vascular pattern. The traces
to the disc emerge as branches of traces going up to the two carpels, bearing out
the carpelloid nature of the disc. Also, I could find no evidence for Boke’s”
interpretation of the corolla-tube as being partly receptacular in origin.
Fruit: The fruit of Rauvolfia is a drupe. It is variously syncarpous or apo-
carpous. The calyx is persistent on the fruit in all the species and the pedicel
tends to become stouter in the fruit. The fruit is comparatively small, syncarpous
and globose in all species of section RauvoLria. In the species of other sections,
however, it is rather large and variously syncarpous or apocarpous. The biggest
fruit is that of the Amazonian R. paraensis, about 40 mm. in diameter, while the
smallest perhaps is that of the Bahian R. blanchetii, about 6 mm. in diameter.
The fruit is either rounded or somewhat flattened, The two lobes in hemi-
syncarpous fruits or the drupelets in the apocarpous fruits differ considerably in
their mutual relationship as well as in shape. They may either be parallel or
divergent to differing degrees. These characters offer trustworthy taxonomic
criteria for specific diagnosis. With the gradual ripening of the fruit there is a
change in the color of the pericarp from green to red and black. This is true for
almost all the species. The exocarp is usually thin and membranaceous. The
$, Woodson & Moore, in Bull. Torr. Bot. Club 65:147. 1938.
Boke, N. H., in Am. Jour. Bot. 35:422-423. 1948
1956]
RAO—REVISION OF RAUVOLFIA 263
mesocarp is fleshy, rarely slightly fibrous, and the endocarp stony. R. linearifolia,
the dwarf undershrub of eastern Cuba, has fruits with unusually thin pericarps
resembling superficially a follicle more than a drupe. The stone is usually scle-
rotic, with a smooth or rugose outer surface and a smooth inner surface. It is
generally ovoid, with a stout basal region and a rather flattened distal region. The
stone also offers some reliable key characters. In all the species there is a tendency
towards the abortion of one of the carpels during development. Although a con-
siderable number of extra-American species possess apocarpous fruits, R. sellowii
is the only American species on which they occur.
Seed: The seed is albuminous and is usually symmetrically ovoid. In R. nitida
of section MACROVOLFIA and in all the species of section RAUVOLFIA the seed is
slightly gibbous. This is true also in several African and Asian species. The
seed-coat is membranaceous and easily separable from the endocarp, on the one
hand, and the albumen, on the other. The albumen is carnose, entire, and fairly
abundant.
The embryo is typically dicotyledonous, either erect or slightly to strikingly
arcuate. Usually it is about as long as the seed, with the terete hypocotyl about
equal to the usually ovate, obtuse cotyledons.
Seedling: I have observed germination in the seeds of R. tetraphylla, R. lit-
toralis, and R. serpentina. Owing to the sclerotic endocarp, they require from ten
to thirty days for the first signs of germination. e radicle emerges first, and
three to four days later the crook of the plumule appears and soon becomes erect
and disentangles the cotyledons from the still-persisting endocarp and seed-coat.
he cotyledons expand and serve as the first pair of green leaves for the seedling.
In R. tetraphylla and R. littoralis the cotyledonary leaves are ovate and obtuse, but
in R. serpentina they are orbicular, rounded, or slightly emarginate. Only pairs of
decussate leaves are produced in the subsequent two or three nodes of the actively
growing seedlings. The fourth node, however, almost invariably exhibits the
characteristically whorled phyllotaxy of the adult plant. The plants of R. tetra-
bbylla appear to be precocious and flower when they are about four months old.
Anatomy: The literature contains very little reference to the anatomy of
Rauvolfia. Metcalfe and Chalk?? have remarked briefly on the anatomy of Apo-
cynaceae. The recent importance of the roots of Rauvolfia, from the point of
view of the pharmacognosists, has required a precise knowledge of their anatomy
for accurate determination of the species and for detection of adulterants.
Woodson?* has recently reported on the anatomy of about twenty Rauvolfia
species, of which nine are American. At least in one example there is no verifiable
herbarium material available, resulting in a certain amount of doubt. Even as the
cytologists are increasingly depositing specimens of plants which they investigated
in an herbarium, it is strongly recommended that anatomists hemists
T "erer 2:914. 1950. Oxford.
24 Tu E p . Ke €— A e rmacology. Little, Brown & Co.,
Boston. In prin:
[Vol. 43
264 ANNALS OF THE MISSOURI BOTANICAL GARDEN
too realize the advantage of preparing herbarium specimens for resolving future
questions of identity.
In a genus characterized by copious latex, surprisingly, no laticiferous tubes
have been observed in Rauvolfia roots even though artifacts simulating latex tubes
have been noticed in the Asian R. serpentina and R. cambodiana. This is in keep-
ing with the observation of excised roots of living plants.
The roots are tetrarch and in the mature state have metaxylem to the exclusion
of the pith. The outer bark is of varying thickness and consists of phloem and
phellem. The bark may be flaky or indefinitely ribbed. The color of the bark
also varies in relation to the color of the soil in which the roots grow. The sec-
ondary phloem is characterized by the presence of sclerotic cells, either in the form
of prominent bands or of scattered nests of few to many cells, The xylem consists
of vessels, tracheids, and xylem parenchyma. Growth rings are not present in many
of the species. The diameter of the vessels varies considerably, offering a reliable
guide for species determination.
The phloem and xylem characteristics show a trend—possibly phylogenetic—
indicative of a transition from the arboreal to the sub-herbaceous habit. The
phloem exhibits a change from strongly radially arranged sieve cells collapsible on
desiccation in the arboreal species, to weakly radial, non-collapsible sieve cells in
the shrubby species. There is a parallel change in the prominence of the sclerotic
nests of the phloem. An associated change has been noticed in the xylem, the
tracheary wedges consisting roughly of about 50 per cent of the volume in the
arboreal species and 25 per cent or less in the shrubs. The vessels in the arboreal
species are wide, as much as 250 in diameter, while those of the least ligneous
shrubs are as narrow as 20u. To what degree phylogenetic value can be attached
to these trends is an open question. Based as the study was on only about 20
species for the world, it is not surprising that Woodson has “failed to associate
definite anatomical characteristics with Pichon's fourteen sections".
VARIATION AND HYBRIDIZATION
The multiplicity of specific names for essentially similar plant populations of
Rauvolfia has been due to an inadequate awareness of the range of variability.
Many of the species, particularly in the New World, show variation in leaf and
inflorescence characters which has led astray earlier students of the genus. R.
mollis, from Mato Grosso, and R. paraensis and R. pentaphylla, from the Amazon
area, offer examples for such variation to a limited degree. R. macrophylla, R.
caffra of Africa, and R. verticillata of Asia, also have caused much taxonomic
confusion by their variability. e
Two of the American species that have been the cause of vexatious problems of
synonymy are R. tetraphylla and R. ligustrina. Fairly abundant material of these
two species has afforded me an opportunity for a statistical analysis of variation
in some measureable characters. A perusal of the literature provided me with a
list of characters that had attracted the attention of the earlier students. These
1956]
RAO—REVISION OF RAUVOLFIA 265
characters are: leaf number per whorl, leaf shape and size, degree of anisophylly,
degree of clarity of the vein network, leaf indument; peduncle length and degree
of branching; flower number per peduncle; corolla-tube length and fruit size. Of
these, the characters that I have measured are indicated by their mean values in the
following chart (p. 266).
At first I obtained data for small populations restricted to narrow areas. As
these data did not reveal any pattern of variation, as a matter of convenience I
divided the populations into three major areas: Antilles, Central America, and
South America. The variability in leaf shape in R. tetraphylla is erratic and cannot
be correlated with any other character. The ideograph and the accompanying
mean values (fig. 1 of chart) show to what extent the measured characters of these
populations are alike. An evident variable character is the pubescence of the leaf.
The Central American populations have an overwhelmingly large number of
glabrous plants; the Antillean are the most pubescent; while the South American
populations are intermediate with puberulent leaves. This is true in spite of the
disparity in the sample size from the three areas. However, in each of these regions
it is not unusual to encounter plants exhibiting the extreme conditions of pubes-
cence. Because of inconsistency in the variability of these populations, I would
not consider according even subspecific rank to them
The second species analyzed, R. ligustrina, indicates a quantitative variation in
leaf size in the three areas (fig. 2 of chart). Here again a disparity in the sample
size makes the conclusion somewhat uncertain.
In R. tetraphylla, Y was able to note the trend in variation and to compare my
data from herbarium material with data from plants growing in the greenhouse.
I have observed one plant each from Costa Rica, Cuba, and the Dominican Re-
public, and about twenty plants from Lucknow and the Anamalai hills, both
introduced in India?5 from the Antilles.
A study based only on herbarium material cannot be expected to provide an
explanation for this variability in these two species. However, it may be signifi-
cant that these two species are widespread and have been well known to man and
associated with his environment. As to what extent variability is inherent in these
species and to what extent it may be due to extraneous agents can only be resolved
by carefully planned cytogenetic experiments. The large-scale cultivation of R.
tetraphylla plants in some areas, and their spreading as weeds in India, and more
recently in Australia, should also afford ideal opportunity for a detailed field study
of variation in this species.
Hybridization: Herbarium material is not ideal for providing conclusive data
regarding hybridization. Howevgr, often specimens which could not be re
with certainty to one of two sympatric species pointed to the possibility of intro-
gression. Several such specimens were encountered, particularly of R. tetraphylla
Populations. R. fefrapbylla is sympatric, in the range of its distribution, with
three closely related species: R. ligustrina, R. littoralis, and R. viridis. The fre-
25 Voigt, J. O. Hort. Suburb. Calcut. Cat. Pl. p. 202. 1845.
[Vol. 43
266 ANNALS OF THE MISSOURI BOTANICAL GARDEN
Gen Largest Smallest Leaf indument p —_— Fruit
Reri lea ea ——
— ren x b'dth | eth x b'dth | Glabr. Puber. | Pubes. [P| lovers | sone [Breadth
peduncle
: (cm.) (cm.) (cm.) (mm.) | (mm.)
1. Antilles 7 | 59X2.6 | 3.0 X 1.8 PIS bhi 5 64 | 72
2. Cent. America |255 | 6.2 X 2.9 | 3.0 Xx 1.8 |220 | 35 0 | 13 5 GEES
3. So. America 47 6X2.8 | 3.4 X 1.4 39 4 $ 5 64 1 735
Rauvolfia tetrapbylla
ES:
R.TETRAPHYLLA de
Gri
1
Sample Largest Smallest Length | Number| Length of Fruit
: size leaf ea ` = Geng Caceres faite
Region Fe x b’ath | eth x b'dth uncle Eger tube Height Breadth
peduncle
(cm.) (cm.) (cm.) (mm.) (mm.) (mm.)
1. Cub H 423516 E eg E Er? 5 3.0 5.6 6.5
2. Cent. America 15 3.0 X 2.0 | L8X 1.3 1.3 7 3.0 5.6 6.7
3. So. America 9 PORTEL 12 13 6 3.0 SS 6.4
Rauvolfia ligustrina
Chart showing variation in two species of Rauvolfia
1956]
RAO—REVISION OF RAUVOLFIA 267
quency of fruits with abortive embryos in many of these examples indicates a
genetic abnormality and lends further support to the possibility of hybridization.
An instance of what, in my opinion, is very possibly a complex hybrid has
been described as a distinct species, R. mollissima Mfg. The original description
of this species is based on two specimens: Tonduz 13940 from Costa Rica, and
Karsten 17b from Colombia. I have examined both specimens. Of the two, the
Tonduz specimen inclines more toward R. littoralis in leaf shape and size, inflores-
cence branching, and flower number, though to a large extent, particularly in leaf
indument, it resembles the other suspected parent, R. tetraphylla. Similarly, the
Karsten specimen, while resembling R. tetraphylla, still indicates a close approach
in those characters to R. viridis. I have treated R. mollissima as a synonym of R.
tetraphylla. The situation is obviously complicated by a complex, interbreeding
involving several back-crosses. Also, I have strongly suspected hybridization in
R. paraensis and R. sprucei. I am recording these opinions with the hope that they
will stimulate further observations in the field.
Economic Uses
The many names existing for several Rauvolfia species throughout the world
testify to man's familiarity with these plants. I have listed the known common
names separately under each species.
The most significant use of Rauvolfia plants has been in the healing of certain
chronic human ailments. The raw roots of the Asian Rauvolfia _serpentina have
been used for several centuries in treating intestinal disorders and in heart an
nervous conditions. Recent biochemical and clinical investigations have amply
proved the efficacy of the chemical contents of the roots in alleviating high blood
pressure and in calming excitable subjects. The invaluable nature of the chemical
contents naturally has led to an extensive investigation of the substances involved,
not only in the famed Asian species but also in many of the related ones.
Much information on the botany and chemistry of Rauvolfia serpentina is
available and I do not propose to duplicate it here. Monachino?? has a detailed
contribution on the botany of the species, while the New York Academy of Sci-
ences? has published a symposium on the chemistry, pharmacology, and clinical
nature of the active alkaloid, reserpine, the most valuable of about 30 alkaloids so
far isolated from the roots of R. serpentina. Furthermore, Feuell?* has a more
general account of several important Rauvolfia species which is particularly
valuable for its exhaustive bibliography.
With the recent pharmacological vindication of the early claims from folk-lore
of the curative properties of serpentina roots, attention has now been directed to
some related species. Several additional species have been found to contain the
reserpine in exploitable quantities. Almost all commercially extracted reserpine is
26 Monachino, J., in Econ. Bot. pud
"7 Miner, Re W. (editor). Ann. N. Y. Acad. Sel sc 1-140. 1954.
28 Feuell, A. J., in Colon. Pl. & An. Prod. 5:1-33. 1955.
[Vol. 43
268 ANNALS OF THE MISSOURI BOTANICAL GARDEN
now obtained from the African R. vomitoria and the circum-Caribbean R. tetra-
bbylla. A detailed account by Schittler?? of the chemistry of the several species
so far investigated is expected in a forthcoming publication.
In America R. tetraphylla and R. nitida have been much used in indigenous
medicine. The decoction of the bark and leaves has been used as a gargle and in-
ternally administered in intestinal disorders and in syphilitic infections. The bark
has been further used as an unguent on skin eruptions. The fruits are reputedly
poisonous. In Nicaragua the fruits of R. tetraphylla have been used as a source of
ink and a black dye.
Standley?? and Roig y Mesa?! have recounted these numerous uses for the two
species in Central America and in Cuba respectively, while Descourtilz?? has given
a picturesque account of the deleterious properties of R. nitida, which he refers to
as R. canescens. An accurately colored illustration of the plant in the text leaves
no doubt regarding the identity of the species. A confusion on this count has led
Feuell to attribute all properties of R. nitida, described by Descourtilz, to R. canes-
cens, which is identical with R. tetraphylla of Linnaeus.
Apart from the medicinal importance, some of the arboreal species are useful
as sources of timber. Record and Hess,33 however, do not attach much importance
to the value of Rauvolfia as a source of timber. They describe the wood as of fine
texture, easily worked, but perishable in contact with the ground. Monteiro da
Costa mentions the use of R. pentaphylla wood for “furniture and turnery work.”
R. nitida has been introduced into Florida and has been recommended as an efficient
wind-break on the seashore.
R. salicifolia and R. cubana bushes, with their glistening coriaceous leaves and
fragrant flowers, invite serious consideration as ornamental plants in tropical and
subtropical gardens and parks.
GEOGRAPHY
I will confine myself here to a general discussion of the geography of the genus.
Data concerning the ecology and distribution of individual species have been in-
cluded with the descriptions, while the accompanying maps indicate areas which
could be accurately plotted on the outline maps with the help of the 'Lippincott's
Gazetter of the World' and an assortment of large-scale maps published by the
National Geographic Society and other such agencies.
auvolfia is a pantropical genus, with a distribution extending to the 30th
parallel in the north as well as the south. In addition to tropical America, there
are numerous species in Africa, Asia, and in several of the Oceanic islands. How-
29 Schittler, H., in Woodson et al, Rauwolfia: Botany, Chemistry and Pharmacology. Little,
Brown & Co. Boston
. In press.
` Standley, P. C. LM and shrubs of Mexico. In Contr. Smithson. Inst. 23:1153. 1924.
m: Roig y Mesa, J. T. Plant. Medicin. Aromat. Venen. de Cuba. p. 631. 1945. Havana.
Descourtilz, M. E. Fl. Med. Ant. 3:151. 8
33 Record, S. J. & Hess, R. W. Timbers of the World. p. 66. Yale Univ. Press. 1943.
1956]
RAO—REVISION OF RAUVOLFIA
269
nar NR RO
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MAP 1
Distribution of Rauvolfia throughout the world. (All outline maps used in this paper
are from Goode's Series of Base Maps, University of Chicago Press.)
| [Vol. 43
270 ANNALS OF THE MISSOURI BOTANICAL GARDEN
ever, it is not native to Australia where R. tetraphylla, the American species, has
been introduced and has now become naturalized in a small area in Queensland
(Map 1). The distribution pattern of the sections and the species suggests three
or four centers of differentiation. In each of the major land areas there are one or
two widely distributed species. Significantly, these are also widely variable and
have broad tolerances to varied environment. In America, R. tetraphylla and R.
ligustrina are the two “wide” species. In Africa, R. caffra, R. macrophylla and
R. vomitoria are fairly widespread, while in Asia R. densiflora and R. verticillata,
closely followed by R. serpentina, have a wide distribution.
The American R. tetraphylla flourishes equally well in much-exposed, sunny,
sandy soils near the seashore and in wet and marshy, shady grasslands at elevations
of about 2000 m. The Asian R. serpentina is similarly characterized by a wide
ecological range. Such traits should make these two commercially useful species
easy for domestication and large scale cultivation.
The distribution of the sections and the world species can also be obtained
by a reference to Map 1. The large number of species that are endemic is rather
unusual. Willis** has stated that he has no doubt whatever that in the great
majority of cases endemics are simply the early stages of dispersal of species which
as yet have not had the time nor the opportunity to spread far. In Rauvolfa,
however, evidences of comparative morphology, and to some extent of anatomy,
points to the contrary. It is the widespread species that are relatively the most
advanced, while the endemics are relatively primitive from the standpoint of
comparative morphology.
The problem of plant distribution is an ever-present challenge to the botanist.
Particularly in a genus such as Rauvolfia, with no fossil history for the genus nor for
almost the entire family, an attempt to solve the problem is fraught with disillu-
sion and disappointment. It is perhaps for this reason that both Pichon and Wood-
son, though they have included considerable information on the geography of many
genera, have not attempted to integrate the information and present ideas on the
origin and area of the family.
In the past, with an innocent belief in the existence of innumerable convenien
land bridges or in the drifting apart of the originally concentrated land masses, it
was fairly easy to advance hypotheses concerning various problems of plant dis-
persal. Thus Macfarlane, with an unquestioned belief in land bridges, had no
further difficulty in believing that many of the apocynaceous genera had their
origin in West Africa, from where they migrated to America, on the one hand,
and to Asia, on the other. Both Good?* and Schmidt? have recently reviewed
such ideas in plant and animal geography respectively. Schmidt has pointed out
5 Willis, J. C. The birth and spread of plants. In Boissiera 8:509. 1949. :
"4 Macfarlane, John M. The Evolution and Distribution of Flowering Plants. Philadelphia. 1933-
ES Ronald, in “ tury of Progress in the Natural Sciences." Calif. Acad. Sci. p. 747.
37 Schmidt, Karl P. ibid. p.767. 1955.
1956]
RAO—REVISION OF RAUVOLFIA
271
WER A S| EIR WOODSONIA
; © 2.RLITTORALIS
e qued des
D
af
yl
e 4.R.VIRID
x - 4 OSR.MORICANDII
T ^| © 7.R-MOLLIS |
R-LIGUSTRINA
MAP 2
Distribution of eight species of the section RAUVOLFIA
272
ANNALS OF THE MISSOURI BOTANICAL GARDEN
O9.R.MATTFELDIANA
4 10.R.PACHYPHYLLA — |
A 11.R.LEPTOPHYLLA
I2-R.POLYPHYLLA
=
ei3RSPRUCE — |
o I4-R.PARA ENSIS
_Aá%$4+ —À
€ I5-R.-MACRANTHA |
016-R-PENTAPHYLLA
Distribution of eight species of series LATIFOLIA of the section MACcROVOLFIA
[Vol. 43
1956]
RAO—REVISION OF RAUVOLFIA 273
that there is no evidence from geology to support the past existence of land bridges
other than what are present today, and he gives evidence from geodesic studies to
disprove the idea of continental drift.
It cannot therefore be either through land bridges or drifting continents that
the spread of Rauvolfia from the place of its birth can be explained. Plant dis-
persal commonly is effected by animals and birds. Here again there is very little
information available concerning Rauvolfia. Biswas38 has noted that fruits of R.
serpentina are eaten by birds and scattered by them. The brightly colored fruits
of the other species may similarly be attractive to birds. Many species, not only in
America but also in Africa and Asia, have a distribution following rivers and
streams. The stones of Rauvolfia are hard and can withstand considerable buffet-
ing in water currents. It may well be that birds and water currents together are
responsible for the wide dispersal of the genus.
Another tempting problem is that of the center of origin of the genus. Can?
has recently made a critical evaluation of the several critera, first used by Adams,*?
as indicators of the center of origin of a genus or species. In the absence of evi-
dence from several sources such as Paleontology, Cytology and Zoogeography, the
only recourse is to appeal to the present distribution and comparative morphology
of the species. Some supporting evidence may be obtained from the distribution
data of related genera, for example, Aspidosperma, Plumeria, Tabernaemontana,
Thevetia, and Vallesia. Of these, only Tabernaemontana has a pantropical distri-
bution, while the others are almost exclusively New World genera.
I have estimated about 80 species of Rauvolfia for the world, with 34 species
in the western hemisphere and the remaining 46 species distributed fairly equally
in Africa and Asia in the eastern hemisphere. Species concentration and the maxi-
mum number of endemics are evident in tropical America. The eight species of
section RAUVOLFIA are the most advanced while the species constituting section
Macnovorria include an assemblage variously primitive and advanced. Thus spe-
cies concentration, species differentiation, and distribution pattern of related genera
all indicate the possibility of tropical America being the birthplace of the genus.
RELATIONSHIPS
Rauvolfia is generically distinct, with its whorled leaves, terminal cymose
inflorescences, relatively small flowers with eglandular calyx and esquamellate
corolla, and the ovary with one or two ovules developing into one-seeded variously
connate or free drupelets. However, within the genus, the frequent morphological
parallelism in the several species of the major land areas makes a clear-cut classi-
fication of the sections slightly difficult. .A comparison of the flowers of R. ser-
bentina and R. vomitoria (figs. C and D), with those of R. grandiflora (fig. 23)
and R. praecox (fig. 32) respectively will best exemplify this situation.
38 Biswas, K., in Sci. & Cult. 18:579. 1953.
wei? St anle ey x: Foundations of Plant Geography. — 14. New York. 1944.
40 Adams, Charles G., in Biol. Bull. 3:115-131.
[Vol. 43
. 274
ANNALS OF THE MISSOURI BOTANICAL GARDEN
4 4 LA
PUE
O22-R-SANCTORUM |
23-R-GRANDIFLORA" ~~ |"
© 24.R.PAUCIFOLIA |
25.R.SESSILIFOLIA
SENI NEDDELLIANA
aQ 27.R.STEYERMARKIIL Leet Je k x
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O 29.R.BAHIENSIS
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MAP 4
Distribution of eighteen species of series ANGUSTIFOLIAE of section MACROVOLFIA
1956]
RAO—REVISION OF RAUVOLFIA 275
Comparative morphology, combined with the geography of the species, serves
to produce a reasonably satisfactory key reflecting the natural relationships amongst
the species. Some of the characters that provide necessary taxonomic criteria are:
shape and size of the corolla and the extent of pubescence within, the position and
proportion of the anthers in the corolla tube, the dorsal surface of the anthers, and
the extent of carpellary cohesion in the ovary and in the fruit. None of these
characters solely can be used in delimiting a section.
N
A
-
Se
SO)
Gi
St Mac YA
CON Vg Ai
S NE () N aS jl
wa ^ NS > AS
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Fig. C. Rauvolfia serpentina Benth.
[Vol. 43
276 ANNALS OF THE MISSOURI BOTANICAL GARDEN
A survey of the above-mentioned and other features in all the species lead to
the conclusion that advance in the genus has taken a path of simplification by
reduction. Section MACROVOLFIA, with 26 species, exhibits these various char-
acters at different stages of development. Species of series LATIFOLIAE, most of
them confined to the Amazon valley, are prominent by their comparatively broad,
terminally clustered leaves and fairly large flowers. The fruit, however, is very
variable. It is large and fully fused in R. paraensis and R. pentaphylla, while it is
half-fused in the closely related R. sprucei and R. macrantba. In series ANGUSTI-
FOLIAE again, R. grandiflora, with its large flowers and deeply inserted anthers and
hemisyncarpous fruits, is at one end of the scale, while R. sellowii, with its short
A mem. s
a
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bs
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Lr]
x
Fig. D. Rauvolfia vomitoria Afzel.
1956]
RAO—REVISION OF RAUVOLFIA 277
flowers and free drupelets, occupies the other end. All the other species can be
appropriately placed between these two extremes. R. nitida, with its short flowers
and syncarpous fruits and slightly arcuate embryos, represents a further step in the
evolutionary progress of the genus. In all the other species of section Macro-
VOLFIA and in most of the species of the other sections, except CYRTOSIPHONIA,
the embryo is straight. In several species in each of the sections this tendency for
a curved embryo is noticeable. What advantage this imparts to the species is a
matter of speculation.
Section RAUVOLFIA, containing mostly undershrubs with variously pubescent
or glabrous, membranaceous leaves, tiny flowers in comparatively poorly branched
inflorescences, and syncarpous fruits with convolute embryos, represents the most
advanced group.
The African species forming section AFROVOLFIA, the Malaysian species form-
ing section CYRTOSIPHONIA, and the Hawaiian species forming section OcHRosi-
omes, all show morphological features extraordinarily similar to the species of
section MacRovoLriA. Likewise, species of section ENDoLoBus of Africa and
Madagascar show close relationship with AFRovoLFia as well as with Macro-
VOLFIA, except that in species of section ENDOLOBUS the corolla-lobes are inflexed
in the bud. Also the two African species, R. oreogiton and R. volkensii, consti-
tuting section OPHIOXYLANTHUS, with large flowers and the anthers inserted
almost halfway down the tube, are in proximity with species of the Asian section
OPHIOXYLON.
The species of Hawaii and New Caledonia appear to be evolving in their own
fashion. They are less related to the species from the major land masses. Sherff,*!
in a provisional classification of the Hawaiian Rauvolfias, recognized seven species,
mainly based on variation in leaf characters but to some extent on calyx and
corolla size. Obviously, some factor is influencing a reduction in the size of calyx
and corolla. My own experience with the other Rauvolfia species makes me cau-
tious in attaching much taxonomic value to leaf characters in founding new species.
The Hawaiian species show some relationship with the species of section Mac-
ROVOLFIA of America. They appear to be intermediate between species of series
LATIFOLIAE and series ANGUSTIFOLIAE, having generally broad leaves with a strong
tendency to persist only at the terminal nodes, fairly large flowers with character-
istic foliaceous calyx-lobes, and a rather large corolla. Unlike MACROvOLFIA, the
Hawaiian species, constituting the section OCHROSIOIDES, have anthers with smooth
backs. In this they resemble the African species of section AFROVOLFIA. is
The New Caledonian species, constituting the section HEURCKIA, are distinct.
They have strikingly coriaceous leaves 4-6 in a whorl, and the flower with leathery
calyx-lobes and twisted corolla. The stigma-head has no distinct collar, but the
two apiculate lobes are very prominent. The fruit is apocarpous in all the species
and the drupelets are strongly divergent.
*! Sherff, Earl E., in Field Mus. Nat. Hist. Bot. 23:321-331. 1947.
[Vol. 43
278 ANNALS OF THE MISSOURI BOTANICAL GARDEN
My proposed classification of the sections is based on that of Pichon.* As I
have indicated above, no one character can be given undue weight in delimiting
the sections. In my opinion, Pichon's classification suffers from this defect. He
has given much importance individually to the extent of fusion of the fruits, the
size of the leaves, and the size of the disc around the ovary.
As I have shown above, even in very closely related species the fruit may remain
in different degrees of fusion. For example, of the three Cuban species, R. lineari-
folia shows fruits fused only at the base and the lobes divergent, while R. cubana
and R. salicifolia have fruits more than two thirds fused and lobes parallel. Fur-
thermore, all three species have narrow leaves, but they have very close relationships
with the rest of the American species.
Referring to the Asian Rauvolfias, Pichon** himself has stated that his section
OPHIOXYLON differs from DissoLAENA only in having slightly fused fruits. Con-
sequently, I have merged DissoLAENA with OPHIOXYLON. On the same basis, I
have merged RHOPALANTHUs with ENDOLOBUS and DiLOBOCARPUS with CYRTOSI-
PHONIA, and amongst the American sections HEsPEROVOLFIA and MACROVOLFIA.
Another section which disappears is CRYPTOGYNE, characterized by species with
broad discs concealing the ovary. Except for this, the two species of this section,
R. amsoniaefolia and R. chaudocensis, have many characters in common with the
species of section CYrTOSIPHONIA. Hence, I have included CrYPTOGYNE with
CYRTOSIPHONIA. ile I have examined representative specimens for most of the
species, I have not been able to study any specimens for species of section OPHI-
OXYLANTHUS, nor at least three of the species included in section OPHIOXYLON.
I would therefore not claim any finality for this proposed classification of the
sections. A further examination of the material for the species I have mentioned
may very likely lead to a recasting of the system.
‘ACKNOWLEDGEMENTS
I am grateful to the people and the government of the United States of Amer-
ica for the award of a Fulbright Fellowship during 1953-54, which enabled me to
visit this country and to spend a year of graduate study at Cornell University,
Ithaca, New York. I am greatly indebted to Dr. Robert E. Woodson, Jr., of the
Missouri Botanical Garden, for much understanding kindness and generosity and
considerable guidance in the course of this investigation. My thanks are due to
the Ciba Pharmaceutical Products Inc., for the grant of a fellowship during the
tenure of which this work was completed. It is also a pleasure to acknowledge
the various privileges and facilities accorded to me by Dr. Edgar Anderson, the
director, and other authorities of the Missouri Botanical Garden where this researc
has been carried out. My fellow students have rendered me much help by testing
and discussing the taxonomic keys. I should like to express my thanks to the
curators of the various herbaria who have lent specimens for my study, and to
#2 Pichon, M., in Bull. Soc. Bot. Fr. 94:31—39, 1947.
43 Pichon, in Mem. Mus. Nat. d'Hist. Nat. 27:235. 1948.
1956]
RAO— REVISION OF RAUVOLFIA 279
Dr. K. N. Kaul, director of the National Botanic Garden, Lucknow, India, and
Dr. Jorge León of Instituto Interamericano de Ciencias Agrícolas, Turrialba, Costa
Rica, for seeds of R. tetraphylla from those areas. Finally, I appreciate very much
the editorial assistance of Miss Nell Horner in the preparation of the manuscript.
MATERIALS
This study is based on herbarium specimens from the following herbaria. The
abbreviations for them are taken from Lanjouw & Stafleu’s ‘Index Herbariorum’
part I (Regnum Vegetabile, vol. 2, 2nd. ed., 1954).
—Arnold Arboretum, Harvard University, Cambridge, Massachusetts.
EAP —Escuela Agricola Panamericana, Tegucigalpa, Honduras
—Conservatoire et Jardin Botanique, Genéve, Switzerland.
GH —Gray Herbarium of Harvard University, Cambridge, Massachusetts.
K —Herbarium, Royal Botanic Gardens, Kew, England.
lands.
L Bieta) Leiden, Netherlan
M —Botanishe Staatssammlung, München, Germany
MO issouri Botanical Garden, St s, Mi
NY New York Botanical Garden, New York,
—Muséum National d’Histoire Naturelle, bonas. de Phanérogamie, Paris,
France.
RB —Jardim Botanico Rio 9 Goss Rio de Janeiro, Brazil.
S —Naturhistoriska Riksmuseum, Botanical Department, Stockholm, Sweden.
UC —Herbarium of the Uaec: of California, Berkeley, California.
UPS — Institute of Systematic Botany, Botanical Garden and Botanical Museum of the
University of Uppsala, ebe Sweden
US —National Museum, Smithsonian Institution, Washington, D. C.
Ww —Naturhistorisches Museum, Wien, Aust
TAXONOMY
Rauvorri [Plum.] L. Sp. Pl. ed. 1. 208. 1753; Gen. Pl. ed. 5. 98. 1754;
Pichon, in Bull. Soc. Bot. Fr. 94:32. 1947. L. Gen. Pl. ed. 7 (Reichard). 118.
1778 (as RAvworriA). L. Gen. Pl. ed. 8 (Schreber). 160. 1789; L. Sp. Pl.
ed. 4 (Willdenow). 1217. 1798; A. DC. in DC. Prod. 8:336. 1844; Benth.
& Hook. Gen. Pl. 2:637. 1876; K. Sch. in Engl. & Prantl, Nat. Pflanzenfam.
45:153. 1895 (as RAUWOLFIA).
Miser a Sp. Pl. ed. 1. 1043. 1753.
Dissolaena Lour. Fl. ch. 137. 1790.
Cyrtosipbonia Mig. Fl. Ind. Bat. 2:401. 1856.
Heurckia Muell.-Arg. in Flora 55:168. 1870.
Lactescent shrubs or trees with dichotomous or verticillate branches, the nodes
with small interpetiolar deciduous stipules and persistent glands confined to the
leaf-axils or ascending the petiole. Leaves whorled, 3-5 at each node (opposite in
seedlings), petiolate or sessile, simple, entire, penninerved, membranaceous or cori-
aceous, glabrous or pubescent, often anisophyllous. INFLORESCENCES terminal or
lateral and interpetiolar, often geminate, 2-, 3- or 4-chotomously branched, bracte-
ate, few- to many-flowered dichasia. FLOWERs pedicellate or sessile; calyx cam-
[Vol. 43
280 ANNALS OF THE MISSOURI BOTANICAL GARDEN
panulate, 5-lobed, the lobes eglandular within; corolla salverform, infundibuliform,
urceolate or campanulate, usually white, glabrous externally and variously villous
internally, the lobes 5, equal, sinistrorsely contorted, the apices usually just over-
lapping in the bud, rarely inflexed; stamens 5, epipetalous, the anthers free from
each other and the stigma, included, fully fertile, the filament slender, short, the
pollen-grain spherical or subspherical, triculpate; pistil 2-carpellary, superior, the
ovary apocarpous or variously syncarpous and bilocular, with 1—2 ovules on a
ventral or axile placenta, the disc annular or cupuliform, entire or dentate, the
style single, slender, usually glabrous, rarely villous at the base, the clavuncle or
the stigma-head conspicuous, cylindrical, calyptriform or sub-mitriform, obscurely
bi-apiculate. FRUITS apocarpous or variously syncarpous drupes, often only one
carpel developing, 1-seeded, the seeds stout or flat, with a membranaceous testa and
a linear hilum, albuminous, the albumen carnose, entire, easily separable from the
testa, the embryos nearly as long as the seeds, or half as long, straight or slightly
to deeply arcuate, the cotyledons 2, flat, fleshy, ovate, obtuse, the hypocotyl terete,
about as long as the cotyledons.
TYPE SPECIES: Rauvolfia tetraphylla L.
KEY TO THE SECTIONS
A. Leaves with glands in the axil and on the petiole. Corolla-tube urceolate. Fruits fully
syncarpous, globose; embryo U-shaped. Antilles, Central America, and South America. ,
E EE ect. I. RAUVOLFIA
AA. Leaves with glands in the axil only. Corolla-tube salverform, infundibuliform, or
campanulate. Fruits apocarpous or variously syncarpous, globose or bilobed; embryo
curved.
B. Corolla-lobes not inflexed in the bud.
C. Throat of the corolla-tube very narrowly pilose within (a ring of less than 1 mm.),
above the anther-tips.
D. Anthers clearly dorsifixed. Stigma-head with distinct collar. Fruits apocarpous
or variously syncarpous, the drupelets slightly divergen
E. Anthers with a callosity at the back; filaments short and indistinct. Antilles,
ntral America, and South America... Sect. II. MACROVOLFIA
EE. Anthers with a smooth back; filaments slender and distinct.
: ers 6 to 8 times shorter than the corolla-tube.
G. Corolla-tube 7-12 mm. A waii. Sect
GG. Corolla-tube 2-6 mm. long. Tropical Africa and Madagascar.
III. OcHROSIOIDES
FF. Anthers 2 to 4 times shorter than the corolla-tube. Southeast Asia and
i G
apocarpous ew Caledonia..................... Sect. VI. HEURCKIA
CC. Throat of the corolla-tube pilose in a fairly broad band within (from 2 to 6 mm.),
above the anther-tips.
H. Ovary apocarpous or syncarpous only at the base. Fruits apocarpous
or syncarpous only at the base, the drupelets divergent. Southeast
Asia and Malaysia... Sect. VII. OPHIOXYLON
HH. Ovary syncarpous almost to its middle. Fruits syncarpous almost to
tip, the drupelets parallel. East Africa.....Sect. VIII. OPHIOXYLANTHUS
frica and Madagascar.
Sect.
the
BB. Corolla-lobes deeply inflexed in the bud. Tropical
IX. ENDOLOBUS
EES EES Us Up odus,
1956]
RAO—REVISION OF RAUVOLFIA 281
RAUVOLFIA IN AMERICA: KEY TO SECTIONS AND SERIES
A. Leaves oem — — in oe eng si on the petiole, acopio pes ca
low
or pubesce F mall, corolla- mm. long (4-5 mm. in R. andii).
iie fil faeit KS 8 mm. ve Ma 18 mm. in R. littoralis), globose. Em
E. ped I. Racecar
B. en ate hg arely 3 ^ B. the kasoe leaf at each node "on cm. g.
resa ces SE mes shorter than the iated large leaf............. Ser. 1. TETRAPHYLLAE
BB. Leaves 3 at eac h mode, Se 4, the largest let s each qe 3-5 cm. long. Inflores-
cences about as long as the associated l r TERNIFOLIAE
AA. Leaves with mene ve glands Ger | ^ e axil, Pp apa te or coriaceo
glabrous. Flowers large, corolla-tube 4—24 mm. long. Fruits apocarpous or Sege
syncarpous, 0 mm. SC? bilobed (globose in R. paraensis ae R. uer veka Em-
hm straig Sect. II. MaAcROVOLFIA
Flo AM twigs leafy only at the tips; bud-scales or their scars present immediatly
above each node, the current gro deg Piper, of but one verticillate node.
— Dco. as edi as Lt Jin times in R. polypbylla), obtuse to tounde d
a Ser, LATIFOLIAE
es Flowering a le dy duod "ge: bud-scales or their scars usually a — lane
nodes, the current growth consisting of several verticillate nodes, es 34
ons mes as hos as broad (about twice as ah ng in R. steyermarkii), tac ied
ate at both ends or at least at the base. Talora terminal or lateral
r. 4. ANGUSTIFOLIAE
SECTION I. RAUVOLFIA
$ ANISOPHYLLAE Mgf., in Fedde, Rep. Spec. Nov. 20:113. 1924.
$ AMEROVOLFIA Pichon, in Bull. Soc. Bot. Fr. 94:32, 38. 1947.
SERIES 1. TETRAPHYLLAE A. S. Rao, ser. nov
Tee quaternata rarius ternata vel quinata in quoque nodo, folio majore 5-16
mm. longo. Inflorescentiae folio majore 2- vel 4-plo breviores. Species typica:
R. tetraphylla L.
KEY TO THE SPECIES
A. othe parece anisophyllous, the smallest leaf of each node qus si as long as the
al.
arg Inflorescences mostly later eil x-lobes acute, ct or ri
B. a. leaves at the node narrow, 3 to 4 times as long as b rod, Tanceoate, glabrous,
duncles.
distinctly petiolate. Sef ane Aa oo MUS Oy Lean i
BB. Largest leaves at the node broad, about twice as long as broad, ovate, ovate-elliptic, ahr
obovate, Ke or pie indistinctly pone Inflorescences condensed, wit
be = pan Calyx-lobes acute, obtuse, or roun :
iech acuminate, E dedu at the base , glabrous or E
ER nt. Inflorescences -branched, 16- to many-flowered.
hus o oun, Cara Riss a o Colombia, Ecuador, and Venue SH 2. R. littoralis
CC. Leaves ovate or ovate-elliptic, acute or obtuse, abruptly Suen at the
glabrous or pubescent. Inflorescences unbran ched, or rarely 1- nta xir x
an
to Tä cont p acute. Mexico to Colombia, xo eis m en $
AA. Leaves lightly anisophylious: the SS "leaf of each node erg id more than half as
ge, as the largest Inflorescences mostly terminal. Calyx-lobes acuminate or
pidate.
tic, 2-3 times as long as broad, secondary veins arcuate.
eaves ovate ege gg 7 Colombia, gege and Lesser pue! gen A. R. viridis
Leaves narrowly elliptic, 4—5 times as long as broad, secondary veins transverse. P
Corolla-tube 3-5 mm. long. Northeastern Brazil
[Vol. 43
282 ANNALS OF THE MISSOURI BOTANICAL GARDEN
Ee V.C b
z
ut o 4
NN,
IESUS
OC)
Fig. 1. Rauvolfia woodsoniana Standl. (Brenes 6801)
Ld
.
RAUVOLFIA WOODSONIANA Standl. in Field Mus. Nat. Hist.—Bot. 18:942.
1938. (T.: A. M. Brenes 6801!)
Plants with 2-, 3-, or 4-chotomous branches. Twigs slender, faintly angular,
glabrous, the nodes with pectinate glands in the leaf-axils and the base of petioles.
LEAVES strikingly anisophyllous, in whorls of 4 or sometimes 3 at each node, shortly
petiolate, the larger leaves lanceolate, gradually tapering at both ends, 7-12 cm.
long, 2-3 cm. broad, the smaller leaves narrowly ovate-elliptic to lanceolate, 1.5-
4.0 cm. long, 1.0-1.5 cm. broad, membranaceous, glabrous, the secondary veins
delicate, indistinct on the upper surface, but distinct on the under surface, 7-10
pairs, arcuate; petioles distinct, 4-8 mm. long. INFLORESCENCEs lateral, inter-
petiolar, 1, or 2-3 at the nodes, several-flowered; peduncles relatively slender, 2—4
cm. long, 2-, 3., or 4-chotomously branched, the branches divergent, minutely
bracteate; pedicels slender, 2-3 mm. long. FLOWERS small; calyx deeply 5-lobed,
the lobes ovate, rounded, minutely ciliolate, about 1.5 mm. long, glabrous; corolla
broadly urceolate, the tube 2~3 mm. long, about 1.5 mm. in diameter at the base,
glabrous without, but pilose within near the throat and around the stamens, the
lobes rotund, about 1.5 mm. long; stamens 5, inserted near the throat, the anthers
Ovate, acute, about 1 mm. long, the filament distinct, about 0.5 mm. long; ovary
1956]
RAO—REVISION OF RAUVOLFIA 283
bi-carpellary, syncarpous, subspherical, about 1.5 mm. high, about 1.5 mm. in
diameter, glabrous, the ovules 2 in each locule on an axile placenta, the disc annular,
about 1 mm. high, the style slender, about 1.5 mm. long, glabrous, the stigma-head
subcapitate, about 0.5 mm. high, faintly biapiculate. FRUITS not seen.
Flowering in April. La Calera de San Ramon, Costa Rica.
This species is easily separated from its related species by its lanceolate leaves
and the loose inflorescences with slender, elongated peduncles.
Costa RICA. ALAJUELA: San Ramon, Brenes 6801 (EAP, F).
2. RAUVOLFIA LITTORALIS Rusby, Descr. New Sp. S. Am. Pl. 84. 1920. (T.:
H. H. Smith 1905!)
R. multiflora Riley, in Kew Bull. 1927:124. 1927. (T.: L.A.M. Riley 125!)
R. macrocarpa Standl. in Trop. Woods 16:11. 1928, nomen; in Publ. Field Mus.—Bot.
4:254. 1929. (T.: Cooper & Slater 200!)
Shrubs and trees with milky latex, up to 9 m. tall. Branches dichotomous,
terete, glabrous, sparingly lenticellate, the nodes with slender, interpetiolar decid-
uous stipules and glands in the leaf axils and on petioles. Leaves whorled, strik-
ingly anisophyllous, 4 or rarely 5 at the nodes, shortly petiolate, usually obovate,
rarely ovate-elliptic or broadly elliptic, acute to acuminate, gradually tapering at
the base, rarely abruptly so, the largest leaves 6-16 cm. long, 3-6 cm. broad, the
smallest leaves 1-4 cm. long, 0.5-2.0 cm. broad, membranaceous, glabrous or
rarely minutely puberulent, the secondary veins distinct on both surfaces, arcuate,
10-12 pairs; petioles 2-8 mm. long. INFLORESCENCES terminal or lateral, inter-
petiolar, paired at the nodes, several- to many-flowered; peduncles relatively stout,
1.5—4.0 cm. long, 2-, 3- or 4-chotomously branched, minutely bracteate; pedicels
slender, 2-4 mm. long. FLOWERS small; calyx deeply 5-lobed, the lobes ovate,
obtuse or rounded, about 1 mm. long, glabrous; corolla broadly urceolate or almost
campanulate, white, the tube stout, 2-3 mm. long, 1.5-2.0 mm. in diameter at the
base, glabrous without, villous within almost to the base, throat densely pilose, the
lobes orbicular, 1.0—1.5 mm. long, glabrous or rarely ciliate at the margin; stamens
5, inserted near the throat, the anthers ovate, acute, about 1 mm. long, the filament
distinct, about 0.5 mm. long; ovary bicarpellary, syncarpous, subspherical, about
1.5 mm, high, 2.0 mm. in diameter, glabrous, the ovule 1 in each locule on an
axile placenta, the disc annular, about 0.5 mm. high, the style columnar, about 2
mm. long, glabrous, the stigma-head depressed-capitate, about 0.5 mm. high,
faintly biapiculate. rRurrs spherical or subspherical, 8-15 mm. in diameter, green
at first but changing as they ripen from pale red to deep red and black, 2-seeded, the
stones ovate, stout and prominently ridged at the base, flattened and smooth above;
seeds ovate, albuminous, with a membranaceous testa, the albumen copious, carnose,
the embryo strongly arcuate, the cotyledons ovate, obtuse, 2-3 mm. long, the
hypocotyl terete, about equal to the cotyledons in length.
Chiefly near sea-shore from 10 to 100 m. altitude. In thickets and somewhat
[Vol. 43
284 ANNALS OF THE MISSOURI BOTANICAL GARDEN
MM
Fig. 2. R. littoralis Rusby (H. H. Smith 1905)
swampy areas. Flowering and fruiting from June to September. Costa Rica to
Colombia, Ecuador and Venezuela.
Common Names: Colombia—Anguito, Contra Solita, and Cruceto. Panama—Fruta
del Diablo.
This species is obviously closely related to R. tetraphylla L. It can d
distinguished by the usually glabrous, obovate leaves with acuminate apex and
very distinct, strongly arcuate secondary veins, and by the usually geminate, many-
flowered inflorescences.
1956]
RAO——REVISION OF RAUVOLFIA 285
Riley, in describing R. multiflora, mentions that it is distinct from all other
Central American species. He may not have seen either Rusby's description or
representative specimens of R. littoralis. Except that the leaves are slightly smaller,
R. multiflora does not show any other significant difference from the type of
Rusby's R. littoralis. Standley has described R. macrocarpa only on fruiting mate-
rial. I have obtained material from the type locality, through the kindness of
Mr. A. M. Bouché, and have given particular attention to the fruits. R. littoralis
fruits profusely, and occasionally giant fruits have been noticed. It may also be
significant that seeds of R. macrocarpa which I have dissected were found sterile.
I am convinced, therefore, that the type specimen of R. macrocarpa is only an
abnormal specimen of R. littoralis. I am accordingly putting it in synonymy.
MBIA: siehe" Baranoa, Dugand 4549 (US); Barranquilla, vicinity of Las
Pendales, Elias 1543 (F, MO, US). Bottvar: Manati, Dugana 591 (F). MAGDALENA:
Magdalena, Dugand & Beien E tc (F); Santa Marta, Smith 1905 (A, F, G, GH, L, MO,
P, S, US), 1906 (A, F, G, GH, L, NY, P, S, US). VALLEE DEL cauca: Coli Soto Herrera
941 (US).
Costa RICA. PUNTARENAs: San Rafael de Esparta, León 4499 (MO).
Ecuapor. MANABI: El Recreo, Eggers e aem 15568 (F, K, M, MO, S, US).
PANAMA. CANAL ZONE: Cocoli, Riley 125 (MO, US); G. Whit e 120 (MO); Mira-
oe: Gorgas Memorial Lab., P. White 130 (MO , US), west end of island, vicinity of lake,
7 (MO). cuimiQur: Progreso, Cooper & Slater 200 (A, F, US).
3. RAUVOLFIA TETRAPHYLLA L. Sp. Pl 208. 1753. (T.: Linn. Hb., photo
293-4!)
. hirsuta Jacq. Enum. Pl. Carrib. ae 1760; Select. Stirp. Am. Hist. 47. 1763, ex char.
. tomentosa Jacq. l.c. ex char. et ici
. canescens L. Sp. Pl. ed. 2. 303. EE ex char., non R. canescens Descourt. Fl. Ant.
oct sie IA
subpubescens L. Mant. 2:345. 1771, ex char.
bei bl 4:805. 1819, ex cbar.
eteropbylla R. & S. geg Veg. (T.: Bertero s. n.!)
latifolia var. minor Muell.-Arg. l.c. 396 (in ër as to > Jameson 514!)
heterophylla var. puberula A. Gray, in Pr 5:187. 1861, nom. nud.
odontophora Heurck & Muell.-Arg. in Hack. "oh eh 150. 1870. (T.: Spruce
6302
canescens var. intermedia Mgf. in Fedde, Rep. Spec. Nov. 20:115. 1924, in clave.
canescens var. GER léi Le, in clave.
. mollissima Mgf. (T.: Tonduz 13940!)
Le.
. hirsuta var. glabra elt: -Arg.) Woodson, in Ann. Mo. Bot. Gard. 26:299. 1939.
apo PRRUPMM. pan
Sub-shrubs and shrubs with milky latex, 0.5-1.5 m. tall; branches 2-, 3- or
4-chotomous, terete, softly pubescent to glabrous, the nodes with interpetiolar,
linear, deciduous stipules and pectinate glands in the axil and on the petiole.
LEAVES in whorls of 4, rarely 3 or 5, strikingly anisophyllous, shortly petiolate,
very variable in shape, ovate, ovate-elliptic, or oblong-elliptic to obovate-elliptic,
acute to obtuse and broadly acute to obtuse at the base, the largest leaves at the
node 5-15 cm. long, 2-4 cm. broad, the smallest 1-4 cm. long, 0.8—3.0 cm. broad,
[Vol. 43
286 ANNALS OF THE MISSOURI BOTANICAL GARDEN
membranaceous, glabrous to tomentulose on both surfaces, secondary veins distinct
to obscure, arcuate, 5-12 pairs, the vein network distinct or obscure, extremely
close-knit when distinct; petiole 2-5 mm. long. INFLORESCENCES terminal and
lateral, few- to rarely many-flowered; peduncles slender, 1-4 cm. long, rarely 2- to
3-branched, glabrous to puberulent, minutely bracteate, the pedicels slender, 2-5
mm. long, glabrous to puberulent. Flowers small; calyx deeply 5-lobed, the lobes
ovate, acute, about 1 mm. long, glabrous to puberulent without and glabrous
within, corolla urceolate, white, the tube slender, 2-3 mm. long, glabrous to puberu-
lent without and rather densely villous within near the throat, the throat slightly
constricted, the lobes obovate to subrotund, rounded, about 1 mm. long; stamens
Fig. 3. Rauvolfia tetraphylla L. From living plant at the Missouri Botanical Garden.
1956]
RAO—REVISION OF RAUVOLFIA 287
5, included, inserted near the throat, the anthers ovate, acute, about 1 mm. long,
the filament about 0.5 mm. long; ovary 2-carpellary, syncarpous, subspherical,
about 1.5 mm. high and 2 mm. in diameter, the ovules 1—2 in each locule on an
axile placenta, the disc annular, 0.5 mm. broad, the style slender, 1.5—2.0 mm. long,
glabrous, the stigma-head subcapitate, about 1 mm. high, obscurely bi-apiculate.
FRUITS spherical to subspherical, 5-8 mm. in diameter, glabrous, gradvally turning
from green to red and black as they ripen, 2-seeded, the stones ovoid, flattened
ventrally, convex dorsally, distinctly rugose; seeds ovate, slightly curved, albu-
minous, testa membranaceous, the albumen carnose, the embryo deeply arcuate,
the cotyledons cordate-ovate, obtuse, 2.0—2.5 mm. long, the hypocotyl terete, about
as long as the cotyledons.
Plants of extremely varied habitat, near road-sites, in waste places, on hills, on
lake banks, on rocky cliffs, in dry fields and in hedges, in moist rich soil in full sun
or under shade of trees; at altitudes from sea-level to about 2,000 meters. Flower-
ing and fruiting practically throughout the year. I have noticed in the green-
house, young plants scarcely 3—4 months old bearing flowers. Greater Antilles,
Mexico to Colombia, Ecuador, Peru and Venezuela.
Common Names: Colombia—Anguito, Cruceto, Venenito. Costa Rica—Chalchupa,
Cobataco, Guataco. El Salvador, Nicaragua, and Panama—Amatillo, Comida de Culebra,
Guataco, Hierba de San Jose, and Señorita. Mexico—Coralilla, Corazillo, and Corralio.
Venezuela—Boboro.
R. tetraphylla can be distinguished from its related species by its very unequal
leaves with usually obscure secondary nerves, the mostly lateral few-flowered in-
florescences much shorter than the leaves, and the small flowers with acute calyx-
lobes. It has been often confused with R. nitida Jacq., particularly in the Antilles,
but differs from it in having dull leaves with few arcuate secondary veins, the
inflorescences with slender, rarely branched peduncles, and the flowers with acute
calyx-lobes.
The confusing nomenclatorial history of R. tetraphylla and its clarification by
Rendle have already been discussed. The extremely variable leaf characters have
occasioned the publication of several specific names for this polymorphic species.
I have made a statistical analysis of variation in this species. Any study based on
the herbarium material alone or a few plants from the greenhouse cannot be con-
clusive. R. tetraphylla offers a very interesting example for extensive observation
in the field and in the botanical garden. This species has been known to have been
introduced to India in the last 150 years, and to Australia in the last 50 years or
less, and in both countries it has become naturalized. Considering the paucity of
information on the behavior of introduced plants in their new homes, a detailed
botanical study of this species in these regions should be of great interest.
ANTILLES.— 4
BARBADos: St. George, Jordan's estate, Barrow 154 (NY); Goodding 380 (NY).
*5 Voigt, l. c.
[Vol. 43
288 ANNALS OF THE MISSOURI BOTANICAL GARDEN
Cub AMAGUEY: Camaguey to Santayana, Britton 2401 (NY), N. L. 8 E. G.
Britton s Powell A E (MO, US); Cayo eee Shafer 2847, 2853 (NY, US); vicin-
ity e Tiffin, 2886 (NY, US). Havana: Havana, Baker 5130 (NY, US); vicinity of
— r, Britton, Earl & Wilson 6234 (NY), Britton, E "em & Gager 6276 (NY), Palmer
& Riley 840 (US), Ekman 348, 825, 944 13419, 16892 (S), Leon 1337, 5201 (NY),
Shafer 173 (NY), Van Hermann 908 (NY, US), e 1369 (NY); Santiago de las
Vegas, Cook 91 (US). ISLE OF PINES: Kibri Gerona, Curtiss 517 (A, G; GH, L, M,
OI (P); vicinity of Daiquiri, Britton & Cowell 12688 (NY); ët Eggers 5411 (P);
SU iso de Cuba, Havard 68 (NY), Leon 138, A Ria ), Gonzalez d a sn
O); Pico Toduin) south slopes, lg IOIO (US); Yara to Gëtt, Shafer 123 30, :
12366, along Río Canto, 1647 (NY, US), Underwood & ag 130 (NY); precise locality
nown, Ekman 5679, 7063, 0760 (S). PINAR DEL RIO: Bay of Mariel, Britton &
Earle (ee panes Corrientes Bay, Britton & Cowell 9976, 99764 (NY); vicinity of
Gu Britton & Cowell 9749 (NY), Shafer Veer “MO, NY, US); Herra-
dura, = de z E G. Britton, Earle & Gager 632 23 (NY, US), Van Hermann 06 (NY,
US); Los Acostos, Zanna cate 5272 (NY); vicinity of Sumidero, Shafer I pae (NY, P,
US). SANTA CLARA (LAS VILLAs): Cienfuegos, Combs 81 (GH, MO, NY, P, US), Hodge
& Howard 4139 (GH), Jack 4661 (A, US), 5232, 5555 (A, P), 7297 (A); "Jagua, How-
ard, Briggs, et al. 253, 392 A ON v de Së Leon E (NY). PRECISE LOCAL-
OT KN
, MC sf, `W).
M cR REPUBLIC: Late pat Fuertes 238 (GH, L, P, NY, S, US, VeL
A. EA ).
Abe Ekman 8535 (S, US); E ME Ekman 10452 (S, US); En-
. Leonar
Gonaives, Nash & Taylor 1522 (NY); Tortue Island, La Vallée, E
s.n. (A, GH, US); precise locality unknown, Jaeger s. n. (GH, M, P, > UPS, Me
unica: precise locality unknown, Boos s. n. (W); Swartz s. n. (M, S, W).
SAINT THOMAS: precise locality not known, Perrin s. n. (NY).
TRINIDAD: Trinidad Botanic Garden la psa 3 570 P S); road to Galera Point,
(S); St.
O). topaco: Thiebaut 1223 (P e
CENTRAL AMERICA
BRITISH HONDURAS: Beine: Gentle 36 (F); Corozal, Gentle 495 (MO); El Cayo and
vicinity, Chanek 120 (F).
Costa Rica. GUANACASTE: camino de Concepción, Echeverria 4166 (UC); El Coyo-
lar, Standley s. n. (US); Fadl, Echeverria a (UG 3E ES Kupper 1424 (M);
e de Cabello, age 15695 (F); Isla de San Lucas, Quinos (F); La Cruz, M
I (MO, UC, US); Las ee Send & Sie 46663 (US). yo (US) ; Los Loros,
i 21467 (F ). PUNTARENAs: Puntarenas, Beetle 26225 (UC, Ek Brenes 12365 (E),
Howell 10155 (US), León 502 (F), Maxon & Harvey 7857 (US), W. W. & H. E. Rowlee
123, 124 (US); Stork 528 (US), 3397 (F); San José, Tonduz 13916 (P, US); precise
locality not known, Kru koff s. n. (MO); Canton de Osa, vicinity of La gäe, Allen 5287
(P, UC, US), vertiente del Pacifico, P. Birlley fl. 17342 (US).
ATEMALA. ALTA VERAPAZ: La L d. ned 2493 €: 3399 (MO, S),
Co
Ge
N
w
Mercedes Aguilar 138 (A, ne CHIMALTEN. Patzún, Heyde & Lux 6417 (G, F,
MO, US). ESCUINTLA: -Anubis, near Obispo enscher 12371, 12601 (F), Escuintla,
Pacheco s. s. (F), Pedro Pirales 2046 (US), See Ruan S) ; Chichipin, Morales
o 544 (U
Ruano 551 (US); San José, Standley 64227 (F); Las Fianzas, Salas 36 (US). GUATE-
MALA: Ruebl s.n. (MO, US), Merck & Co. s.m. (MO). HUEHUETENANGO: Agua
1956]
RAO—REVISION OF RAUVOLFIA 289
Caliente hoer J. M. & M. T. Greenman 5958 (MO); vicinity of Cuilco,
Steyermark 50757 (F); Nentón, Ce de "e e chumatanes, Steyermark 51431 (F, MO).
e 'xillerman $059 (F). APA: Jutiapa, Standley 75129 (F, MO),
Sen o (F): TALHULEU: Ayutla, bad: 68829 (F); Reüilkaleut Bernoulli & Cario
1835 (S), Standley 66786. san marcos: Ocós, Steyermark 37825 (F), Río Naranjo,
Rojás 175 (US). santa rosa: Chicquimulilla, southeast, Standley 7885 , MO);
7684 (US), Deam 6287 (A, F, MO, US, W); trail between Rio Hondo and waterfall,
neat 29489 (F, MO); Zacapa, anie d 72027 (F), 73654 (F, MO).
COMAYAGUA: mayagua, Williams & pre 14680 (F), Standley :
Chacon I (F); El Bano, J. Sei Rodrigue 2320, 2 200 (F). conrEs: Barbar
Molina 3800, 3867 (F); La Lima, Johansen 37 (F, MO, US), Williams & Molina ipis
(F); Progreso, near San Pedro Sula, KE 69 (F); ps Pedro Sula, dern aye 333 (A);
FRANCISCO MORAZAN: between Suya ves do capte Standley 14200 (F); pr magne
Fogg 21751 (MO); Río oa Gam 1600 (F), Williams 8 Molina 13266 (F, MO,
US); Zamorano, Molina 30 (F), J. Valerio Rodrigues 230 (F), 1241 (F), milo I aen
3896 (F), Williams 14070 an OLANCHO: ity of egene yeh 18186, 18374
(F). VALLE: Amapala, J. Valerio ici Soe (F, MO); vicinity of y iir Stand-
e 20765 (US). Yoro: Aguan River valley, ió et al. 8078, 8655 (F, MO, P, S,
ech SALVADOR. AHUACHAPAN: vicinity of pen Standley 20219 (US).
LIBERTAD: vicinity of pecans pets 23332 (US); near Comasagua, Carlsom 260 (F,
UC), 552 (F). La UNION: Acajutla, Stork, Eyerdam > Beetle agri (A, P, UC), Stand-
ley 21935 (S, US); vicinity of is Union; Beetle 26262 (A, MO, P, UC, US), Standley
20783 (S, US); Laguna de Maquigue, Standley 20981 (US). RAN SALVADOR: Apulo,
Williams oi Molina 16760 (F, US); along the road from San Martin to ri zd Ilopango,
Standley 22558 (US); vicinity of Tonacatepeque, Standley 19529 (US); o de San
Jacinto, near San Salvador, Standley 20601 (US), Velasco 9006 (US), mida Calderón
36 Me SANTA ANA: vicinity of Metapán, Standley & Padilla Y 5033 (F). sa
vicinity of San Vicente, Standley 21309 Gei precum ——
Stendley 21935 (S, et Stork, Eyerdam & Beetle 8732 (A, P, UC), of A
Standley 23440 (US); Nahuizalco, Hartman 20, 63 (S); Sobionste, Sadis EA (US).
PRECISE LOCALITY UNKNOWN: Renson s. n. (
NICARAGUA. CARAZO: vicinity of Toae, Standley 8490 (F). CHINANDEG
Ameyer, Maxon, Harvey & Valentine 7104 (US) Maxon et al. 71 56 (US); Chichigalpa,
Standley 11306, 11396 (F). CHONTALES: Juigalpa, Standley 9376 (F). DA:
Granada, Baker 173 (MO, UC), 608, 846, (US), Levy 214 (P, W), Maxon e e? 7613
)»
ESTELÍ: vicinity of co Standley 20402 (F). MANAGUA: Managua, Chaves 88 (A),
Garnier 328 (MO), J. M. & M. T. Greenman 5687 whee Maxon et al. 7221 (US), d one
7545 (US); Laguna de Mane Maxon et al. 7740 (US); Cosiguina Volcano, owell
I0257 Za A Ometepe Island, Lake Nicaragua, Shimek d Smitb 73 (US), C. L. "idle in s. f.
( seid PRECISE LOCALITY ep Si ne s. n. (P).
ana oucbé s.n. (MO), i 57 X Fo vios ae
oe (A. US): Jaa Diaz, Siti. por (US) ; Panama City, enm
5145 (MO); Panama Vieja, Asplund s. n. (UPS), Panama Vieja to » Bella mu. Als 831
(F, MO, " US); Taboga Island, Standley 27100, 27851 (US), Woods. Allen & Seibert
1530 (A, MO).
Mex
C,
Manzanillo, Eyerdam & Beetle 8711 (A, MO, P, UC), Palmer 1030 (US), West 351 5
(UC); Paso del Rio, Emrick 186 (F). incu Acapulco, ter s.n. (US);
[Vol. 43
290 ANNALS OF THE MISSOURI BOTANICAL GARDEN
pulco, sea cliffs, Clark 7172 (MO); eT C. L. Smitb 6018 (A, MO, P); e
J.N.S . Rose & Painter 9305 (MO, US); Mazatlán, Stork & Horton 8606 (MO, P,
UC); Sen. Dus Hinton 10874 (A, MO, P); p Coyuca, Hinton et al. 5799 (F);
south of Taxco, Clark 7105 (MO). jJaLisco: La Palma, Jones (MO, US). Mexico:
Amatepec, Matuda 31291 (MO); Tamacaltepec, Hinton Leg (F, MO, US), Hinton et al.
3582 (US), Hinton 3826 (P); San Luis de Tu rrubales, S. Jimenez Canossa 2 (F); Maria
Nurs. Island, Octavis Solis 89 (US), Mason 1839 ( US); Mexico, NR 89 (W).
MICHOACAN: Apatzingan, Hinton 12028 (US), Leavenworth 475 (F, MO), Lender
EI Hoogstraal 1352 (F, MO). wnayarīit: Acaponeta, Rose, Eë Sg & Russell 14280
(US); beo of ZE b peg S. ig P SCH US). sAN Luis Potosi: Rascon,
Purpus 5300 US); Tamaso le 5068 (G); Tamazunchale, Kenoyer
s.n. (MO), SE 5038 (UC); Valles, redi q (MO), Vines 3326 (US). SINA-
A: Culiacán, Brandegee s.n. (UC, US); «e del $. n. nd Mosen
Gen
E 3 ns Ortega 7013 (F, P), 7210 (F, MO, P),7 360 (F), 7482 (MO), Rose
14032, s. n. (US) ; Rosario, Rose 1575, 14622 (US) ; San Ce Ortéga 4023, 4126 (US);
Santa Fe, Dani 4678 (US). EH Tlaxcoapan, Hahn s.n. (P); pmi Lyon-
net 2185, 2655 (US). .vERA CRUZ: Antigua, hes 6143 (F, MO, UC, US); Ojapa,
Orcutt 5193 (MO, US); Papantla, ee 15135 (MO, UC, (UN Pueblo Viejo, Palmer
I (F, MO, US); Rio de SE Maria, Pd Se (UC, US); Rinconada, Ross s. n.
(MO); San Francisco, C. L. Smith 1 336 (UC); Vera Cruz, Galleotti 7114 (P); Warten-
berg, near Cartoyuca, Case, 1858 (P). oaxaca: Cuicatlán, Conzatti 3991 (US),
Nelson 1872 (US); Oaxaca, Arson s.n. (L, S), Conzatti 2165 (F), Lä um (G),
4861 (M, MO, P, UC, US, W); San Geronimo, Purpus 7147 (A, F, MO, UC, US);
bere Orcutt 3455 (F, MO, US) ; Tomelin A es Smitb $55 i UC, Ver Tux-
tepec, Martinez-Calderón 90 (MO, UC, US). Yuc AN: Campeche, Fogg 21757 (MO),
C. L. Lundell 900 (F, MO, UC, US); Cabal, Sem 16 (F); Chichankanab, Gaumer
4490 (F), 1869 (A, F, MO, UPS, US); Bx ©: L. e A. A. Lundell 7464 (MO, US),
Steere 1104 (MO), 1 363 (F); Izamal, Geumer. 610 (A, F, MO); Mérida, Schott 431 (F);
"pd locality not known, Lundell 4768 (A, F), Foss 21765 (MO), Gaumer 24047 (F,
SOUTH Ax CA
Britis Guiana. Georgetown Botanical Garden, Gre, & Persaud s. n. (F).
COLOMBIA. BOLIVAR: Cartagena, Billberg 205 (S), Cufodontis 31 (W), Killip 8
Smith 14161 (A, GH, US), Schott 851 (F); Cienaga, D R. Castaneda 984 (MO); Tur-
D U CA:
. CUNDINAMARCA:
Cespedel $.9». (P), T: $.". (P). HUILA Ais » Perez Arbelaez 735 (MO, US).
GDALENA: E qe near Codici zi, Haacht 2316 (A, M O, US), Pittier 1574 (US);
Sierra Nevada de Santa Marta, Barkley 18 C 511 (MO); Sears Marta, Bertero s. n. (P),
Juan Giacometto 1027 (MO), "Purdie e 407 (P), H. H. Smith 1648 (A, F, GH, L, MO, P,
S, US), 1649 (F, GH, MO, P, US), 1654 (P), 2097 (A, F, GH, L, MO, P, S, UC, US).
SANTANDER: Santander, J. A. Molina & Barkley 18 S 256 (MO). PRECISE LOCALITY UN-
N W).
CUADOR. GUAYAs: Salao, Eggers 14535 (M, S, US); Guayaquil, Anderson s. n. (S);
Asplund 5125 (P, UPS, US), Jameson 514 (G), Spruce 6302 (P, S, m" ANABI: Manabi,
Laid 20 MO, US), Solis 10641 (F); Savana ad Zë Mille 989 (F
MAZONAS: Bagua, Dik: s.n. (MO). LAMBAYEQUE: dus Ramon |
Ferreyra 7610 Katie Lopez Miranda 288 (MO, SH Pell, Soukup 4198 (US). LIBER-
tad, A. Raimondi 741 (MO). Santa Lucia, Ule 6805 (L)-
tunm e geg tween Cauchaque and Buenos ge dan mon Ferreyra 10949 (MO).
SAN MARTIN: Huallaga, cerca a Bella Vista, Ramon Ferreyra 10085 (MO).
VENEZUELA. ARAGUA: H. Pittier 14038 US). DISTRITO FE : Caracas and
Puerto Cabello, near El Ca E. Pittier 75 (US); Catia and Las Trincheras, E. Pittier 8
(US); Curucuri Valley near Maiquetia, CN Pittier 13388. (A, F, MO, P, US); Curucuri
1956]
RAO—REVISION OF RAUVOLFIA 291
vicinity, H. Pittier 10282 (GH, US); between Caracas and La Guaira, Rose 21920 (US);
La Guaira, Curran & Haman 854, 886 (GH, US). Lara: Carora, Jahn 179 (US); Rio
Tocuyo, M ema 314 (US). Mrmipa: El Molino, ebe. 56218 (F, MO); Tovar,
Fendler 2206 (GH); MIRANDA: Chaco, Williams 11189 (F). TRUJILLO: La Cerba, Reed
961 (US); La Concepción, Reed 1094 (US); San ig se Mendoza, H. Pittier 1 3329
( US). ZULIA: Isla San Carlos, Curran & Haman $00 US). northern Venezuela,
Gg eid not known, Curran & Haman 784 (GH), 29 (GH, US), Goudot s. n. (P).
MADRAS: Madras, Woodward s.n. (MO). UTTAR PRADESH: Dehra Dun,
Bhatnagar s.n. (MO); Lucknow, Hiralal s. n. (MO), otodan s. 1. (MO).
AUSTRALIA. QUEENSLAND: iot pi near Rockhampton, Rass s.n. (A), Stan-
dish s. n. (A), cult. C. S. I. R. O. 5428 (MO).
4. RAUVOLFIA virIDIS R. & S. Syst. Veg. 4:805. 1819. (T.: Humboldt & Bon-
pland s. n., photo !)
R. psycbotrioides H. B. K. Nov. Gen. et Sp. 3:231. 1819. we aged 69!)
R. nitida Lam. Enc cyc. Tab. 2:304. 1819, non R. nitida Jacq.
R. lamarkii A. DC. in DC. Prodr. 8:337. 1844. (T.: Postal S: 9., SÉ !)
: latifolia A. DC. in DC. Prodr. 8:339. 1844. (T.: Sieber 74!)
R. latifolia var. minor Muell.-Arg. in Linnaea 30:396. 1860. (T.: Sieber 2681)
R. lanceolata Griseb. Fl. Br. W. Ind. 408. 1864, nom. nud., non R. lanceolata A. DC.
1844.
Es 5% ck
FEN
ei 2
MU Sr
| g 527)
ee Z
Fig. 4. Rauvolfia viridis R. & S. (Humboldt 69)
[Vol. 43
292 ANNALS OF THE MISSOUR! BOTANICAL GARDEN
Shrubs up to 2 m. tall; branches terete, puberulent when young, but glabrous
later, the nodes with very few pectinate glands in the axil and on the petiole.
LEAVES slightly anisophyllous, quaternate, or rarely ternate, shortly petiolate, ovate
or broadly ovate-elliptic, acute to acuminate at the tip, tapering at the base, the
largest leaves 5-15 cm. long and 3-5 cm. broad, the smallest leaves 2-4 cm. long
and 1.5-3.0 cm. broad, membranaceous, glabrous above and minutely puberulent
beneath generally or along the midrib and the secondary veins, the secondary veins
particularly distinct beneath, sharply arcuate, 8—15 pairs, the tertiary veins and the
veinlets forming a distinct network; petiole 2-8 mm. long. INFLORESCENCES ter-
minal or rarely lateral, several- to many-flowered; peduncles slender, 1-4 cm. long,
2- or 3-branched, glabrous or puberulent, minutely bracteate, the pedicels slender,
2-4 mm. long. FLOWERs small; calyx deeply 5-lobed, the lobes ovate, acuminate
or cuspidate, 1.0—1.5 mm. long, glabrous; corolla tubular or narrowly urceolate,
white, the tube slender, 2.5-3.0 mm. long, glabrous or rarely scantily puberulent
without and poorly pilose within near the throat, the throat slightly constricted,
the lobes ovate, obtuse, 2.0—2.5 mm. long, about 1 mm. broad, glabrous; stamens
5, included, inserted near the throat, the anthers ovate, acute, about 1 mm. long,
the filament distinct, about 0.5 mm. long; ovary 2-carpellary, syncarpous, sub-
spherical, about 1.5 mm. high and 2.0 mm. in diameter, the disc annular, 0.5 mm.
wide, the style slender, 1.5-2.0 mm. long, the stigma-head depressed-capitate, 0.5
mm. high, obscurely bilobed. rom spherical or subspherical, slightly flattened,
5—7 mm. in diameter, glabrous, 2-seeded, the stones ovate, flat ventrally, convex
dorsally, distinctly rugose; seeds ovate, slightly curved, albuminous, testa sub-
membranaceous, the albumen fleshy, the embryo deeply arcuate, the cotyledons
ovate, obtuse, 2-3 mm. long, the hypocotyl terete, about as long as the cotyledons.
Scattered shrubs from near sea-level to about 1000 meters altitude, on coral
cliffs and along hill-sides on sandy shady beaches, near mangrove swamps but on dry
soil. In arid meadow regions at higher altitudes. Flowering and fruiting from
April to October. Lesser Antilles, Colombia and Venezuela.
Common Names: Colombia—Cruceto, Mata Penos, and Venenito. Lesser Antilles—
Antigua Balsam, Bitterbush, Billyache, Bois Lait and Milky Bush, Snakeberry Tree. Vene-
zuela—Bole de Berraco.
This species can easily be distinguished from its related species by the
slightly anisophyllous leaves with the very distinct vein network and the mostly
terminal inflorescences, with the corolla-tube and the lobes almost equal in the
flowers. It can also be distinguished from R. nitida Jacq. by its usually broad
leaves with deeply arcuate secondary veins, the slender peduncles and the acuminate
calyx-lobes.
McVaugh* has recently shown that Roemer & Schultes’ name, R. viridis, has
the priority over the till.now more familiar Kunth's name, R. psycbotrioides.
Markgrafí? was correct in his recognition of the identity of R. latifolia A. DC
45 McVaugh, R., in Taxon 4:84. 1955.
16 Markgraf, F., in Fedde, Rep. Spec. Nov. 20:114. 1924.
1956]
RAO—REVISION OF RAUVOLFIA 293
with R. lamarkii A. DC., but yet he maintains R. lamarkii as distinct from R.
psychotrioides mainly perhaps on geographical grounds. The characters that Mark-
graf used in his key for separating the two species, such as short- and long-acu-
minate, glabrous or hairy on the midrib, the pilosity in the throat of the corolla,
the shape of the stigma-head, all intergrade and cannot be used. The distribution
of the species from Puerto Rico to Venezuela and Colombia is nothing unusual.
LESSER ANTILLES.—
ANTIGUA: Box 872 (MO, UC, US), Aria 342 (M, W).
605 (US).
GRENADA: Belair, Beard 1201 (GH, S, US), Broedwey s. n. (GH, MO, NY, P); La
Pointe, Eggers 6461 (A, P, US); ap side of Chatham Bay, C. P. Cooper III 202
(NY).
GUADELOUPE: Le Moule, H. Steble 951 (NY), 981 (P), H. 8 M. Steble 6994, 7022
(US). PRECISE LOCALITY UNKNOWN: Bena 533 Ze, zë 6 (P), 2614 (GH, MO,
NY, US), Forstrom 807 (S), Perrotet s. n. jf d xs 211 (PX LG Richard $. n.
um
QUE: Case Pilote, Hahn 1503 Se S), Mouret 226 (P); St. Anne, H. & M.
sii dE (US); Valle des Pierre, Hahn 1459 (G E G, GH, NY, P, US). PRECISE LOCALITY
KNOWN: Duss 1225 (NY, US), Sieber E ei W).
“Mow SERRAT: a 46 (NY, US).
PUERTO Rice: CULEBRA ISLAND: Britton & Wheeler 60a ( US). GUAYAMA:
ig Goll et al. 5 7 (NY). HuMacao: Fajardo lighthouse, ellen 1186 (NY, US),
Sintenis 1194 (GH, M, P, S, US), 1195 (P). MAYAGUEZ: Guanica, Fogg 21775 (MO).
PONCE: Ponce to Playa, Las Cucharas, N. L. 8 E. G. Britton & Marble 1951 (GH, US);
Cayo Muertos, N. L. Britton, Cowell & Stewardson Brown 5048 (N ). SAN JUAN,
San Juan, Sintenis 2806 (L); precise locality unknown, Plée 610 (P), ‘Sergent 195 (US),
Underwood & Griggs 628 (NY, US). Kee: ISLAND: Cerra Encanta, Shafer 2541,
vicinity of Isabela Segunda, Shafer 2560 (NY, US).
St. BARTHELEMY: Forström s. n. (S), Goes s. n. (S, UPS), Questol 228. (NY), 907
m
Crom: A. E. Ricksecker 407 (GH, MO, NY, UC, US), J. J. Ricksecker 333 (P),
E 284 (US), 208 (GH, NY, P), 350 ( (GH).
T. Eustatius: Boldingh 1235B (L), q e
ST. Jan: Bethania, hillside woods, N. L. Brit SN M de I93 (NY, US).
j Y).
S).
Sr. Thomas: Bórgeson 27B6B (S), 66B (NY), N. L. 8 E. G. Britton 6 Shafer 124
(NY), N. L. & E.G. Britton & Marble 401, 1206 (NY, US), Eggers 183 (G, L, M, P, UPS),
Eggers s. n. (GH, W), Ebrenberg s. n. (MO), Jelmarsson s.n. (S), Holton s.n. (NY),
Poiteau s. n. ES
Sr. Vincent: HH & G. W. Smith 557 (GH, NY).
TRINIDAD: Sieber 268 (MO, P, W); roBAco: Broadway s.n. (GH, MO, P, US);
Buccoo Point, Goente 298 (MO).
VIRGIN Let. EEN Gorda, Fisblock 75 (NY).
Bour ron
Tico: Barranquilla, Dugand 125,958 (F), 4822 (MO), Dugand
g eo pon 3217 (US), Elias 14,189 (US), Paul C34 (US); Puerto Colombia,
Elias 1014 (F). BOLIVAR: Cartegena, Dugand & Jaramillo 3374 (US), Heriberto 38, 354
(US), Sa s. n. (F). MAGDALENA: Santa Marta, Espina 47 (US), Perez-Arbelaez 4816,
5860 (US).
VENEUZELA. ARAGUA: Ocumare, H. Pittier 14035 (US), Williams 10170 (F). cara-
BOBO: road from Puerto Cabello to San Felipe, H. Pittier 8872 (GH, US); Maracay, Vogel
[Vol. 43
294 ANNALS OF THE MISSOURI BOTANICAL GARDEN
1305 (M). DISTRITO FEDERAL: Caracas, Curran t$ Haman 1093 (GH, US). MIRANDA:
Guatire, H. Pittier 7644 (GH, US); Santa Lucia, Holt 548 (P); prim locality not
own, H. dali E de '(GH, P, US). monacas: Río Caribe, C wis & Haman
1269 (GH UC). Nueva ESPARTA: Isle Margarita, Johnston 135 (F, G, GH, UC,
US, W), Hon o Ce së 43 (F, GH, MO, P, US). sucre: Cristóbal Colón n, Broadway
103, 109 (GH, US); Cumaná, se n t 69 (P); southwest of Cuchivano, Steyermark
62789 (F, MO). YARACUY: en o San Filipe, Steyermark 55855 (F). ZULIA:
Perija, Tejira 13 (GH, US); d. Fendler 612 (K, GH, MO); precise locality not
nown, Kuntze v74 (F).
5. RAUVOLFIA MORICANDI A, DC. in DC. Prodr. 8:340. 1844. (T.: Blanchet
1007!)
Plants with branches ternate or quaternate, cylindrical or obscurely angular,
glabrous. LEAVES ternate or rarely quaternate, slightly anisophyllous, sparsely
glandular at base, short-petiolate, narrowly elliptic, caudate-acuminate, gradually
attenuate at both ends, entire, 3-8 cm. long, 1.0—2.5 cm. broad, submembrana-
ceous, glabrous, dark green above and pale green beneath, secondary veins delicate,
indistinct above, distinct beneath, transverse, many; petioles 2-8 mm. long. 1N-
FLORESCENCES terminal, 6- to 15-flowered, peduncles slender, 1-3 cm. long, 2- or
3-branched, bracteate, bracts minute, pedicels filiform, 4-6 mm. long. FLOWERS
small; calyx deeply 5-lobed, the lobes deltoid, acuminate, about 1 mm. long;
corolla salverform, the corolla-tube slender, 4-5 mm. long, about 1 mm. in di-
e itm.
TEN
GC
AE
eme
=
=
=
e SS
Za.
Fig. 5. Rauvolfia moricandii A. DC. (Blanchet 1007)
1956]
RAO—REVISION OF RAUVOLFIA 295
ameter at the base, glabrous without, sparsely villous within along the upper half,
the throat constricted, the corolla-lobes ovate, obtuse, 2.0—2.5 mm. long, about 1
mm. broad; stamens 5, inserted near the throat, the anthers ovate, acuminate, the
filaments short, about 0.5 mm. long; pistil bicarpellary, hemisyncarpous, the ovary
subspherical, about 1 mm. high and about 1 mm. in diameter, with 2 ovules in
each locule on an axile placenta, the disc annular, narrow, about 0.5 mm. high,
the style common, columnar, 2.0—2.5 mm. long, glabrous, the stigma-head sub-
cylindrical, about 1 mm. high, obscurely biapiculate. FRUITS not observed.
Northeast Brazil: Bahia.
This must be a comparatively rare species. Amongst all the material borrowed
from the different herbaria, including that of the Rio de Janeiro garden, I have not
come across any more specimens of this species. The paucity of the petiolar glands,
the rather long corolla-tube, and the hemisyncarpous ovary, characteristic of this
species, are exceptional to the section to which it is presently assigned. It is con-
ceivable that with the study of more material this species may well be shifted to
the section MACROVOLFIA.
BRAZIL. BAHIA: precise locality not known, Blanchet 1007 (G).
SERIES 2. TERNIFOLIAE A. S. Rao, ser. nov.
Folia in quoque nodo ternata rariusve quaternata, folio majore 3—5 cm. longo.
Inflorescentiae folium majorem subaequantes. Species typica: R. ligustrina R. & S.
KEY TO THE SPECIES
A. Leaves mucronulate, the largest leaf at each node less than twice as long as broad.
Twigs, leaves and peduncles with a afa pubescence. goin. Sieg Brazil. z
BE R. blancbetii
AA. Leaves acute, the largest leaf at each node mops than twice as long as broad. Twigs,
leaves, and peduncles glabrous or with a whitish pubesce:
B. Leaves (at Mp: t the young ones), young twigs, me? iade with a whitis
cence, rarely Greg Inflorescences spreading; peduncles 2- or 3 grange ge?
5-8 mm. long. Calyx-lobes ovate, acute. Paraguay and south-central Brazi
. mollis
BB. Les ves, young twigs, and peduncles pubes or leaves puberulent along the ai
only beneath. erbe "un gested; peduncles unbranched or rarely 1- to
debat pedicels 2-4 e Calyx- Lew anceolate, acuminate. uba;
Mexico to Colombia, Bolivia, + Gi and i aioe a Brazil 8. R. ligustrina
6. RAUVOLFIA BLANCHETTI A. DC. in DC. Prodr. 8:340. 1844. T.: Blanchet
2718!, 2769!)
Shrubs; branches 2-, 3- or 4-chotomously branching, terete, fuscous-pubescent,
the nodes with very few axillary glands ascending the petiole. LEAVES ternate to
quaternate, slightly anisophyllous, shortly petiolate, ovate to ovate-elliptic, acute,
mucronulate, abruptly attenuate at the base, 1—4 cm. long, 1-3 cm. broad, mem-
branaceous, glabrous to puberulent above and pubescent beneath, the secondary
veins 4-8 pairs, rather indistinct, the vein network hardly distinct; petioles 1-3
mm. long. INFLORESCENCES terminal and lateral, few-to many-flowered; peduncles
slender, dichotomously branching, 1-3 cm. long, puberulent, the pedicels slender,
[Vol. 43
296 ANNALS OF THE MISSOURI BOTANICAL GARDEN
Fig. 6. Posdb blanchetti A. DC. (Blanchet 2718)
2-4 mm. long, puberulent. FLOWERs small; calyx deeply 5-lobed, the lobes ovate,
acute, about 1.5 mm. long, glabrous; corolla tubular or indistinctly urceolate, the
tube slender, 2-3 mm. long, glabrous or rarely puberulent without, densely villous
within near the throat, the throat scarcely constricted, the lobes ovate to sub-
rotund, obtuse, 1.5—2.0 mm. long, the stamens 5, included, inserted near the throat,
the anthers ovate, acute, about 1 mm. long, the filaments distinct, about 0.5 mm.
long; ovary 2-carpellary, syncarpous, spherical, about 1.5 mm. in diameter, the
ovules 1-2 in each locule on an axile placenta, the disc annular, about 0.5 mm.
broad, the style columnar, 1.5-2.0 mm. long, glabrous, the stigma-head subcapitate,
about 0.5 mm. high, obscurely bilobed. rmurrs spherical when young, 2-seeded.
I have not seen ripe fruits. Mueller-Argoviensis!! has depicted mature fruits
in his illustration of R. blanchettii. The fruits are obviously like those of the other
members of this section.
47 Mart. Fl. Bras. 61: tab. 9, fig. 1. 1860.
1956]
RAO—REVISION OF RAUVOLFIA 297
This species can be distinguished from all the related species by its fuscous-
pubescent twigs, the mostly ternate, ovate-elliptic, mucronate-tipped, puberulent
leaves, the inflorescences mostly few-flowered and with dichotomously branching,
pubescent peduncles, and flowers with ovate, acute calyx-lobes.
Plants of northeastern Brazil.
BRAZIL. BAHIA: Monte de la Jacobine and Villa de pate Blanchet 2718 (G, F, P),
n (G); Joazeiro, Martinus 2370 (M). ESPIRITO SANTO: Itapemirim, Glaziou 11183
7. RAUVOLFIA morus S. Moore, in Trans. Linn. Soc. Ser. II. 4:393. 1895. (T.:
S. Moore 950, photo!)
R. divergens Mgf. in Fedde, Rep. Spec. Nov. 20:115, 119. 1924. (T.: Fiebrig 4664!)
Shrubs 1-2 m. tall; branches di- or trichotomous, terete, glabrous, or with a
whitish pubescence, the nodes with a few axillary glands ascending the petiole.
Fig.7. Rauvolfia mollis S. Moore (E. Hassler 7400)
[Vol. 43
298 ANNALS OF THE MISSOURI BOTANICAL GARDEN
LEAVES ternate to very rarely quaternate, slightly anisophyllous, shortly petiolate,
ovate-elliptic to narrowly elliptic, acute, attenuate at the base, 1-5 cm. long,
0.5—3.0 cm. broad, membranaceous to subcoriaceous, glabrous above, generally
puberulent to puberulent only along the midrib beneath, the secondary veins and
the vein network obscure; petiole 1-5 mm. ong. INFLORESCENCES terminal and
lateral, few- to many-flowered; peduncles glabrous to puberulent, slender, 1—4 cm.
long, dichotomously branched, the branches usually divergent, minutely bracteate,
the pedicels slender, 5-8 mm. long, glabrous to puberulent. FLOWERs small; calyx
deeply 5-lobed, the lobes ovate, acute, 1.0-1.5 mm. long, glabrous or ciliate; corolla
urceolate, white; the tube slender, 2.0—3.5 mm. long, glabrous without, villous
within near the throat, the throat a little constricted, the lobes obovate to rotund,
rounded, 1.0-1.5 mm. long; stamens 5, included, inserted near the throat, the
anthers ovate, acute, about 1 mm. long, the filament distinct, about 0.5 mm. long;
ovary 2-carpellary, syncarpous, subglobose, about 1.5 mm. in diameter, the ovules
1-2 in each locule on an axile placenta, the disc annular, about 0.5 mm. broad, the
style columnar, 1.5-2.0 mm. long, glabrous, the stigma-head depressed-capitate,
constricted in the middle, about 0.5 mm. high, obscurely bilobed. Frurrs spherical
to subspherical, 4—6 mm. in diameter, glabrous, 2-seeded, the stones ovoid, flat ven-
trally, bulged dorsally, distinctly rugose; seeds ovate, slightly curved, albuminous,
the testa membranaceous, the albumen carnose, the embryo deeply arcuate, the
cotyledons ovate, obtuse, 2.0—2.5 mm. long, the hypocotyl terete, about as long as
the cotyledons. i
In dry fields and in inundated secondary forests. Flowering from September to
December; fruiting from December through February. Paraguay and central-
southern Brazil.
R. mollis can easily be mistaken for the more widespread R. ligustrina. How-
ever, it can be distinguished by the often whitish pubescence of its twigs, leaves,
and peduncles, by the more branched, conspicuously divergent inflorescences, and
the flowers with ovate calyx-lobes.
The leaf shape and the whitish pubescence of the twigs, leaves and peduncles
are extremely variable. This is particularly evident in a suite of specimens of
Malme from the type locality of R. mollis. The type specimen of R. divergens
Mgf. does not indicate any distinctive character to justify its retention as a distinct
species. Iam, therefore, merging it with R. mollis.
AZIL. MATO GROSSO: Corumba , ;
Cuyaba, Malme 2679, 26794 (S); Santa ook Gg e E EE
(UPS), A 2885 (S). RIO DE JANEIRO: Rio de Janeiro, Dusén 139 (S).
RAGUAY. BOQUERON (CHACO): 21° latitude, Fiebrig 1278 (G, P, S), 1350 (G, P);
between Rio Apa and Aquidaban, Fiebrig 4634 (G, K); Villa Sana, Fiebrig 4664 (G, GH»
: M, S); San Genaro, Isla Margarita, Meyer 18.600 (MO). PRECISE LOCALITY UNKNOWN:
Anisits 2230, 2655 (S), Hassler 7400 (A, G, MO, P, $, UC, W).
1956]
RAO—REVISION OF RAUVOLFIA 299
Fig. 8. Rauvolfia ligustrina R. & S. (Humboldt 1480)
8. RAUVOLFIA LIGUSTRINA R. & S. Syst. Veg. 4:805. 1819. (T.: Humboldt 8
Bonpland s. n.)
R. ternifolia H. B. K. Nov. Gen. et Sp. 3:232. 1819. (T.: Sot 1480!)
R. parvifolia Bert. ex Spreng. Syst. 1:834. 1825. (T.: Bertero s. n.
R. parvifolia var. cubana A. DC. in DC. Prodr. 8:340. 1844, nom. P
> barvifolia var. tomentella Maell. -Arg. in PRE 30:394. og a Sieber 326!)
R. alphonsiana Muell.-Arg. l.c. (T.: Poeppig s. n.!)
R. indecora Woodson, in Ann. Mo. Bot. Gard. 24:12. 1937. (Stork 2800!)
Shrubs 1-3 m. tall; branches dichotomous, terete, glabrous, the nodes with
axillary glands ascending the petiole. Leaves ternate, slightly anisophyllous,
shortly petiolate, ovate to ovate-elliptic, acute to acuminate and broadly acute to
rarely obtuse at the base, the largest leaves at the nodes 3-5 cm. long, 1-5 cm.
broad, the smallest 1—3 crn. long, 0.5—1.5 cm. broad, membranaceous, glabrous or
puberulent along the midrib beneath, secondary veins distinct, arcuate, 4-10 pairs,
[Vol. 43
300 ANNALS OF THE MISSOURI BOTANICAL GARDEN
the vein network obscure; petiole 1-3 mm. long. INFLORESCENCES terminal and
lateral, few- to rarely many-flowered; peduncles slender, 1-3 cm. long, 2- or
rarely 3-branched, the secondary peduncles rather congested, glabrous, or minutely
puberulent, the pedicels slender, 2-4 mm. long, glabrous. FLOWERS small; calyx
deeply 5-lobed, the lobes lanceolate, acuminate, 1.5—2.0 mm. long, glabrous; co-
rolla urceolate, white, the tube slender, 2.0-3.5 mm. long, glabrous without and
pilose within near the throat, the throat slightly constricted, the lobes ovate to `
subrotund, rounded, 1.0-1.5 mm. long; stamens 5, included, inserted near the
throat, the anthers ovate, acute, about 1 mm. long, the filament distinct, 0.5 mm.
long; ovary 2-carpellary, syncarpous, subspherical, about 1.5 mm. high, 2 mm. in
diameter, the ovule 1 in each locule on an axile placenta, the disc annular, 0.5 mm.
broad, the style columnar, 1.5—2.0 mm. long, glabrous, the stigma-head calyptri-
form, about 1 mm. high, obscurely bilobed. FRurts spherical to subspherical, 5-7
mm. in diameter, glabrous, 2-seeded, the stones ovoid, flattened ventrally, convex
dorsally, distinctly rugose; seeds ovate, slightly curved, albuminous, testa mem-
branaceous, the albumen carnose, the embryo deeply arcuate, the cotyledons ovate,
obtuse, 2.0-2.5 mm. long, the hypocotyl terete, about as long as the cotyledons.
Chiefly in moist situations, near sea-shore in coastal thickets, in river plains,
savannas, and wet meadows; from sea-level to 1000 m. altitude. Flowering and
fruiting from April to September. Cuba; Mexico to Colombia, Bolivia, Vene-
zuela, Surinam, and northeastern Brazil.
Common Names: Brazil, Maranhoa—Paratudo; Paraiba—Mamao de pri Colombia
—Contra and Venenito. El Salvador—San Jose. Mexico—Chirillo and Venen
R. ligustrina can easily be recognized by its ternate, mostly ovate-elliptic,
acuminate leaves, the rather congested, mostly few-flowered, lateral inflorescences,
and the flowers with lanceolate, acuminate calyx-lobes.
Roemer & Schultes’ name, R. ligustrina, has priority over the Kunthian name
R. ternifolia, as shown by McVaugh.!8 The several specific names here are mainly
due to an inadequate appreciation of variation in leaf characteristics. Markgraf,
while correctly recognizing the identity of R. alpbonsiana with R. parvifolia, still
maintained the distinction between R. parvifolia and R. ternifolia. He has used as
his key characters leaf size and inflorescence flower-number. Similarly, Woodson
differentiates his R. indecora “by its nearly isophyllous, subsessile leaves, which are
minutely puberulent beneath, and its larger drupes.” These characters, however,
are very variable and do not help in maintaining the distinctions.
ANTILLES.—
CUBA. HAVANA: Havana, Baker 2640 (NY, UC), Leon 7191 (MO, NY), Leon 8
Roca 7251, 7252 (MO, NY), Van Hermann 673 (NY), Wilson CR (A, G, NY, P, UC,
US, W). MATANZAs: Matanzas, Alain wee (MO), Baker 2421 ORIENTE: A
Wier omg 5927 (S). PINAR DEL RIO g 1733 (NY), Shafer La (MO, NY, US).
CLARA (LAS VILLAS): CE Cok 180 (GH, MO, NY, P, US); Sagua, N. L.
18 McVaugh, R., in Taxon 4:84. 1955.
1956]
RAO—REVISION OF RAUVOLFIA 301
Britton & Wilson 306, 369 (NY), Howard 5582 (GH, NY); Santa Clara, Baker 4063
(NY, UC » US), Ekman 16850 (S), Leon 9223, 9447 (MO). PRECISE LOCALITY UN-
D: ru . Britt E
Broadway Mii (G), 2680 (G, US), 9010 (A, MO), 9265 (A, G); St. Augustine, Baker 1
(MO). PRECISE LOCALITY UNKNOWN: Sieber 326 (P, MO), Trinidad Botanical Garden
2772 (US).
CENTRAL AMERICA,—
Costa Rica, GUANACASTE: Bebedero, Brenes 12579 (F); Nocoya, Jorge León 4286,
O).
L SALVADOR. LA PAZ: La a We? IQ (US). La UNION: Laguna de Maquigue,
Standley 20910 (US). sAN MIGUE a de Olomega, Standley 20996 (US), Tucker
949 (US). sowsoNATE: Acajutla, Pre di Calderón 1654 (US).
GUATEMALA. ESCUINTLA: San Jose, Kellerman 4570 (US), Standley 64198 (F,
o» Merck & Co. s.n. (MO). SUCHITEPEQUEZ: south of Tiquisate, Steyermark 47806
F)
XICO. CHIAPAS: Paderon, Matuda 16273 (F, MO), 16927 (F), 2734 (A, oi
Us), 2687 (A, F, MO). oaxaca: Oaxaca, Mitede extra 5 (MO). jaLisco: Tuxpá
Mexia 1042 (A, F, G, MO, P, UC, US).
Ges
Bot mia, BEN o Madre de Dios, Kublmann 581 Vu junction of R. Beni and
R. Madre de Dios, es 2388 (F, GH, MO, W), 2389 (F, GH, P
BRAZIL. n HIA: Bahia, RB 47482 (RB). ceara: Ceara, Allemáo 974 (P), Drouet
2707 (F, GH, S, US), Ducke s. n. (RB), Hoehne 7544 (F). MARANHAO: Ducke s. n.
(RB), Hoebne 3430 (F); near Lorenda, ten $ 2047 (A, F, G, M, MO, S, US). PARA:
Montelegre, ide bnt 3 (P, RB, S, US). Panama: J. C. de M Mord 890 (MO), Xavier 103
MO). PER : Fernando de Noronha, Mosley s. n. (P), Ridley, Lea 6 Ramage 83
(GH); Pernambuco, Gardner 1063 (G, GH, S, US, W). piauí: Netto 40 (F).
CoLom ATLANTICO: Dugand 4541 (US), Dugand & Bariga 2307 (US), Elias
(
1425 (F, MO, P US): Río "Kë Dugand E Jaramillo SC: 3262 (US); Barran-
quilla, Elias oe (E, G, MO, US). BOLIVAR: SE Otto 804 (W ) ; Florencia, Perez
Arbelaez 636 (US); San Martin, Curran 28, 75 (GH, US), 408 (GH), Dugand 528 (F,
O); Río "Tucarsics; foothills of Sierra Nevada, Sage 1025 (MO); San Pedro Alejan-
drino, R. R. Castañeda 74 (F); Monteira, Zainum, Molina t$ Barkley BO. 115 (MO, US),
Pond] 4140 (GH, MO,U
SURINAM: Surinam River, Focke 1287 See
VENEZUELA. ARAGUA: Maracay, Vogel 1455 (M). DISTRITO FEDERAL: Betel, Clar-
endon 67 (US); around Caracas, H. Pittier pes (G, US); Losa, Tamayo 197 (MO).
MERIDA: Tovar, Fendler 2347 (G, GH, MO, P, US).
Section II. MACROVOLFIA (Pichon) A. S. Rao, emend.
$ GRANDIFLORAE Met. in Fedde, Rep. Spec. Nov. 20:116. 1924.
$ HESPEROVOLFIA Pichon, in Bull. Soc. Bot. Fr. 94:32, 33. 1947.
SERIES 3. LATIFOLIAE A. S. Rao, ser. nov.
Rami floriferi apice foliosi, cataphyllis super quoque nodo manifestis, hoc aucto
nodo verticillato unico. Folia ca. bis longiora quam latiora (usque ter longiora in
R. polyphylla) basi obtusa vel rotundata. Inflorescentiae terminales. ci
typica: R. polyphylla Benth.
[Vol. 43
302 ANNALS OF THE MISSOURI BOTANICAL GARDEN
KEY TO THE SPECIES
A. Calyx-lobes ovate. Corolla-tube 6—8 mm. lon
B. Tari ovate or ovate-elliptic. Peduücles NE to or up to 4 times as long as the
e ban membranaceous or subcoriaceous, 1.5—2.5 cm. broad, petioles 1.0—1.5 cm.
Peduncles slender, 3—4 times as long ps petio oles. Northeastern Brazil...9 R. DE
CC. Leaves delicately membranaceous or strik T coriaceous, 3—6 cm. broad, petioles
2.0—3.5 cm. long. Peduncles stout, as hind as he petioles or up to 2.5 times as long
D. Leaves strikingly coriaceous. Peduncles as =% as, or 1.5 times longer than, the
petioles, pedicels 0.8—1.0 cm. long. Corolla-tube lilac-colored, 1.5—2 times a
long as the corolla-lobes. Venezuela 10. R. pachyphylla
DD. Leaves delicately membranaceous. Peduncles 2.0—2.5 times as long as the petioles
pedicels 0.3—0.5 cm. long. Se tube cv and as long as or 1.25 times longer
than the corolla-lobes. Colombia 11. R. leptophylla
BB. Leaves lanceolate. Peduncles shorter than the petioles. Northwestern Brazil.
12
R. polyphylla
AA. — —— or reen — 10-24 m
with 5 rs of distinctly arcuate ahead veins 8—12 mm. apart
n "Petioles 1; ES : A “Tons od 20-to many-flowered. Calyx- lobes
eum 1.0-1.5 m er
G. Leaves mostly SCH adi e, membranaceous. Anthers 2.0-2.5 mm. long;
ovary ida. easy Fruit bilobed, reniform. Weer yes il
and Peru
R. feig
GG. € mostly at node, subcoriaceous. Anthers 1.0-1.5 mm. je
geed "ric globose. Northwestern Brazil 14. R. “paraensis
FF. Em 4 5-1.0 cm. long. ' Inflorescences 1- to 5-flowered. Calyx-lobes
lanceolate, 2.5-3.5 mm. long. Northwestern hol. and Peru.....15. R. furem
EE. Leaves with 12-20 pairs a scarcely arcuate or almost transverse mo sb
mm. apart. Northwestern Brazil R. pentaphylla
9. RAUVOLFIA MATTFELDIANA Mgf. in Fedde, Rep. Spec. Nov. 20:117, 120.
1924. (T.: Ule 7114!)
Shrubs 2-5 m. tall; branches 2-, 3- or 4-chotomous, terete to slightly 4-angled,
sparsely lenticellate, glabrous, the base of young branches with caducous catap ylis
2-3 mm. long, about 1 mm. broad, the nodes with pectinate glands confined to the
axil. LEAVES in verticils of 4 or 5 at the tips of twigs, slightly anisophyllous, short-
petiolate, ovate-elliptic, acute to acuminate, attenuate at the base, the largest leaves
4-7 cm. long, 1.5-2.5 cm. broad, the smallest 2-4 cm. long, 1.0-1.5 cm. broad,
membranaceous to subcoriaceous, glabrous, secondary veins and the vein network
obscure above but distinct beneath; petioles slender, 1.0-1.5 cm. long. INFLORES-
CENCEs terminal, lax, comparatively few-flowered, corymbose; peduncles slender,
2-4 cm. long, usually geminate, 2- to 3-branched, minutely bracteate, the bracts
subulate, about 2 mm. long, the pedicels slender, 10-15 mm. long, glabrous.
FLOWERS rather small; calyx deeply 5-lobed, the lobes ovate, acute, 1.5-2.0 mm.
long, glabrous; corolla tubular, slender, 6-7 mm. long, glabrous without and pilose
within along the upper half, the throat not constricted, the lobes linear-ovate,
obtuse, 3—4 mm. long, 1.5-2.5 mm. broad; stamens 5, included, inserted near
t, the anthers ovate, apiculate, 1.5—2.0 long, with a callosity on the back at
the point of insertion, the filament obscure; ovary 2-carpellary, hemisyncarpous,
obovoid, about 2.5 mm. high and 2 mm. in diameter, the ovules 1—2 in each locule
on a ventral placenta, the disc annular, about 0.5 mm. broad, the style columnar,
3.0-3.5 mm. long, glabrous, the stigma-head cylindrical, a little narrowed in the
1956]
RAO—REVISION OF RAUVOLFIA 303
middle, obscurely bilobed. Frurrs deeply bilobed when young, 2-seeded.
In the fields near hilly areas; flowering in November. Northeastern Brazil.
The terminal verticils of rather small leaves with the distinct vein network
beneath, the few-flowered, slender-peduncled inflorescences, the rather long pedicels
of the fairly small tubular flowers, are characteristics which help in distinguishing
this species from its related species.
BraziL. BAHIA: Bahia, Serra de Sincora, Ule 7114 (G, L). MINAS GERAES: Minas
Geraes, Tapanahoacanga, Pohl 3462 (W).
Fig. 9. Rauvolfa mattfeldiana Mgf. (Ule 7114)
[Vol. 43
304 ANNALS OF THE MISSOURI BOTANICAL GARDEN
Fig. 10. Rauvolfia pachyphylla Met. (Ule 8736 photo, and
e
Pittier 9465).
10. RAUVOLFIA PACHYPHYLLA Mgf. in Fedde, Rep. Spec. Nov. 20:117, 121. 1924.
(T.: Ule 8736, photo!)
Aspidosperma quadriovulatum Pitt. in Bol. Cient. y Tecn. Mus. Com. Venez. 1:66. 1925.
(T.: Pittier 9465!)
Shrubs 1-2 m. tall; branches 2- or 3-chotomous, terete, prominently lenticel-
late, the nodes with many distinct, pectinate glands confined to the axil and with
a verticil of cataphylls or their scars immediately above. LEAVES 4-6 at node
usually at the tips of twigs, long-petiolate, elliptic to obovate-elliptic, acuminate,
abruptly attenuate at the base, 10-15 cm. long, 5-7 cm. broad, strikingly cori-
aceous, glabrous, the secondary veins distinct on both surfaces, 6-8 pairs 10-15
mm. apart, arcuate, faintly joining at the margin to form a marginal vein on either
side; petioles stout, 2-3 cm. long. INFLORESCENCES terminal, many-flowered;
peduncles 2-3, dichotomously branched, stout, 3-4 cm. long, glabrous, minutely
bracteate, the pedicels stout, 8-10 mm. long, glabrous. FLowErs rather small;
calyx 5-lobed, the lobes ovate, acuminate, glandular-dentate at the margin, about
1.5 mm. long, 1 mm. broad, the corolla tubular, lilac to dark violet in color
1956]
RAO—REVISION OF RAUVOLFIA 305
(Ule), about 7 mm. long, 2 mm. in diameter, glabrous without, pilose within
near the throat, the throat a little constricted, the lobes ovate, obtuse, about 4 mm.
long, 3 mm. broad; stamens 5, included, inserted near the throat, the anthers ovate,
acuminate, about 1 mm. long, the filament obscure; ovary 2-carpellary, syncarpous,
with 2 ovules in each locule on an axile placenta, the disc annular, the style colum-
nar, the stigma-head cylindrical, constricted at the two ends, biapiculate.
Between rocks, amongst undershrubs, 900 to 1,600 m. altitude. Flowering
May, June. Venezuela and Mount Roraima in British Guiana.
The type specimen of R. pacbypbylla is presumably lost in the general de-
struction of the Berlin Herbarium. However, the type photo which I had for
comparison includes Markgraf's drawings of the flower dissections. This was
helpful in tracking down the identity of Pittier's Aspidosperma quadriovulatum.
Woodson* had already determined this as a Rauvolfia
VENEZUELA. BOLIVAR: Ptari-Tepui, southeast-facing slopes, Steyermark 60022 (F).
DISTRITO FEDERAL: quebrada de San Lázaro, near Caracas, H. Pittier 9465, 11062 (US).
11. Rauvorria leptophylla A. S. Rao, spec. nov.
Arbor circa 15 m. alta; ramulis sparse lenticellatis griseis plus minusve quad-
rangulatis basi ramulorum novorum caducis ibique cataphyllis praeditis. FOLIA
d T)
SET £
iii
Fig. 11. Rauvolfia leptophylla A. S. Rao (R. R. Castañeda 309)
————
e Woodson, R. E., in Ann. Mo. Bot. Gard. 38:129. 1951.
[Vol. 43
306 ANNALS OF THE MISSOURI BOTANICAL GARDEN
tenuissima in apices ramulorum 5-verticillata glabra late elliptica inaequalia, major-
ibus 9—12 cm. longis 4-5 cm. latis, minoribus 5-6 cm. longis 3.0—3.5 cm. latis,
nervis 9—15 in utroque latere arcuatis; petiolo 2.0-3.5 cm. longo eglandulo. mN-
FLORESCENTIAE terminales corymbosae multiflorae, pedunculis binis 5—6 cm. longis
glabris, pedicellis 3—5 mm. longis gracilibus. FLORES majusculi; calycis lobis 1.5-
2.0 mm. longis circa 1 mm. latis glabris; corollae tubo flavido 6—7 mm. longo extus
glabro intus in dimidio superiore praesertim supra insertionem staminum piloso,
lobis lineari-ovatis obtusis, 4-5 mm. longis 1.0—1.5 mm. latis; staminibus 5 inclusis
sub faucibus insertis, antheris ovatis acutis circa 1 mm. longis subsessilibus; ovariis
bicarpellatis semiconnatis obovoides circa 2.5 mm. altis 2.0 mm. diametro 4-
ovulatis; disco annulare circa 0.5 mm. alto, stylo 3.0—3.5 mm. longo, clavunculo
calyptriformi circa 1 mm. alto, obtuse bilobato. FRUCTUS desunt. l
Trees about 15 m. tall; branches terete or faintly 4-angular, sparsely lenticel-
late, glabrous, the base of young branches with caducous cataphylls, the nodes with
pectinate glands confined to the leaf axils. LEAVES in verticils of 5 at the tips of
twigs, slightly anisophyllous, long-petiolate, elliptic, acuminate, attenuate at the
base, the largest leaves at the node 9—12 cm. long, 4—5 cm. broad, the smallest 5-6
cm. long, 5.0—3.5 cm. broad, delicately membranaceous, glabrous, secondary veins
9-15 pairs, arcuate, equally obscure on both surfaces; petiole stout, 2.0-3.5 cm.
long. INFLORESCENCEs terminal, many-flowered, corymbose; peduncles slender,
5-6 cm. long, usually geminate, 2- or 3-branched, glabrous, minutely bracteate,
the pedicels slender, 3-5 mm. long, glabrous. FLOWERS rather large, calyx cam-
panulate, deeply 5-lobed, the lobes ovate, acute, 1.5-2.0 mm. long, about 1 mm.
broad, glabrous; corolla salverform, yellowish, the tube slender, 6—7 mm. long,
glabrous without, villous within along the upper half, scarcely constricted at the
throat, the lobes linear-ovate, obtuse, 4-5 mm. long, 1.0-1.5 mm. broad; stamens
5, included, inserted near the throat, the anthers ovate, acute, about 1 mm. long,
subsessile; ovary 2-carpellary, hemisyncarpous, obovoid, about 2.5 mm. high, 2
mm. in diameter, the ovules 2 in each locule on an axile placenta, the disc annular,
about 0.5 mm. high, the style columnar, 3.0—3.5 mm. long, glabrous, the stigma-
head calyptriform, about 1 mm. high, obscurely bilobed. Fruits not seen.
ds oe MAGDALENA: San Sebastián de Rábago, Romero R. Castaneda 900, TYPE
12. RAUVOLFIA POLYPHYLLA Benth. in Hook. Jour. Bot. 3:241. 1841. (T:
Robert Schomburgk 891!)
R. eo var. connivens Benth. ex Muell.-Arg. Mart. Fl. Bras. 61:31. 1860. (T:
ce
R. polyphylla var. divergens Benth. ex Muell.-Arg. Le, (T.: Spruce 1837!)
Trees or shrubs with di- or tri-chotomous, terete to slightly quadrangular,
glabrous branches, the nodes with pectinate glands confined to the axil. LEAVES in
verticels of 4—5 at the tips of twigs, slightly anisophyllous, long-petiolate, ovate-
lanceolate, acute to acuminate at the tip, obtuse to abruptly attenuate at the base,
1956]
RAO—REVISION OF RAUVOLFIA 307
the largest leaves 11-15 cm. long, 3-5 cm. broad, the smallest 4-6 cm. long, 1-3
cm. broad, membranaceous, glabrous, secondary veins 5—10 pairs, arcuate, equally
evident on both surfaces; petioles fairly slender, 1-3 cm. long. INFLORESCENCES
terminal, few-flowered, corymbose; peduncles slender, 10-15 mm. long, glabrous,
bracteate, the bracts linear-lanceolate, about 1.5 mm, long, the pedicels slender, 2-5
mm. long, glabrous. FLOWERS white, odorous; calyx deeply 5-lobed, the lobes
Fig. 12. Rauvolfia polyphylla Benth. (R. Spruce 3821, and 1896)
[Vol. 43
308 ANNALS OF THE MISSOURI BOTANICAL GARDEN
ovate, acute to acuminate, 1.5-2.0 mm. long, 1.0-1.5 mm. broad, glabrous; co-
rolla tubular, white, a little dilated near the throat, 6-8 mm. long, glabrous with-
out, villous within along the upper half, rather densely near the anther tips, the
lobes linear-ovate, obtuse, 4-5 mm. ong, 2.0—2.5 mm. broad; stamens 5, included,
inserted near the throat, the anthers ovate, acute to acuminate, 1.0—1.5 mm. long,
with a dorsal callosity at the point of insertion, the filament about 0.5 mm. long;
ovary 2-carpellary, fused at the very base, obovoid, about 2.5 mm. high, 2 mm. in
diameter, the ovule 1 in each locule on a ventral placenta, the disc annular, about
1 mm. broad, the style linear, 3.0—3.5 mm. long, glabrous, the stigma-head cylin-
drical, constricted at the two ends, about 1 mm. high, obscurely bilobed. FRUITS
deeply bilobed, compressed, pear-shaped, 15—19 mm. long, 14-15 mm. broad,
glabrous. e
Chiefly near dry river beds and on the banks of streams; flowering from
September to November and fruiting in December. Northern Brazil and Guiana
rder.
This species can easily be recognized by its ovate-lanceolate leaves and the
peduncles much shorter than the petioles.
The two varieties, obviously based on the characters of the leaves and the fruits,
do not deserve to be maintained, as both the characters are variable and are of no
taxonomic significance.
B : AMAZONAS: Mañaos, Río Taruma, Igapo ad ripas infra cataractum minorum,
Ducke 626 (F, MO, RB, US); Cucuhy, Rio Negro, Igarape Macacumy, Ducke s. n., RB
30117 (RB); secus Río Negro, inter Barcellos et San Gabriel, Spruce 1896 (P, W); inter
Barra et Barcellos, Spruce 1837 (G, GH, MO, W); ad flumen Guainia et Río Negro supra
ostium flumini Cusquiari, Spruce 3621 (P, W); Brasilia australis, without precise locality,
Riedel s. n. (G, P).
Guiana: Robert Schomburgh 891 (W).
13. RAUVOLFIA SPRUCEI Muell.-Arg. in Mart. Fl. Bras. 61:34. 1860. (T.: Spruce
17321)
R. lauretiana Woodson, in Ann. Mo. Bot. Gard. 18:541. 1931. (T.: G. Klug 35!)
. Trees 8-25 m. tall; branches terete, sparsely lenticellate, bark dark brown,
crackled or longitudinally striated, the nodes with glands confined to the axil.
LEAVES in whorls of 4, rarely 3 or 5, at the tips of twigs, slightly anisophyllous,
long-petiolate, ovate to rhombic-ovate, acute to acuminate, abruptly attenuate at
the base, the largest leaves 9—15 cm. long, 5-8 cm. broad, the smallest leaves 5-8
cm. long, 3-5 cm. broad, membranaceous, glabrous, the secondary veins 7-10 pairs,
arcuate, rather obscure above, very distinct beneath; petioles stout, 1.5—3.5 cm.
long. INFLORESCENCEs terminal, many-flowered; peduncles 1—4 cm. long, dichoto-
mously branched, the secondary peduncles often longer than the primary peduncle,
glabrous, bracteate, the bracts ovate, acuminate, about 1.5 mm. long, the pedicels
slender, 5-12 mm. long, glabrous. FLOWERS conspicuous; calyx deeply 5-lobed,
the lobes broadly deltoid, broadly acute to obtuse, 1.0—1.5 mm. long, about 2 mm.
broad, the margin minutely ciliate, glabrous; corolla tubular, whitish with crimson
1956]
RAO—REVISION OF RAUVOLFIA 309
Fig. 13. Rauvolfia sprucei Muell.-Arg. (Spruce 1732, Mexia 6415)
streaks, 12-20 mm. long, about 2 mm. in diameter, glabrous without, sparsely
pilose within near the throat, the throat scarcely dilated, the lobes obovate, obtuse,
5-8 mm. long, 3-5 mm. broad; stamens 5, included, inserted a little below the
throat, the anthers ovate, acuminate, 2-3 mm. long, with a callosity on the back
at the point of insertion, subsessile; ovary 2-carpellary, hemisyncarpous, cylindrical,
about 3 mm. high, 2 mm. in diameter, sulcately marked, the ovules 2 in each locule
on an axile placenta, the style filiform, 5-12 mm. long, glabrous, the stigma-head
calyptriform, about 1.5 mm. high, biapiculate. FRUITS reniform, bilobed when
both carpels develop, but often ovoid, with only one carpel developing, 20-25 mm.
high, 14—16 mm. broad, the stones ovate, compressed, faintly rugose, 19-24 mm.
[Vol. 43
310 ANNALS OF THE MISSOURI BOTANICAL GARDEN
long, 9—15 mm. broad, the seed ovate, albuminous, the testa membranaceous, the
albumen fleshy, the embryo erect, about as long as the seed, the cotyledons ovate,
obtuse, 5-9 mm. long, 3-5 mm. broad, the hypocotyl terete, as long as the coty-
ledons, superior.
Trees on the banks of rivers and overflowed creeks; 100—110 m. altitude. Flow-
ering from September to October; mature fruits in January. Northwest Brazil
and northern Peru.
R. sprucei is closely allied to R. paraensis, and indeed it is often difficult to
distinguish one from the other. However, R. sprucei usually bears leaves in 4's,
the leaves are quite membranaceous, and more often rhombic-ovate. The peduncle
is regularly dichotomous in branching, and very often the secondary peduncles are
longer than the primary peduncle. The anthers are longer, the ovary sulcately
marked. The fruits are reniform, bilobed.
AMAZONAS: Barra, Spruce 1732 ës GH, K, W) Ilha de Bacaba, Fróes 21307
(F, x mouth «€ La uge e GH DS A, G, M, MO, S UC, US);
Ex adi 5107 (NY, S); Mishuyacu, near
Iquitos, Klug 35 "(US), gms Tuer Mexia 6415 (GH, K, MO, S, US); Tarapoto,
Spruce 3856 (K, W).
14. RAUVOLFIA PARAENSIS Ducke, in Arch. Jard. Bot. Rio de Janeiro 4:167.
1925. (T.: Ducke, RB 43!)
R. amazonica Mgf. in Notizblatt 9:960. 1926. (T.: Ducke, RB 917 & RB 11, 395!)
Trees up to about 20 m. tall; branches trichotomous, terete, sparsely lenticel-
late, the nodes with pectinate glands confined to the axil, Leaves in whorls of 5,
rarely 2, 3, 4, or 6, at the tips of twigs, slightly anisophyllous, long-petiolate, ovate
to ovate-elliptic, acute to acuminate, abruptly attenuate at the base, the largest
leaves 9-18 cm. long, 5-9 cm. broad, the smallest leaves 8-12 cm. long, 3-7 cm.
broad, membranaceous to subcoriaceous, glabrous, the secondary veins 6-10 pairs,
arcuate, more distinct beneath; petioles stout, 1.5—4.5 cm. long. INFLORESCENCES
terminal, many-flowered; peduncles geminate, 2- or 3-chotomously branched, 2-18
cm. long, glabrous, bracteate, the bracts ovate, acute, about 1 mm. long, the
pedicels slender, 5-10 mm. long, glabrous. FLOWERS conspicuous; calyx deeply
5-lobed, the lobes broadly deltoid, acute, about 1.5 mm. long, 2 mm. broad, gla-
brous; corolla tubular, white with scarlet streaks (according to Ducke), 12-20
mm. long, about 2 mm. in diameter, glabrous without, sparsely pilose within near
the throat, the throat a little dilated, the lobes ovate, obtuse to rounded, 3-6 mm.
long, 2—4 mm. broad; stamens 5, included, inserted near the throat, the anthers
ovate, acute, 1.0-1.5 mm. long, with a callosity on the back at the point of in-
sertion, subsessile; ovary 2-carpellary, hemisyncarpous, cylindrical, 2.5-3.5 mm.
high, about 2 mm. in diameter, smooth, the ovules 2 in each locule on an a
placenta, the disc annular, about 1 mm. broad, the style filiform, 6-15 mm. long,
glabrous, the stigma-head drum-shaped, about 1 mm. high, bilobed. FRUITS glo-
bose, 3—4 cm. in diameter, 2-seeded, one of them often abortive, the stones fusi-
1956]
RAO—REVISION OF RAUVOLFIA 311
Fig. 14. Rauvolfia paraensis Ducke (Ducke RB 43)
form, compressed, rugose, 3.0—3.5 cm. long, about 1 cm. broad, the seed elliptic,
albuminous, the testa membranaceous, the albumen carnose, the embryo about
half as long as the seed, erect, the cotyledons ovate, obtuse, 4-5 mm. long, 2.0-2.5
mm. broad, the hypocotyl terete, equal in length to the cotyledons.
Trees in non-inundated primary forests; flowering from September to January,
ripe fruits in May. Northern and northwestern Brazil.
Common Name: Brazil, Pará—Gogo de Guariba.
This species is closely related to R. sprucei and R. pentaphylla. It differs from
the former in having mostly elliptic, coriaceous leaves in 5's; the primary peduncles
usually much longer than the secondary peduncles; the flowers with shorter anthers
and smooth ovaries and the fruits globose. It differs from the latter species in
having fewer distinctly arcuate secondary veins and slightly larger fruits.
[Vol. 43
312 ANNALS OF THE MISSOURI BOTANICAL GARDEN
Markgraf has differentiated his new species R. amazonica from R. paraensis
thus:—
2. Blatter 3 zu bis 4, gróber, lingliche-elliptisch (vorn kurz zugespitzt), mit 6 bi
Paar os Kronzipfel Y bis V5 s Een wie die Kronróhre; Fruchtknoten
oppelt so hoch wie die stumpfen Kelchzip R. paraensis
3. Blatter zu 3, Tore 2 vorn E d mit 5 bis 8 Paar Seitennerven; Kron-
zipfel etwa 94 46 lang wie die Kronróhze, Ne doppelt so hoch wie die
spitzen Kelchzipfel R. amazonica
I have examined a good number of Ducke's specimens from the Rio de Janeiro
Garden Herbarium and elsewhere. I have measured and analyzed the leaf and
flower characteristics on which Markgraf has based his R. amazonica. I find a
considerable variation in the number of leaves per node, their shape and structure
clearly varying with age of the twigs. There is a similar variation in the flower
In view of this, I agree with Ducke, who collected and described R. para-
ensis, in including R. amazonica (RB 917, 11395) under that name. Hence, I am
merging R. amazonica with R. paraensis.
BRAZIL. AMAZONAs: Borba (Rio Madeira), Ducke 74 (A, F, MO, US), 30115 Ge
Esperança, ad ostium fluminis Jauary, Ducke 1118 (K, MO, RB, US); Santa Izabel, R
egro, Ducke 23950 (A, RB, S, US); Sao Paulo de Olivença, basin of Rio So lo
Krukoff 8996 (A, F, MO, P, US). PARA: vi y Ducke 785 (F, MO, US), 43 (RB, US);
Boa Vista, Rio Tapajos, Ducke 11391 (RB, S, US); Macajubim, Ducke 11395 (RB);
Santa Izabel, Estrada de Ferro do Braganca, Dscke A (G, P, RB, S, US); Juruty Velho
(Civ. Para), Ducke 2159F (G, with 917, P, RB, S, US); Villa Rave: Rio TE. Pires
(
RINAM. Bro nsweg, Zaandam 6885 (L), Lanjouw 1255 (S, MO); precise locality
not known, RB ani 35, 27412 (RB). :
15. RAUVOLFIA MACRANTHA K. Sch. ex Markgraf, in Fedde, Rep. Spec. Nov. 20:
117,120. 1924. (T.:Ule 5174!)
R. micrantha K. Sch. ex Ule in Engl. Bot. Jahrb. 40:136. 1907. (nom. nud., sphalm).
Shrubs up to 6 m. tall; branches 2-, 3-, or 4-chotomous, terete, glabrous, the
bark longitudinally wrinkled, the nodes with axillary glands. LEAVEs in whorls o
4, slightly anisophyllous, short-petiolate, ovate to broadly ovate-elliptic, acute to
acuminate at the tip, abruptly attenuate at the base, the largest leaves 8-10 cm.
long, 4—7 cm. broad, the smallest 3—6 cm. long, 2-4 cm. broad, membranaceous,
the secondary veins 5—7 pairs, arcuate, distinct on both surfaces; petioles slender
0.5-1.0 (1.5) cm. long. INFLORESCENCES terminal, few-flowered; peduncles
geminate, slender, 1-3 cm. long, bracteate, the bracts lanceolate, about 1.5 mm.
long, the pedicels slender, 5-8 mm. long, glabrous. FLOWERS conspicuous; calyx
deeply 5-lobed, the lobes lanceolate, acuminate, 2.5—3.5 mm. long, glabrous; co-
rolla salverform, white (Ule), 15-20 mm. long, about 2 mm. in diameter, glabrous
without, villous within for more than half of the upper part, a little dilated near
the throat, the lobes ovate, broadly acute to obtuse, 5-6 mm. long, 3-4 mm.
broad; stamens 5, inserted about 3 mm. below the throat, in the region of the
dilatation, the anthers linear-ovate, acuminate, 2-3 mm. long, with a dorsal callosity,
1956]
RAO—REVISION OF RAUVOLFIA 313
Fig. 15. Rauvolfia macrantha (Ule 5174 and Krukoff 6698)
subsessile; ovary 2-carpellary, almost apocarpous, cylindrical, about 2.5 mm. high,
2 mm. in diameter, glabrous, the ovules 2 in each locule on a ventral placenta, the
disc annular, about 1 mm. broad, the style linear, 10-12 mm. long, glabrous, the
stigma-head cylindrical, narrowed at the two ends, about 1.5 mm. high, obscurely
bilobed. FRurrs obcordate, deeply bilobed, often only one carpel developing, then
ovoid, rounded, 2.5—3.5 cm. high, about 2.5 cm. broad, 2-seeded, the stones elliptic,
a little flattened, faintly rugose, 2.0-2.5 cm. long, about 1 cm. broad, the seed
[Vol. 43
314 ANNALS OF THE MISSOURI BOTANICAL GARDEN
elliptic, compressed, albuminous, the testa membranaceous, the albumen fleshy, the
embryo as long as the seed, the cotyledons ovate, obtuse, 5-6 mm. long, 3-4 mm.
broad, the hypocotyl terete, equal in length to the cotyledons.
In high forest, on river banks; flowering August to November; mature fruits
in December. Northwestern Brazil.
R. macrantba is closely related to R. sprucei but can be distinguished from it
and all the other related species by its few-flowered inflorescences with short,
slender peduncles and the flowers with lanceolate, acuminate calyx-lobes, and the
obcordate fruits.
BRAZIL. AMAZONAS: Japura, Ducke 6772 (RB); uu ars erage! near Livra-
mento, on immediate shore of river, Kru of e (A, MO, S, US); Sao Paulo de Olivenca,
near Palmares, Krukoff 82 262, 8422 (A, F, G, MO); bein of creek, o. Krukoff 8871
(A, F, G, MO, US) ; Marary, wé Juruá, Ule 5174 (G, L).
16. RAUVOLFIA PENTAPHYLLA (Hub.) Ducke, in Arch. Jard. Bot. Rio de Janeiro
3:244. 1922. (T.: Ducke 11038!)
Couma pentapbylla Hub. in Bol. Mus. Goeldi 7:124. 1913, nom. nud.
R. duckei Mgf. in Fedde, Rep. Spec. Nov. 20:121. 1924. (T.: Ducke 16544!)
Trees about 25 m. tall; branches verticillate, terete, glabrous, the nodes with
prominent axillary glands. Leaves in 5's, slightly anisophyllous, short-petiolate,
ovate to obovate-elliptic, acute to acuminate, abruptly attenuate at the base, the
largest leaves 10-15 cm. long, 4-6 cm. broad, the smallest 4-7 cm. long, 2-5 cm.
broad, firmly membranaceous to coriaceous, the secondary veins 12-20 pairs, trans-
verse, very little curved at the margins and uniting to form marginal veins, equally
distinct on both surfaces; petioles stout, 1.5—3.0 cm. long. INFLORESCENCES
terminal, many-flowered; peduncles geminate, stout, 3-6 cm. long, bracteate, the
bracts ovate, acute, about 1 mm. long, the pedicels slender, 2-6 mm. long, glabrous.
FLOWERS conspicuous, odorous (Ducke); calyx deeply 5-lobed, the lobes ovate,
acute, about 1.5 mm. long, 2 mm. broad, glabrous; corolla infundibuliform, white
with purplish streaks (Ducke), the tube slender, 12-18 mm. long, about 2 mm. in
diameter, glabrous without, villous within near the throat, the throat a little
ilated, the lobes ovate, obtuse, 6—12 mm. long, 3-5 mm. broad; stamens 5, in-
serted near the throat, the anthers ovate, acute, 1.0-1.5 mm. long, with a dorsal
callosity,, subsessile; ovary 2-carpellary, hemisyncarpous, cylindrical, about 2.5 mm.
, 2 mm. in diameter, glabrous, the ovule 1 in each locule on an axile placenta,
the disc annular, about 1 mm. high, the style columnar, 10-15 mm. long, glabrous,
the stigma-head cylindrical, bulged in the middle, BET bilobed. FRUITS sub-
globose to globose, 2.5-3.0 cm. in diameter, 2-seeded, the stones elliptic, 2.0-2.5
cm. long, about 1 cm. broad, faintly rugose, the seed slightly compressed, albu-
minous, the testa membranaceous, the albumen carnose, the embryo erect, the
cotyledons elliptic, 4-5 mm. long, 2.5-3.0 mm. broad, the hypocotyl terete, equal
in length to the cotyledons.
1956]
RAO—REVISION OF RAUVOLFIA 315
Fig. 16. Rauvolfia pentaphylla Ducke (Ducke 11038)
In non-inundated forests; flowering from June to December; mature fruits in
March, April. Northern Brazil.
Common Names: Brazil, Amazonas—Itapeua Grande, Marfim, and Muira Jussara.
Ducke has pointed out that the last name is also used for Aspidosperma duckei, which is
a tree resembling R. pentaphylla.
This species very much resembles R. paraensis, but can be distinguished by its
leaves with numerous, quite transverse secondary veins, and the smaller fruits. As
in R. paraensis, here also there is considerable variation in the leaf and flower
Characteristics. Hence, there is no reason for the continued distinction of R.
duckei. Ducke has cited several numbers in his original description. Of these ;
have not seen 11032, but 11038 I find well represented in the several herbaria.
[Vol. 43
316 ANNALS OF THE MISSOURI BOTANICAL GARDEN
The Macbride type photograph collection includes a photo of Ducke 11038 from
the Berlin Herbarium. I am, therefore, designating Ducke 11038 (RB) as the
type of R. pentaphylla.
L. AMAZONAs: Mañaos, eg 492 (A, F, MO, US), Ducke RB 22426 (P, RB,
T "Dicho RB 23941 (RB, S, US). amará: Porto Platon, Rio Araguari, Pires & Silva
4784 (MO). PARÁ: up ees Black 47-9 “942 (NY); Cajutuba, M. da Costa 273 (F, MO);
Gurupa, Ducke 11038 = 3298 (G, P, RB, US), Ducke 16544 = RB 13299 (G, P,
RB, US).
SERIES 4. ANGUSTIFOLIAE A. S. Rao, ser. nov.
Rami floriferi prorsus foliosi, cataphyllis non manifestis, hoc aucto nodis verticil-
latis pluribus. Folia 3- vel 4-plo longiora quam latiora (ca. 2-plo longiora in R.
steyermarkii) basi attenuata. Inflorescentiae aut terminales aut laterales. Species
typica: R. grandiflora Mart.
KEY TO THE SUBSERIES
A. e linear or obovate, 0.2—2.5 cm. broad. Corolla-tube 8-20 mm. Eis constricted
End due cor Se debs dana. obtuse. Cuba series 1. CUBANAE
AA. L ellip broad. Corolla-tube long, constricted a t the
cete ep ve PW acute. Andes of Pers Eo x e Argent
Subedi 2. ANDINAE
AAA. Leaves obovate or ege: 3—7 cm. broad Kees less than 3 cm. Pe d in R.
weddelliana ez R. paucifolia, but not obovate). rolla-tube 4—20 mm. long, not
constricted at ër A corolla bei Fais acute or obtuse. Greater Audits and
northern Mi ons c South Americ Subseries 3. pmi
SUBSERIES 1. CUBANAE A. S. Rao, subser. nov.
Folia linearia vel obovata 0.2-2.5 cm. lata. Corollae tubus 8-20 mm. longus,
faucibus constrictis; lobis obovatis obtusis. Species typica: R. cubana A. DC.
KEY TO THE SPECIES
A. Hohen subshrubs 1—5 dm. tall. Leaves linear. Corolla purplish-violet; calyx-lobes
naceous, acuminate. Fruits deeply 2-lobed, the lobes acute or slightly beaked.
Geer R. linearifolia
AA. Woody shrubs 2-8 m. tall. Lezves obovate Corolla white; calyx-lobes ën
obtuse. Fruits bilobed at the top, the lobes obtus or broadly acute, but not beaked.
B. Leaves mostly 4-nate. Cor etal 8-10 mm. long, Bier in the throat, the lobes
alf as long et 18. R.
hal long as the tu salicifolia
BB. Leaves mostly 3-nate. QE Ae 12-20 mm. long, villous in the throat, ui lobes
almost as long as the tube, Western provinces of Cuba and the Isle of Pines...19. R. cubana
17. RAUVOLFIA LINEARIFOLIA Brit. & Wils., in Mem. Torr. Bot. Club 16:94.
1920. (T.:Sbafer 1754!)
Shrubs 1—5 dm. tall; branches slender, terete, 2-chotomously branching, gla-
brous, the nodes with minute glands confined to the leaf-axils. LEAVES in whorls
of 3, scarcely anisophyllous, shortly petiolate, linear, acute to acuminate, attenuate,
at base, 2-6 cm. long, 0.2—0.5 cm. broad, membranaceous, the midrib distinct on
both surfaces but the secondary veins obscure; petioles slender, 1-3 mm. long.
1956] :
RAO—REVISION OF RAUVOLFIA 317
Fig. 17. Rauvolfia linearifolia Brit. & Wils.
(Ekman 9552).
INFLORESCENCES terminal, few-flowered, cincinnate; peduncles slender, 1-5 cm.
long, glabrous, bracteate, the bracts lanceolate, about 1.5 mm. long, the pedicels
slender, 2-5 mm. long, glabrous. FLOWERS conspicuous; calyx deeply 5-lobed, the
lobes lanceolate, 1.5-2.0 mm. long, glabrous; corolla salverform, pale purplish-
violet (Ekman), the tube slender, 8-10 mm. long, about 1.5 mm. in diameter at
the base, glabrous without, villous within along the top half, constricted at the
throat, the lobes ovate-elliptic, obtuse to rounded, 4-5 mm. long, 1.5-2.5 mm.
broad; stamens 5, inserted near the throat, the anthers linear-ovate, about 1.5 mm.
long, the filament about 0.5 mm. long; ovary 2-carpellary, connate at the base,
obovoid, about 1.5 mm. high and 1.5 mm. in diameter, the ovule 1 in each locule
on a ventral placenta, the disc annular, about 1 mm. high, the style linear, 5-6 mm.
long, glabrous, the stigma-head drum-shaped, about 1 mm. high, obtusely bilobed.
FRUITS deeply bilobed, the lobes widely divergent, flattened, semi-lunate, acuminate
or slightly beaked, 9-12 mm. long, 2.5-3.0 mm. broad, the stones elliptic, smooth,
the seed elliptic, flattened, albuminous, the testa membranaceous, the embryo erect,
smali (probably abortive), the cotyledons elliptic, rounded, about 1 mm. long, 0.5
mm. broad, the hypocotyl terete, equal in length to the cotyledons.
On limestone hills and in the savannas; flowering from April to September;
ripe fruits in October and November. Oriente Province in Cuba.
[Vol. 43
318 ANNALS OF THE MISSOURI BOTANICAL GARDEN
Rauvolfia linearifolia is the shortest of all the American Rauvolfias. It is
characterized by its narrow, linear leaves, the few-flowered, slender-peduncled in-
florescences, the flowers with lanceolate calyx-lobes, and the compressed, widely
divergent, slightly beaked fruits. In all the fruits analyzed the embryos were very
small and indicated abortive development. The plants appear to propagate more
often vegetatively from the root-stocks. The plants of this species are endemic,
and apparently rare in Cuba.
UBA. ORIENTE: Sierra de Nipe, Ekman 1733, 5013, 5939, 0552, 9085, 15304 (S);
Seer 1754 (NY).
18. RAUVOLFIA SALICIFOLIA Griseb. in Mem. Am. Acad. N. S. 8:519. 1863. (T.:
Wright 13861)
Shrubs or trees up to 8 m. tall; branches 2-, 3- or 4-chotomous, terete, glabrous,
the nodes with distinct pectinate glands in the leaf-axils. Leaves in whorls of 4,
scarcely anisophyllous, shortly petiolate, narrowly obovate to oblanceolate, broadly
acute to obtuse, cuneate at the base, 2-7 cm. long, 1-2 cm. broad, coriaceous,
slightly lustrous above, opaque beneath, the midrib distinct on both surfaces but
the secondary veins obscure; petioles slender, 3-8 mm. long. INFLORESCENCES
LEUR i02 10,3 RE eme TER im
Fig. 18. Rauvolfia salicifolia Griseb. (Wright 1386)
1956]
RAO—REVISION OF RAUVOLFIA 319
terminal, few-flowered; peduncles rather stout, 2-6 cm. long, glabrous, bracteate,
the bracts ovate, acute, about 1 mm. long, the pedicels rather stout, 2-4 mm.
long, glabrous. FLOWERS conspicuous; calyx campanulate, deeply 5-lobed, the
lobes occasionally unequal, ovate, obtuse, about 1.5 mm. long, 1 mm. broad, coria-
ceous, the margin minutely glandular-dentate; corolla salverform, white (Wright),
the tube slender, 8-10 mm. long, about 1.5 mm. in diameter at the base, glabrous
without and within near the throat and sparsely villous below the stamens, con-
stricted at the throat, the lobes oblique-obovate, rounded, 5-6 mm. long, 3.0-3.5
mm. broad; stamens 5, inserted near the throat, the anthers ovate, acute, about 1
mm. long, subsessile; ovary 2-carpellary, hemisyncarpous, obovoid, about 2.5 mm.
high, 2 mm. in diameter, the ovules 2 in each locule on an axile placenta, the disc
annular, about 1 mm. high, the style linear, 5-6 mm. long, glabrous, the stigma-
head subcapitate, about 1 mm. high, obscurely bilobed. Frurrs obcordate, slightly
bilobed, the lobes obliquely obovate, obtuse, broadly acute to obtuse, 13-14 mm.
long, 6-8 mm. broad, the stones slightly flattened, faintly rugose, the seed albu-
minous, the testa membranaceous, the albumen fleshy, the embryo about half as
long as the seed, erect, the cotyledons ovate-elliptic, 3.0-3.5 mm. long, 1.5-2.0
mm. broad, the hypocotyl terete, equal in length to the cotyledons.
On limestone hills, in open forests and amongst Pines; flowering from June to
November; ripe fruits in January and February. Oriente province in Cuba.
Common Name: Cuba, Oriente—Corazon de Paloma.
Rauvolfia salicifolia is closely related to R. linearifolia and R. cubana. It can
be distinguished from the former by its broader, obtuse, coriaceous leaves, the
rather stout peduncles with white flowers, the calyx-lobes stiff and obtuse, and
the fruits which are obscurely bilobed and obtuse. It can be distinguished from
the latter by its narrower and longer leaves, and smaller flowers with the corolla-
lobes about half as long as the tube. Like R. linearifolia, this species is endemic to
Oriente province, Cuba.
CUBA. ORIENTE: Alain 3310, 3319, 3673 (MO), Carebia 3823 (NY), Clement 3566,
3605, 4390 (MO), Ekman 3322, 3514, 4147, 6145, 6688, 9078 (S), 6329 (P, S), Howard
5942 (GH, NY, US), 6104 (GH, MO, NY, UC), Corta 2.038 (W), Leon 8 dn
23046 (MO), Leon, Clement 3 Alain 22523 (MO), Leon, Victorin $ Clement gh d
(MO), Roig 65 (MO), Shafer 3529, 3687 (GH, NY), 4238 (GH), 8144 (NY), 9304
(NY, US), Wright 1386 (G, GH, MO, P, S, W), 2946 (G, GH, MO, P, S).
19. RaAuvoLriA CUBANA A. DC. in DC. Prodr. 8:339. 1844. (T.: Sagra 535,
photo!)
Shrubs or small trees, 2-5 m. tall; branches 2- or 3-chotomous, terete, glabrous,
the nodes with distinct pectinate glands limited to the leaf-axils. LEAVEs in whorls
of 3, scarcely anisophyllous, shortly petiolate, obovate, obtuse, cuneate at the base,
1-6 cm. long, 1-2 cm. broad, coriaceous, lustrous above, opaque beneath, the mid-
rib distinct on both surfaces but the secondary veins obscure; petioles 2-4 mm.
[Vol. 43
320 ANNALS OF THE MISSOURI BOTANICAL GARDEN
long. INFLORESCENCES terminal, few-flowered; peduncles slender, 2-5 cm. long,
glabrous, bracteate, the bracts ovate, acute, about 1 mm. long, the pedicels slender,
6-11 mm. long, glabrous, FLoweERs rather large; calyx deeply 5-lobed, the lobes
ovate, obtuse, 2-3 mm. long, 1.0—1.5 mm. broad, coriaceous, the margins glandular-
dentate; corolla salverform, white, odorous (R. de la Sagra), the tube slender, 12-
20 mm. long, about 2 mm. in diameter, glabrous without, villous within near the
throat and along the base of the stamens, constricted at the throat, the lobes obovate-
elliptic, rounded, 10-14 mm. long, 5-7 mm. broad; stamens 5, inserted near the
throat, the anthers ovate, acute, 1.0-1.5 mm. long, the filament about 1 mm. long;
ovary 2-carpellary, hemisyncarpous, cylindrical, about 2.5 mm. high, 2 mm. in
diameter, the ovules 2 in each locule on an axile placenta, the disc annular, about
1 mm. high, the style filiform, 10-14 mm. long, glabrous, the stigma-head cylin-
drical, about 1 mm. high, 2-apiculate. Frurrs broadly obcordate, bilobed, the
lobes obovate, flattened, broadly acute to obtuse, 10-12 mm. high, 5-7 mm. broad
(often only one carpel developing, then the fruit ellipsoid), the stones faintly
rugose, the seed albuminous, the testa membranaceous, the albumen fleshy, the
Fig. 19. Rauvolfia cubana A. DC. (Wright 2047)
1956]
RAO—REVISION OF RAUVOLFIA 321
embryo erect, a little more than half as long as the seed, the cotyledons elliptic,
about 3 mm. long, 2 mm. broad, the hypocotyl terete, as long as the cotyledons.
On limestone hills, in savannas and in swampy regions along river banks;
flowering from April to August; ripe fruits in October to December, western Cuba,
and the Isle of Pines.
Rauvolfia cubana resembles R. salicifolia very much but can be distinguished
from it by its larger flowers with the calyx-lobes larger, the corolla-lobes about as
long as the tube and the throat villous, and the broadly obcordate, smaller fruits.
'R. cubana also is an endemic of Cuba, but confined to the western provinces and
the Isle of Pines.
CUBA. Havana: N. L. Britton, Cowell & C. de LaTerre 13345 (NY), Ekman 10022,
12583 (S), Leon 5208 (NY), Leon, Ekman et al, 9096 (NY), Leon & Roca 7712 (NY),
L. C. Richard 13 (P), Roig 3 (NY), Shafer 72 (NY). PRECISE LOCALITY UNKNOWN:
Sagra ^ 815 (P), Wright 2947 (G, GH, MO, NY, P, S, US, W). ISLE OF PINES:
L. Britton t$ Wilson 14876 (GH, NY, US), Ekman 11642 (S), fenes 212, 615 (GH,
NY, US), Killip 44082 (US). PINAR DEL Alo; N. L. Britton eC ll 9987 (NY), N.
E. G. Britton & Gager 7014 (NY, US), N. L. Britton $ Gager sa lh (US), Leon &
Charles 4914 (NY). SANTA CLARA (Las VILLAs): N. L. Britton, Earle & Wilson 4599
NY), N. L. Britton & Wilson 5744 (US), Combs 245 (G, GH, MO, NY, P), Ekman
17069 (S), Ties 7540 (A, NY, S, US), 8684 (A, NY, S), Leon 9246 (NY), Van Her-
(L).
mann s.
SUBSERIES 2. ANDINAE (Mgf.) A. S. Rao, stat. nov.
$ ANDINAE Mgf. in Fedde, Rep. Spec. Nov. 20:118. 1924.
KEY TO THE SPECIES
A. Leaves 3 times as long as broad. Corolla-tube 5-6 mm. long, the throat villous; calyx-
lobes cuspidate, the margin entire. Andes of Bolivia and northern vers ina.....20. R. schueli
AA. Leaves 4 times as long as broad. Corolla-tube 8—12 mm. ee the oat et
al Per $ à
yx-lobes lanceolate, the margin Geet -dentate. Andes o andina
20. RAUVOLFIA SCHUELI Speg. in Physis 3:337. 1917, ex char. (T.: Spegazzini
$. n.)
R. boliviana Met. in Fedde, Rep. Spec. Nov. 20:122. 1924. (T.: Hieronymus & Lorentz
962, photo!)
Shrubs or small trees 2—6 m. tall; branches 2-, 3- or 4-chotomous, terete,
sparsely lenticellate, the bark grayish and slightly wrinkled longitudinally, the
nodes with glands confined to the leaf-axils. LEAVES in whorls of 3 or 4, scarcely
anisophyllous, long-petiolate, elliptic, acuminate, attenuate at the base, 4-12 cm.
long, 1-4 cm. broad, membranaceous to subcoriaceous, opaque on both surfaces,
the secondary veins many, arcuate, rather obscure above and more distinct beneath;
petioles 1-2 cm. long. INFLORESCENCES terminal, many-flowered, corym
peduncles slender, 2-4 cm. long, glabrous, bracteate, the bracts ovate, acute, about
1 mm. long, the pedicels slender, 3-6 mm. long, glabrous. FLOWERS conspicuous;
calyx deeply 5-lobed, the lobes cuspidate, acuminate, about 2 mm. long; corolla
[Vol. 43
322 ANNALS OF THE MISSOURI BOTANICAL GARDEN
Fig. 20. Rauvolfia schueli Speg. (Venturi 9634)
salverform, white, fragrant (Pearce), the tube slender, 5-6 mm. long, about 1.5
mm. in diameter, glabrous without and sparsely pilose within near the throat,
constricted at the throat, the lobes ovate-oblong, acute, 3.0—3.5 mm. long, about
1 mm. broad; stamens 5, inserted near the throat, the anthers ovate, acute, about 1
mm. long, subsessile; ovary 2-carpellary, hemisyncarpous, ovoid, about 2 mm. high,
1.5 mm. broad, the ovules 2 in each locule on an axile placenta, the disc annular,
about 0.5 mm. high, the style linear, 3.0-3.5 mm. long, glabrous, the stigma-head
cylindrical, with a hairy corona at the base, about 0.5 mm. high, 2-apiculate.
I have not seen any fruits. Spegazzini has described the drupes as reniform,
about 10 mm. in diameter and 5-6 mm. high, blue-black. Markgraf has described
the mericarps as subglobose, 2-seeded, 5 mm. broad, 7 mm. high, black.
Densely branching shrubs or small trees in open woods and in sandy loam;
flowering from May to October. The Andean regions of Bolivia and northern
Argentina.
CoMMON NAME: Bolivia—Lecheron del Monte.
I have not seen Spegazzini’s type of R. schueli. All efforts to locate the speci-
men in the important herbaria of Argentina have been in vain. However, Dr.
1956] e
RAO—REVISION OF RAUVOLFIA 323
Theodoro Meyer of the National University of Tucumán, who has collected the
species from both type localities and has studied them extensively, informs me that
he considers R. boliviana of Markgraf synonymous with R. schueli. Evidently
Markgraf was unaware of Spegazzini's species when he described R. boliviana. I
agree with Dr. Meyer and am including R. boliviana under R. schueli.
TINA. JUJUY: San Pedro, near Río Lavagin, southeast of San Pedro de pier.
omnid 3 Beetle 22524, ` (GH, UCL); Sierra de Santa Barbara, Venturi 9634, (GH, S).
SALTA: Yaquiasmé, Meyer 18.003 (
Borivia: precise locality not known, Pearce s. n., K. no. 2253/54 (K).
Fig. 21. Rauvolfia andina Met, (Weberbauer 7112)
21. RAuvoLFIa ANDINA Met, in Fedde, Rep. Spec. Nov. 20:122. 1924. (T.:
Weberbauer 7112!)
Shrubs 5-7 m. tall; branches terete, sparsely lenticellate, the bark gray, slightly
wrinkled, the nodes with glands confined to the leaf-axils. LEAVES in w
3 or 4, scarcely anisophyllous, long-petiolate, elliptic-oblong, acuminate, attenuate
[Vol. 43
324 ANNALS OF THE MISSOURI BOTANICAL GARDEN
at the base, 8—20 cm. long, 2-5 cm. broad, membranaceous, the secondary veins -
many, obscure above, a little more distinct beneath; petioles 2-3 cm. long. 1N-
FLORESCENCES terminal, many-flowered, corymbose; peduncles stout, 1—5 cm. long,
glabrous, bracteate, the bracts ovate, acute, about 1 mm. long, the pedicels slender,
ong, glabrous. FLOWERs relatively large; calyx deeply 5-lo
lobes lanceolate, acuminate, 2.0-2.5 mm. long, the margin glandular-dentate
corolla salverform, white, the tube slender, 8—12 mm. long, about 1.5 mm. in
diameter at the base, glabrous without and within near the throat, but sparsely
villous below the stamens, constricted at the throat, the lobes linear, acute, 5-6
mm. long, 1.0—1.5 mm. broad; stamens 5, inserted near the throat, the anthers
ovate, acute, about 1.5 mm. long, subsessile; ovary 2-carpellary, hemisyncarpous,
cylindrical, about 2.5 mm. high, 2 mm. in diameter, the ovules 2 in each locule on
an axile placenta, the disc annular, about 1 mm. high, the style linear, 7-8 mm.
iie glabrous, the stigma-head cylindrical with a hairy corona at the base, about
1 mm. , 2-apiculate.
I have seen no fruits. Markgraf has described the fruits as obcordate, 1 cm.
high, 1 cm. broad, about 0.5 cm. thick, red in the immature state, and 2-seeded.
On rocky and stony soil in river valleys, 1200-1700 m. altitude; flowering in
April and May. Andes of Peru.
PERU. CAJAMARCA: Cajamarca, cerca a San Miguel, camino
Ferreyra 7086 (US); Jača, pls of the R. Bade sar near E arg pates
7112 (GH, US);
A
0
SUBSERIES 3. GRANDIFLORAE (Mgf.) A. S. Rao, stat. nov.
$ GRANDIFLORAE Mgf. in Fedde, Rep. Spec. Nov. 20:116. 1924.
KEY TO THE SPECIES
A. Corolla-tube 10—20 mm. long, 2-3 times as long as the corolla-lobes, inflated 2-3 mm.
below the orifice
B. Leaves oblance der < aa elliptic, 5-20 cm. long. Peduncles any-
flowered. Corolla- 2.5—3.0 times as long as the een SCC fin or
agp pre 1.0-1.5 m
Leaves distinetly pe ZE coriaceous, Corolla-tube 10-12 mm. ae Frui
reniform. Colombia and Peru Ce senctorum
CC. Leaves obscurely pails, membranaceous. Corolla-tube 12-20 mm. ie ruits
obcordat astern Brazil. 23. R. grandiflora
BB. Licet horae o or elliptic, 2—4 cm. long. Peduncles 1- to 6-flowered. Corolla-tube
abou as long as ‘the Teire a calyx-lobes lanceolate, 2.5-3.0 mm. long.
a Brasil 24. R. peucifolia
AA. Corolla-tube 4—9 m about as long as or 1.5 times as long as the RARE lobes
(except in R. pol dE mi) not inflated or inflated immediately at the orifice
ves sessile or with petioles 2-10 mm. lon
E. Leaves sessile or with petioles 2—4 mm. l rolla-tube 8—10 mm. lon;
. Leaves membran: Sieg ovate to ablong-ovate, ee gem Brazil.
R. —
. Leay tay ves pp broadly or narrowly elliptic; petioles 2-4 mm.
South-central Seit: Ld BT 36. ve
EE. Leaves with petioles 5-10 mm. long. Corolla-tube 4-7 mm. long.
1956]
RAO—REVISION OF RAUVOLFIA 325
G. Leaves prophe attenuate at the base, upper surface opaque, secondary
veins distinct, 4-5 mm. apart. Inflorescences longer than the subtending
leaf, with SE or quaternately divided peduncles. — Calyx-lobes
oblong-ov vate, obtuse. Venezuela BF KR. b pk
. Leaves gradually attenuate " m base, upper surface lustrous, with. si
disti nct secondary 1- . apart, or
obscure onda e Wi
peduncles, pue as deos as the subtending leaf. Calyx-lobes ovate, acute
d.
unde
H. Laon
num:
tà
mostly in voie z a ëmge upper surface lustrous with in-
erable secondary v . apart. Pedicels stout, 4-8 mm.
long; calyx-lobes LOMA. Acte R. nitida
HH. Leaves mostly in whorls of E disci or obovate, upper po
opaque with obscure secondary veins. Pedicels slender, 10-15 mm.
long; calyx-lobes acute.
I. Leaves membranaceous or subcoriaceous. Cymules d An gat
tube about 2 times as long as the corolla-lobes; ovary f E
base, Nube Brazil ei hunk
II. Leaves coriaceou ymules corymbose; corolla- = Fog as long
^ the rollos ovary e almost to the
Sr e Génie oupe and Dominica R. biauriculata
DD. Leaves e prole 12-40 mm. lon,
J. Leaves obovate or pe BRS coriaceous, opaque on both
deeg
K. Dave usually in whorls of 4, obovate, secondary zeg arcuat
yx-lobes acute; corolla purpurascent. Panama... T 3 Eeer
in wh
n whorls o
veins transverse. Calyx-l obes obtuse; corolla white.
Cost ica £N. qum
JJ. Leaves elliptic, coriaceous or membranaceous, opaque on both
urfaces or lustrous abcve.
L. Leaves coriaceous, lus A Seconda xn - MENT
peduncles slender. Corolla slende
diameter at the base, reddish- vets calya-lobes T
long. Ovary ioca arpous. and Bolivia....... i ri "praecox
LL. Leaves membranaceous, opaque on iod s urfaces. Seconda
"ed tertiary peduncles stout. Coro lla abe stout, about 1.5
. in diameter at the base, white; calyx-lobes about 1.5 mm.
va Ovary apocarpous. Deg ve eet 34. R. sellowii
2. RAUVOLFIA SANCTORUM Woodson, in Ann. Mo. Bot. Gard. 18:543. 1931.
(T.: Killip & Smith 15392!)
Trees 3—4 m. tall; branches slender, drooping, terete, the bark olive-gray, gla-
brous, the nodes with pectinate glands confined to the leaf-axils. LEAVES in whorls
of 5, rarely 4, slightly anisophyllous, long-petiolate, elliptic to elliptic-oblanceolate,
acuminate, cuneate at the base, 8—20 cm. long, 3-8 cm. broad, coriaceous, the
secondary veins equally prominent on both surfaces, many, arcuate; petioles stout,
1-3 cm. long. INFLORESCENCES terminal, solitary or paired, few-flowered;
peduncles slender, 2-5 cm. long, glabrous, bracteate, the bracts ovate, acute, about
1 mm. long, the pedicels slender, 4-8 mm. long, glabrous. FLOWERS conspicuous;
calyx deeply 5-lobed, the lobes ovate, acuminate, about 1.5 mm. long; corolla
infundibuliform, white, the tube slender, 10-12 mm. long, about 1.5 mm. in
diameter, glabrous without and within the orifice, but distinctly pilose near the
tip and base of the stamens, the throat dilated 2-3 mm. below the orifice, the lobes
[Vol. 43
326 ANNALS OF THE MISSOURI BOTANICAL GARDEN
Fig. 22. Rauvolfia sanctorum Woodson (Killip & Smith 15392)
obovate-oblong, about 4 mm. long, 2 mm. broad; stamens 5, inserted 2-3 mm.
below the orifice, the anthers ovate, acuminate, about 2 mm. long, subsessile;
ovary 2-carpellary, connate at the base, obovoid, about 3 mm. high, 2 mm. broad,
the ovule 1 in each locule on a ventral placenta, the disc annular, about 1 mm.
high, the style slender, 4-5 mm. long, glabrous, the stigma-head broadly drum-
shaped, about 1 mm. high, obscurely bilobed. Frurrs reniform, 2-seeded, the lobes
13-15 mm. high, 7-9 mm. broad, the stones broadly ellipsoid, a little compressed,
faintly rugose, the seed albuminous, the testa membranaceous, the albumen fleshy,
the embryo erect, the cotyledons ovate, obtuse, about 4 mm. long, 2 mm. broad,
the hypocotyl terete, as long as the cotyledons, superior. e
In dense forests, along the slopes of hills, 100 to 1500 m. altitude; flowering
and fruiting November to December. Colombia and Peru.
This is closely related to R. grandiflora but differs from it by its stout- petioled,
somewhat leathery leaves, the few-flowered inflorescences, and the kidney-shaped
fruits. The geographical distribution is also different.
1956]
RAO—REVISION OF RAUVOLFIA 327
OLOMBIA. SANTANDER: northern slopes of Mesa de los Santos, Killip & Smith 15315
(GH, NY, US), 15392 (A, GH, MO, NY, S, US).
Peru. LORETO: Mishuhuaca near Iquitos, Killip & Smith 29928 (US).
23. RAUVOLFIA GRANDIFLORA Mart. ex A. DC. in DC. Prodr. 8:341, 1844. (T.:
Martius 908!)
R. affinis Muell.-Arg. in Mart. Fl. Bras. 61:34. 1860. (T.: Sellow s. n., photo!)
R. affinis var. subulata Muell.-Arg. l. c. ex char. (T.: Sellow 315)
[e
a PS
LES
S
Fig. 23. Rauvolfia grandiflora Mart. ( Martius 908)
[Vol. 43
328 ANNALS OF THE MISSOURI BOTANICAL GARDEN
Shrubs 1—6 m. tall; branches 2-, 3- or 4-chotomous, terete, glabrous, the nodes
with pectinate glands confined to the leaf-axils. Leaves in whorls of 3 or 4,
slightly anisophyllous, shortly petiolate to subsessile, obovate to oblong-lanceolate,
acuminate, cuneate at the base, 5-15 cm. long, 1.5—4.5 cm. broad, membrana-
ceous, the secondary veins equally evident on both surfaces, several, arcuate;
petioles slender, 2-5 mm. long, or absent. INFLORESCENCES terminal and lateral,
few- to many-flowered; peduncles slender, 1-4 cm. long, glabrous, the pedicels
slender, 8-12 mm. long, glabrous. rrowrms relatively large; calyx deeply 5-lobed,
ovate, acute to acuminate, about 1.5 mm. long; corolla salverform, white to pale
violet (fide Mendes Magalhaes), the tube slender, 12-20 mm. long, about 2 mm.
in diameter at the base, glabrous without and within near the orifice, but villous
at the tip'and near the base of the stamens, the throat dilated from 3-4 mm.
below the orifice, the lobes ovate, obtuse, 4-8 mm. long, 2-3 mm. broad; stamens
5, inserted 3—4 mm. below the orifice, the anthers ovate, acuminate, about 2 mm.
long, subsessile; ovary 2-carpellary, fused at the very base, obovoid, about 2.5
mm. high, 2 mm. in diameter, the ovules 1—2 in each locule on a ventral placenta,
the disc annular, about 1 mm. high, the style columnar, 8-12 mm. long, glabrous,
the stigma-head cylindrical, about 1.5 mm. long, obscurely bilobed. Frurrs ob-
cordate, 2-seeded, the lobes ovoid, 15-20 mm. long, 10—15 mm. broad, (often
only one carpel developing, then the fruit ovoid), the stones ovoid, slightly com-
pressed, faintly rugose, the seed ovate, albuminous, the testa membranaceous, the
albumen fleshy, the embryo as long as the cotyledons.
Near river banks and on road-sides. Flowering from September to December;
ripe fruits in January and February. Eastern Bazil.
This species can be distinguished from the very closely related R. sanctorum
by its membranaceous, extremely short-petioled, almost sessile leaves, the more
open and many-flowered inflorescences and the obcordate fruits.
There is considerable variation in the size and number of leaves per node, as
also in the branching of the peduncles and the relative proportion between the
corolla-tube and the lobes of the flowers. In view of this, I see no justification
for separate recognition of R. affinis, and R. affinis var. subulata, both of Mueller-
Argoviensis. I am hence treating them as synonymous with R. grandiflora.
BRAZIL. BAHIA: Blanchet 105 (P), 211 (G), 2314 (G, P) ; Muritiba, Blanchet 3468
(W). EsPIRITO SANTO: Rio Doce, Kuhlman 121 (RB); Goitacozes, Rio Doce, Kuhlmann
6 (RB); Minas GERAEs: Belo Horizonte, Governador Valadares, Mendes Magalbaes
4-431 (MO). PERNAMBUCO: Dois Irmáos, Rio Garca, Autenour 27 (RB); RIO DE
JANEIRO: Rio de Janeiro, Baretto 4014 (F), Glaziou 639 (P), 7751 (P), Horto Florestal,
x: sep e th tege 42421 (RB); Santa Maria Magdalena, T
ino » SŁ. Hilaire b?r F, P, : N: Martius
908 (G, GH, L, M, MO, P. v). 97 ( US). PRECISE LOCALITY NOT KNOW
1956]
RAO—REVISION OF RAUVOLFIA 329
Tli
IES
Di
VO
KO
(CN
éi
if
LE?
SS
E
Se N JJ KLEE
y
A, i W)
=> i 7 Ki
SN 1:
SEA NS es
gen
HS n
= i D
z rt] KES
- à
E
CN
Las
Fig. 24. Rauvolfia paucifolia A. DC. (Blanchet 2920)
24. RauvoLriA PAUCIFOLIA A. DC. in DC. Prodr. 8:340. 1844. (T.: Blanchet
2920!)
s ?; branches 2- or 4-chotomous, slender, terete or slightly 4-angular,
lenticellate, the nodes relatively distant, with pectinate glands confined to the
leaf-axils. Leaves in whorls of 4 or sometimes 3, slightly anisophyllous, shortly
petiolate to subsessile, elliptic or obovate, acuminate, attenuate at the base, 2-4
cm. long, 1-2 cm. broad, membranaceous, the secondary veins 6—10 pairs, arcuate,
distinct on both surfaces; petioles slender, 2-4 mm. long, or absent. INFLORES-
CENCES terminal or lateral, few-flowered; peduncles slender, 0.5—2.0 cm. long,
glabrous, minutely bracteate, the pedicels slender, 8-12 mm: long, glabrous.
FLOWERS relatively large; calyx deeply 5-lobed, the lobes lanceolate, acuminate,
2-3 mm. long; corolla salverform, the tube slender, 10-12 mm. long, about 2
mm. in diameter at the base, glabrous without and within near the orifice, but
villous at the tip and near the base of the stamens, the throat dilated 2-3 mm.
below the orifice, the lobes ovate, obtuse, 4-5 mm. long, 2.0-2.5 mm. broad;
stamens 5, inserted 2-3 mm. below the orifice, the anthers ovate, acute, about
1 mm. long, subsessile; ovary 2-carpellary, fused at the very base, subglobose,
[Vol. 43
330 ANNALS OF THE MISSOURI BOTANICAL GARDEN
about 2.5 mm. in diameter, the ovules 2 in each locule on a ventral placenta, the
disc annular, about 1 mm. high, the style columnar, 3-4 mm. long, glabrous, the
stigma-head subcapitate, about 1 mm. high, obscurely bilobed. FRUITS not
known.
On hills near rivers and streams; flowering during September. Eastern Brazil.
Rauvolfia paucifolia is easily distinguished from all its related species by its
slender branches with rather distant nodes and fairly small leaves, and the very
few-flowered short inflorescences.
BrazIL. BAHIA: Jacobina, Serato do Rio Sao Francisco, Blanchet 2920 (G, W).
ESPIRITO SANTO: Serra do Itabapoana, Glaziou 11182 (P). .
25. RAUVOLFIA sESSILIFOLIA S. Moore, in Jour. Bot. (Brit.) 42:103. 1904. (T.:
Robert 494, photo!)
Plants with slender, terete, lenticellate, glabrous branches, the nodes with
glands confined to the leaf-axils. Leaves in whorls of 3 to 4, scarcely anisophyl-
lous, sessile, ovate to ovate-oblong, acute to shortly acuminate, narrowed at the
base, 5-8 cm. long, 2-4 cm. broad, membranaceous, the secondary veins 10—16
pairs, slightly arcuate, evident above, obscure beneath. INFLORESCENCES ter-
minal and lateral, many-flowered, corymbose; peduncles slender, about 3 cm.
long, glabrous, bracteate, the bracts subulate, about 2 mm. long, the pedicels
slender, about 5 mm. long, glabrous. FLOWERS conspicuous; calyx deeply 5-
lobed, the lobes ovate, acute, about 1 mm. long; corolla salverform, the tube
slender, about 10 mm. long, glabrous without and within, except near the sta-
mens, pilose; stamens 5, inserted near the throat, the anthers linear, about 1.5
mm. long; ovary 2-carpelled, oblong, obtuse, about 2 mm. high, the disc annular,
about 1 mm. high. Frurrs not known.
Flowering in September. South-central Brazil.
According to Spencer Moore, R sessilifolia is closely related to R. weddelliana,
but differs from it by its ovate or ovate-oblong sessile leaves, its shorter pedice
and the corolla-tube about twice as long as the lobes.
26. RAUVOLFIA WEDDELLIANA Muell.-Arg. in Mart. Fl. Bras. 61:32. 1860. (T:
Weddel 2966!)
R. elliptica Malme, in Bihang till K. Sv. Vet.-Akad. Handl. Afd. TIL 2419:13, 1899. (T.:
me 1444B!)
Shrubs or subshrubs 0.5—1.0 m. tall; branches few, slender, terete, glabrous,
not lenticellate, the nodes with pectinate glands confined to the leaf-axils.
LEAVES in whorls of 3 to 4, scarcely anisophyllous, shortly petiolate, broadly to
narrowly elliptic, acute to short-acuminate, narrowed at the base, 5-11 cm. long,
1-4 cm. broad, subcoriaceous to coriaceous, the secondary veins 12-18 pairs,
slightly arcuate and meeting on either side to form a delicate marginal vein,
a little obscure above, more distinct beneath; petioles slender, 2-4 mm. long.
1956]
RAO—REVISION OF RAUVOLFIA 331
INFLORESCENCES terminal and lateral, geminate or ternate, many-flowered, co-
rymbose; peduncles slender, 3—4 cm. long, glabrous, bracteate, the bracts lanceo-
late, about 2 mm. long, the pedicels slender, 5-12 mm. long, glabrous. FLOWERS
conspicuous; calyx deeply 5-lobed, ovate, acute, about 2 mm. long, corolla
d
Se
p t
T A.
ES
e
AE
L
LS
aT |
if
4
N
x
uL
be
Fig.25. Rauvolfia weddelliana Muell.-Arg. (Hassler 10696)
[Vol. 43
332 ANNALS OF THE MISSOURI BOTANICAL GARDEN —
salverform, roseate (fide Hassler), the tube slender, cylindrical, 8-9 mm. long,
about 2 mm. in diameter at the base, glabrous without, villous within near the
throat and near the stamens, the lobes ovate, broadly acute to obtuse, 4—5 mm.
long, about 2 mm. broad; stamens 5, inserted near the throat, the anthers ovate,
acute to acuminate, about 2 mm. long, subsessile; ovary 2-carpellary, hemisyn-
carpous, subspherical, about 2 mm. in diameter, the ovules 2 in each locule on an
axile placenta, the disc annular, about 1 mm. high, the style columnar, 3-4 mm.
long, glabrous, the stigma-head subturbinate, about 1 mm. high, obscurely
bilobed. Fruits reniform, 2-seeded, the lobes obovoid, 14-16 mm. long, 10-13
mm. broad, often only one carpel developing, then the fruit obovoid, the stones
distinctly rugose, the seeds obovate, albuminous, the testa membranaceous, the
albumen fleshy, the embryo about as long as the seed, erect, the cotyledons ovate,
obtuse, about 4 mm. long, 3 mm. broad, the hypocotyl terete, about 5 mm. long.
In dry fields and on slopes of hills; flowering from September to February;
ripe fruits in March and April. South-central Brazil and Paraguay.
This species resembles R. paucifolia to some extent, but can be distinguished
from it by its elliptic leaves and long-peduncled, many-flowered inflorescences.
My study of the two type sheets respectively of R. weddelliana and R. ellip-
tica confirms what Spencer Moore had already concluded, that R. elliptica is
conspecific with R. weddelliana. Mueller-Argoviensis has cited, in his original
description, two specimens: Riedel 637 and Weddell 2966. All the Riedel sheets
that I have examined are unnumbered but I have no doubt that these sheets are
isotypic with what Mueller has mentioned. However, the prominently lenticel-
late twigs with oblanceolate leaves, and the flowers with very short corolla-tube
and lobes make this referable to R. babiensis rather than R. weddelliana. Accord-
ingly, I am designating for R. weddelliana, Weddel 2966 (P) as the type.
\
d MATO GROSsO: Santa Anna da Chapada, Malme 1444B (S), Malme s.n. (G,
S, UPS, US); SC locality not known, Robert s. n. (MO); between Goyaz and cud.
Weddell 2966 (P
PARAGUAY. a AY: Sierra de mm Hassler 5044 (A, G, MO, P, S, UC, W);
Sierra de Aman. Hassler 10696 (G, P, W); Cerro Torin, Meyer 18. 541 (MO).
27. RAUVOLFIA STEYERMARKu Woodson, in Fieldiana: Bot. 28?:502. 1953. (T-:
Steyermark 57428!)
Shrubs about 6 m. tall; branches verticillate, terete, glabrous, profusely lenti-
cellate, the nodes with distinct pectinate glands confined to the leaf-axils.
LEAVES in whorls of 3 to 5, scarcely anisophyllous, shortly petiolate, elliptic to
obovate-elliptic, shortly acuminate, obtuse at the base, 5-10 cm. long, 3-4 cm.
broad, subcoriaceous, the secondary veins many, 4—5 mm. apart, slightly arcuate,
equally evident on both surfaces; petioles stout, 5-10 mm. long. INFLORESCENCES
terminal, 3- or 4-nate, much branched, many-flowered; peduncles stout, 4-5 cm.
long, glabrous, bracteate, the bracts ovate, acute, about 1 mm. long, the pedicels
stout, 3-5 mm. long, glabrous. rrowrms relatively small; calyx deeply 5-lobed,
1956]
RAO—REVISION OF RAUVOLFIA 335
the lobes oblong-ovate, obtuse, about 2 mm. long, corolla salverform, greenish-
white, fragrant (fide Steyermark), the tube slender, 5-6 mm. long, about 1 mm.
in diameter at the base, glabrous without, villous within near the throat and the
stamens, the lobes oblique-ovate, rounded, about 2.0-2.5 mm. long, about 1.5
mm. broad; stamens 5, inserted near the throat, the anthers ovate, acute, about
1 mm. long, the filament about 1 mm. long; ovary 2-carpellary, apocarpous,
obovate, about 2 mm. high, 1.5 mm. in diameter, the ovules 2 in each locule on
a ventral placenta, the disc annular, about 1 mm. high, the style columnar, 2.5-
3.0 mm. long, glabrous, the stigma-head subcapitate, about 1 mm. high, obscurely
bilobed. FRUITS not known.
Miro =
lll
EA e H
N ini
EEN
ES
Sr,
Fig. 26. Rauvolfia steyermarkii Woodson (Steyermark 57428)
[Vol. 43
334 ANNALS OF THE MISSOURI BOTANICAL GARDEN
Along river banks, at altitudes between 1820 and 2130 meters; flowering in
July. Venezuela.
Rauvolfia steyermarkii is easily distinguished by its verticillate branches, bear-
ing at their tips the ternate or quaternate, profusely flowered inflorescences.
ELA. TACHIRA: between Tabor and Villapaez, along Río Tachira, by Colom-
EE Cni idee boundary, Steyermark 57428 (F, MO).
28. RAUVOLFIA NITIDA Jacq. Enum. Pl. Carrib. 14. 1760; Select. Stirp. Amer.
47. 1763, ex char., not R. nitida Lamarck, 1819.
R. angustifolia Salisb. di 146. Pes nom nud.
R. nitidissima Steud. Nom. Bot. ed. 1. 682. 1821, nom. nud.
R. canescens Descourt. Fl. dt bye 1827, ex one et icon., not R. canescens L. 1762.
R. lanceolat oe Ger in DC. Prodr. 8:337. 1844. (T.: Wydler 290!), not R. lanceolata
R. nitida Ke Fl. Mex. 53. 1887, ex char.
Shrubs or trees 2-15 m. tall; branches 2-, 3- or 4-chotomous, terete, the bark
eros gray, prominently lenticellate, the nodes with pectinate glands confined
e leaf-axils. LEAvEs in whorls of 4, rarely 2, 3, or 5, ovate-elliptic to oblong-
Sëch acute to shortly acuminate, gradually to abruptly attenuate at the base,
3-15 cm. long, 2—5 cm. broad, subcoriaceous, the secondary veins many, almost
transverse, distinct on both surfaces, the upper surface lustrous, the lower opaque;
petioles 5-10 mm. long. INFLORESCENCEs terminal and lateral, few- to many-
flowered, corymbose; peduncles stout, repeatedly dichotomously branched, 1—4 cm.
long, glabrous, minutely bracteate, the pedicels stout, 2-5 mm. long, glabrous.
FLOWERS relatively small; calyx deeply 5-lobed, the lobes broadly ovate, obtuse
to rounded, about 1.5 mm. long; corolla salverform, white (fide J. G. Jack), the
tube stout, 5-6 mm. long, about 1.5 mm. in diameter at the base, glabrous with-
out, villous within along the upper half, the lobes broadly ovate, rounded, 3-4
mm. long, 2-3 mm. broad; stamens 5, inserted near the throat, the anthers ovate,
acute, about 1.5 mm. long, subsessile; ovary 2-carpellary, hemisyncarpous, sub-
globose, about 2 mm. in diameter, the ovules 2 in each locule on an axile placenta,
the disc annular, about 1 mm. hi gh, the style columnar, about 3 mm. long,
glabrous, the stigma-head subcapitate, about 1 mm. high, obscurely biapiculate.
FRUITS subglobose, faintly emarginate, 10-12 mm. long, 15-18 mm. broad, 2-
seeded, the stones ovoid, stout at the base, compressed above, faintly ridged, the
seed ovate, slightly curved at the base, albuminous, the testa chartaceous,
albumen fleshy, the embryo about as long as the seed, the cotyledons ovate, obtuse;
slightly curved at the tip, about 4 mm. long, 2 mm. broad, the hypocotyl terete,
about as long as the cotyledons.
On sandy and stony soil, on hills, in the forests and in pastures, at altitudes
from sea-level to 600 m. Flowering from September to March; ripe fruits in
March and April. The West Indies.
1956]
RAO—REVISION OF RAUVOLFIA 335
Fig. 27. Rauvolfia nitida Jacq. (A. A. Heller 6190)
Common Names: Bois Lait, Glass Wood, Juan Primero, Lechoso, Palo del Leche,
Palo del Rey, Sucheli Blanco.
This species is easily recognized by its glistening, elliptic leaves with in-
numerable close-knit secondary veins, and the inflorescences with repeatedly
dichotomously branching, stout peduncles carrying innumerable small white
flowers on stout short pedicels.
Rauvolfia nitida is one of the two species that Plumier originally founded
under the trinomial Reuvolfia tetraphylla angustifolia. Since Linnaeus’ time much
[Vol. 43
336 ANNALS OF THE MISSOURI BOTANICAL GARDEN
confusion has occurred on the synonymy of this species. Rendle”? has ably clari-
fied this situation. Recently, however, Feuell?! has been led to a fresh error in
mistaking Descourtilz's R. canescens (= R. nitida) for R. canescens L., and has
consequently ascribed the poisonous properties of R. mifida to R. canescens.
ave seen two specimens from the Trinidad Botanical Gardens. These are
obviously the same that Cheesman?? refers to in his account of Apocynaceae.
Since there has been no further collection of R. nitida from Trinidad, Cheesman
has concluded that this is a doubtful record.
UNITED STATES. FLORIDA: Miami, Coconut Grove, Fennell 651 (MO) (Cultivated).
ANTILLEs.—
BaHaMas: Brace Hm RN NEW PROVIDENCE: Delaport, N. L. Britton & Brace
306 (A, GH, MO, NY
CUBA. HAVANA: pe SC 3696 (Ru: Sagra s.n. (P, W). ISLE oF PINES: N. L.
Britton & Wilson 14864 (NY, US). MarANzas: Matanzas, N. L. & E. G. Britton &
(eas (NY). ORIENTE: Baracoa, Ekman C eg (S), 5145 (P, S), Leon 11863, 12143
); Bayate, Ekman 6112 (S); Cupey, Ekman 6325 (S); Papayo, Ekman ae 4 (S);
i ui, Ekman 9390 (S). SANTA CLARA (LAS VILLAS): Cienfuegos, Combs f GH,
MO, NY, P, US), Jack 5089 (A, S, US), 5154 (A, P, US); Soledad, ër p. ër? P),
5427, 5685 (A, P, US), 5653 (A), 6735 (UŠ), 7426 (A, S, US), Howard 6591 (GH
TA Mina Carlota, Howard ps (GH, MO, NY, UC, US), 5604 (GH, NY). seta
OCALITY UNKNOWN: Rugel 751 (NY), Wright s.n. (S — 1364 (G, GH, MO,
NY. P, US), 400 (S), TN € ep. P). Ak
MAICA I Mountains, below Berurcks, Parkins 1195 (A, GH), hillside, Potsdam
to ious Leap p, N. L. Britton 1131 (NY), Santa Cruz Mountains, Harris S),
TOR (NY), 9743 (NY, US); Tepe near Troy, Harris 11071 (NY, US), 11113 (NY,
+ PRECISE LOCALITY UNKNOWN: Alexander s. n. (GH, NY, P
AU
eege Ge eg Azua, J. N. Rose 4054 (NY, ku ; Barahona, Sek 318 (GH,
E , Von Turn ckbeim 2831 (GH, L, MO, P, S, US, W); PAIS
Salt e A G, ae ie 126 (US), 160 (GH); eg + Abbott s.
5); Higuey, R. A dide 9770 (GH, MO, NY), Higuey to Gato,
Taylor 449 (NY); Ces SE R. A. Howard 12140 uet Santiago de los
Caballeros, Jimenez 2706 (MO); Sierra: del Pala, Eggers 1900 (L, M, P). PRECISE LOCAL-
ITY UNKNOWN: Ead ers 2518 (NY, U s Ekman ¡Po S US), pee s. n. qu Sagra
. (P), Von Tuerckbeim 2692 (M, NY pi 3485 L, M, MO, NY, P, S, US, W),
1385bis (GH), Valeur 678 (MO, NY, P, S, US), Wright 2:9 (US), Wright, Perry 8
Brummel 407 (US).
UERTO (ies HUMACAO: Coamo to Caguas, N. L. Britton $ Cowell 1380 (NY);
vicinity of Coamo Springs, N. L. A E. G. Britton & Brown 6424 (NY, P, US), 6481 (NY),
Underwood & Griggs 460 (NY, U ; Humacao, Blanner 50 (P); near San Germán, Miller
1626 (US); abode, Sintenis Zeie (G, M, P, S). AcUADILLA: Rincon, Sintenis 5680 (L, P).
GUAYAMA: Guayama, Kuntze 511 (NY); Cayey, Sintenis 2372 (G, L, MO, P); Jobos, Goll
H 7 ak 8 us MAYAGUEZ: Monte Mesa, N. L. xus & Hess 2716 (NY, US); Río de
NY
522 (US); road from Yauco to Guayanilla, Underwood e iggs 509 (NY, US). sAN
JUAN: Bayamon, Tbe Hellers 403 (A, NY, US), Sintenis Hm (GH, M, NY, S, US),
endle, A. B., Proc. Linn. Soc. Lond. 3;
S M + I "ia Colon. P e de er ee
52 Cheesman, , in Mor t Trinidad tá See 23:153. 1947.
1956]
RAO—REVISION OF RAUVOLFIA 337
I194bis (G); Vue Pena, Stevenson me (US), Point Cangrejos, Stevenson 522 (US).
PONCE: Ponce to Penuelas, N. L. & E. G. Britton & Marble 1 1753 (GH, MO, NY, US);
Penuelas, Sintenis 4786 (M, S, US, Y); fal Heller 6190 (A, GH, L, MO, GE P, Ab
VIEQUES ISLAND: Santa Maria to Caballo Colorado, Me tan 2680, 2684 (NY ONA
"oie Stevens 6318 (NY). PRECISE LOCALITY UNKNOWN: Ch 200 ^ in Plée
T ds Roe s.n. (P), L. C. Richard s. n. (P), stes n.
OIX: SÉ KEEN Garden, Raunkiaer s. n. (U enzon s. n. (MO), A. E. Rick-
bd eg (GH, MO, NY, UG, US), J. J. Kier 428 (MO, P, US); frederickated,
J. N. Rose 3515 (NY), Thompson 98 (GH, NY), 803, 876 (S).
j S).
Pearl to Bonne Resolütión, E. G. Britton % Marble 1325 (MO, NY,
US); near e Solbjeck. Eager s.n. (S); Signal Hills, Eggers 385 (G, L, M, P, US, W).
PRECISE LOCALITY U ww: Bertero s. n. (M), Krebs s.n. (NY), L. C. Richard s.n.
(P), SER (UP >
IRGIN ISLANDS t. John, Morrow 167 (US); Tortola, Belmont sand dunes, Fishlock
430 (NY), Shafer 1136 1178 (NY, US), N. L. 8 E. G. Britton & Kemp 80 (NY).
NTILLES (WEsT IES): precise locality unknown, Swartz s. n. (S), Sessé & Mogino
675, 1449 (F), Ridley n. (P), Vabl 5209 (L).
UADELOUPE: “rapes n. (L)
MARTINIQUE: Plée s js
. — TRINIDAD: Trinidad Bolas Garden Herb. 1931 (US); woods of Chaguanas, Prestoe
1448 (NY).
29. RAUVOLFIA BAHIENSIS A. DC. in DC. Prodr. 8:339. 1844. (T.: Blanchet
3186!)
Plants with trichotomous, rarely dichotomous, terete, glabrous, prominently
lenticellate branches, the nodes with distinct pectinate glands confined to the
leaf-axils. Leaves in whorls of 3, oblanceolate to obovate, shortly acuminate,
cuneate at the base, 3-12 cm. long, 1.0—4.5 cm. broad, subcoriaceous, the sec-
ondary veins barely visible on both surfaces; petioles 5-10 mm. long. INFLOR-
ESCENCES terminal, geminate or ternate, few-flowered, umbellate; peduncles
slender, 3-4 cm. long, glabrous, minutely bracteate, the pedicels slender, 10—15
mm. long, glabrous. FLOWERS relatively small; calyx deeply 5-lobed, the lobes
Ovate to deltoid, acute, about 2 mm. long; corolla infundibuliform, the tube
slender, 6-7 mm. long, about 2 mm. in diameter at the base, glabrous without,
densely villous nearly to the base within, the lobes ovate, broadly acute, 4-5 mm.
long, about 2 mm. broad; stamens 5, inserted near the throat, the anthers subulate,
about 2 mm. long, subsessile; ovary 2-carpellary, free nearly to the base, obovoid,
about 2.5 mm. high, about 2 mm. in diameter, the ovules 2 in each locule on a
ventral placenta, the disc annular, about 1 mm. high, the style columnar, 3—4
mm. long, glabrous, the stigma-head subcapitate, about 1 mm. high, obscurely
bi i own.
Near river banks and on road-sides; flowering during December. Eastern Brazil.
The densely lenticellate, trichotomous, terete twigs, with the terminal, prom-
inently umbellate inflorescences carrying long-pedicelled flowers, serve to distin-
guish this species from the related Rauvolfias.
[Vol. 43
338 ANNALS OF THE MISSOURI BOTANICAL GARDEN
E ULIIFEEREFEFTLITI
Fig. 28. Rauvolfia babiensis A. DC. (Blanchet 3186)
BRAZIL. , BAHIA: basin of Rio Santa Ana, road to Pontal, Krukoff 12698/63 (NY);
precise locality not known, Blanchet 3186 (G, W); Riedel s. n. (G, GH, P, W).
30. RAUVOLFIA BIAURICULATA Muell.-Arg. in Linnaea 30:396. 1860. (T.: Sieber
s. n. !)
Shrubs or trees 3-9 m. tall; branches 2- or 3-chotomous, terete, glabrous, the
bark gray, wrinkled longitudinally, slightly lenticellate, the nodes with distinct
pectinate glands in the leaf-axils only. Leaves in whorls of 3, rarely 4, shortly
petiolate, oblanceolate to obovate, shortly acuminate to caudate, cuneate at the
base, 5-12 cm. long, 2.0—3.5 cm. broad, coriaceous, the secondary veins obscure on
both surfaces, the two surfaces opaque; petioles 5-10 mm. long. INFLORESCENCES
terminal, few-to many-flowered, corymbose; peduncles repeatedly 2-branched,
1956]
RAO—REVISION OF RAUVOLFIA 339
Fig. 29. Rauvolfia biauriculata Muell.-Arg. (Eggers 674)
slender, 3-5 cm. long, glabrous, minutely bracteate, the pedicels slender, 5-10 mm.
long, glabrous. FLowers relatively small; calyx deeply 5-lobed, the lobes deltoid,
acute, about 1.5 mm. long, the margin minutely glandular-dentate; corolla salver-
form, roseate (fide Howard), the tube slender, 4-6 mm. long, about 1 mm. in
diameter at the base, glabrous without, villous within along the upper half, the
lobes obliquely oblong-elliptic, obtuse, 4-5 mm. long, 2-3 mm. broad; stamens 5,
inserted near the throat, the anthers ovate, acuminate, about 2 mm. long, subsessile;
[Vol. 43
340 ANNALS OF THE MISSOURI BOTANICAL GARDEN
ovary 2-carpellary, fused almost to the top, obovoid, about 2.5 mm. high, 2 mm.
in diameter, the ovules 1 in each locule on an axile placenta, the disc annular,
about 1 mm. high, the style columnar, about 3 mm. long, glabrous, the stigma-
head cylindrical, about 1 mm. high, obscurely bi-apiculate. FRUITS obcordate,
2-seeded (often only one carpel developing, then the fruit ellipsoid and 1-seeded),
the lobes ellipsoid, 12-15 mm. long, 5-8 mm. broad, the stones distinctly rugose,
the seed albuminous, the testa membranaceous, the albumen fleshy, the embryo
erect, the cotyledons ovate, obtuse, about 3 mm. long, 2 mm. broad, the hypocotyl
terete, about as long as the cotyledons.
In dense humid forests, on hills, at altitudes between 500 and 800 m. Flower-
ing from June to December; ripe fruits in January to March. Antilles: Dominican
Republic, Dominica, and Guadeloupe.
CoMMON NAME: Bois Lait-montagne.
Rauvolfia biauriculata resembles greatly R. bahiensis, but can be differentiated
from it by its slightly lenticellate, wrinkle-barked twigs with markedly leathery
leaves, and the terminal dichotomously branching peduncled, corymbose inflores-
- cences bearing flowers with the corolla-tube almost as long as the lobes.
I have seen the two Ritter and Sieber specimens that Mueller has cited in his
description. The Sieber sheet carries distinctly the annotation "Trinitas." How-
ever, Cheesman?? considers this a doubtful record, as he has found no evidence
for the occurence of this species in Trini
ANTILL DOMINICAN REPUBLIC: precise locality unknown, Ritter s.m. (W).
DOMINICA: ‘Placa Valley, Eggers 674 (G, GH, L, M, P, UC, US, W). GUADELOUPE:
ort Baines Jaunes, Stehle, Quentin et Bena 5601 (US), Steble 702 (US); trail from St.
Cloud to Soufriére above Baines Jaunes, Howard 11794 (MO); Grand Bois, Quentin 726
). PRECISE LOCALITY UNKNOWN: Duss 8 (P), 2544 (GH, MO, NY, UPS, US), L. C.
Richard s. n. (P). a lieber s.n. (W).
31. RAUVOLFIA PURPURASCENs Standl. in Trop. Woods 16:11. 1928; Field Mus.
Publ. Bot. 4:255. 1929. (T.: G. P. Cooper 516!)
Stout liana?; branches subterete, glabrous, the bark wrinkled longitudinally,
scarcely lenticellate, the nodes with distinct pectinate glands confined to the leaf-
axils, LEAvEs in whorls of 4, long-petiolate, obovate, broadly acute to shortly
acuminate or obtuse, cuneate at the base, 5-16 cm. long, 3-7 cm. broad, membrana-
ceous to coriaceous, the secondary veins several, arcuate, joining at the margins
to form marginal veins, distinct on both surfaces, the two surfaces opaque; petioles
stout, 12-20 mm. long. INFLORESCENCES terminal, lax, few- to many-flowered,
paniculate; peduncles slender, 3—6 cm. long, glabrous, minutely bracteate, the
pedicels slender, 3-6 mm. long, glabrous. FLOWERs relatively small; calyx deeply
5-lobed, the lobes ovate, acute, about 2 mm. long; corolla salverform, purpurascent
(fide G. P. Cooper), the tube slender, 6-7 mm. long, about 1 mm. in diameter at
the base, glabrous without, villous within near the throat and the stamens, the lobes
9? Cheesman, E. E., in Flora of Trinidad and Tobago 23:153. 1947.
1956]
RAO—REVISION OF RAUVOLFIA 341
Fig. 30. Rauvolfia purpurascens Standl. (Cooper 516)
obovate, broadly acute to obtuse, 5-6 mm. long, 2-3 mm. broad; stamens 5, in-
serted near the throat, the anthers ovate, acute, about 1.5 mm. long, subsessile;
Ovary 2-carpellary, hemisyncarpous, cylindrical, about 3 mm. high, 2 mm. in di-
ameter, the ovules 2 in each locule on an axile placenta, the disc annular, about 1
mm. high, the style columnar, 3—4 mm. long, the stigma-head subcapitate, about
1 mm. high, obscurely bi-apiculate. FRUITS unknown.
Flowering from January to March. Panama: Cricamola Valley. The branches
bearing whorls of 4 long-petioled, obovate leaves with distinct, arcuate secondary
veins, and the terminal, loose, paniculate inflorescences distinguish R. purpurascens
from the related species of Rauvol fia.
^ outil BOCAS DEL TORO: region of Almirante, Cricamola Valley, Cooper 516 (F,
» US).
[Vol. 43
342 ANNALS OF THE MISSOURI BOTANICAL GARDEN
NN]
Fig. 31. Rauvolfia sarapiquensis Woodson (Skutch 3556, and Mora s. n.)
32. RAUVOLFIA SARAPIQUENsIs Woodson, in Ann. Mo. Bot. Gard, 28:271. 1941.
(T.: Skutch 3556!)
Trees about 15 m. tall; branches terete to subangular, the bark dark gray,
glabrous, the nodes with distinct pectinate glands confined to the leaf-axils. LEAVES
in whorls of 3, long-petiolate, oblong-oblanceolate to oblong-elliptic, shortly acu-
minate, attenuate at the base, 15-25 cm. long, 4-7 cm. broad, coriaceous, the
secondary veins several, almost transverse, equally evident on both surfaces, the
two surfaces opaque; petioles stout, 1.5—2.5 cm. long. INFLORESCENCES terminal,
ternate, many-flowered, corymbose; peduncles stout, 5-6 cm. long, 2- or 3-
branched, glabrous, minutely bracteate, the pedicels stout, 4-5 mm. long, glabrous.
1956]
RAO—REVISION OF RAUVOLFIA 343
FLOWERS relatively small; calyx deeply 5-lobed, the lobes broadly ovate, obtuse,
about 2 mm. long, fleshy; corolla salverform, white (fide Skutch), the tube stout,
5-7 mm. long, about 1.5 mm. in diameter at the base, glabrous without, villous
within near the throat and the stamens, the lobes elliptic to ovate-elliptic, obtuse,
about 4 mm. long, 2 mm. broad; stamens 5, inserted near the throat, the anthers
ovate, acute, about 1 mm. long, the filament about 1 mm. long; ovary 2-carpellary,
hemisyncarpous, obovoid, about 2.5 mm. high and 2 mm. broad, the ovules 2 in
each locule on an axile placenta, the disc annular, about 1 mm. high, the style
linear, about 2 mm. long, glabrous, the stigma-head subglobose, about 1 mm. high,
obscurely bilobed. Frurrs slightly bilobed, the lobes obovate, about 18 mm. high,
8 mm. broad, the stones distinctly rugose, the seed albuminous, the testa membran-
aceous, the albumen fleshy, the embryo erect, the cotyledons oblong, obtuse, about
5 mm. long, 2 mm. broad, the hypocotyl terete, about as long as the cotyledons.
On hill slopes at altitudes of about 1500 m.; flowering in February. Costa Rica.
The oblong-elliptic, rather leathery leaves with almost transverse secondary
veins, and the many-flowered, much-branched inflorescences distinguish this species
from the rest of the Central American species of Rauvolfia.
osta Rica. Vara Blanca de Sarapiqui, north slope of central Cordillera, between
Poas and Barba volcanoes, Skutch 3556 (A, MO, US); Buena Vista de San Carlos, H. B.
Mora s. n. (MO).
33. RAUVOLFIA PRAECOX K. Sch. ex Markgraf, in Fedde, Rep. Spec. Nov. 20:112.
1924. (T.: Ule 62561) ; ex Ule in Engl. Bot. Jahrb. 40:400. 1908, nom nud.
Trees 15—30 m. tall; branches subangular to terete, 2-, 3- or 4-chotomous, the
bark gray, lenticellate, the nodes with distinct pectinate glands confined to the leaf-
axils. LEAVEs in whorls of 3 or 4, long-petiolate, narrowly elliptic, acute or acu-
minate to obtuse, attenuate at the base, 5-15 cm. long, 2—5 cm. broad, coriaceous,
the secondary veins several, slightly arcuate, meeting at the margins to form two
marginal veins equally evident on both surfaces, the upper surface lustrous, the
lower opaque; petioles slender, 2-3 cm. long. INFLORESCENCES terminal, often at
the tips of defoliated twigs, many-flowered, paniculate; peduncles ternate, 2- or 5-
branched, 2-4 cm. long, glabrous, minutely bracteate, the pedicels slender, 6—9
mm. long, glabrous. FLOWERS relatively small; calyx deeply 5-lobed, the lobes
ovate, acuminate, about 1 mm. long; corolla salverform, reddish-violet (fide Ule),
the tube slender, 5-7 mm. long, about 1 mm. in diameter at the base, glabrous
without, scantily pilose within near the throat and the stamens, the lobes ovate,
broadly acute to obtuse, 4-6 mm. long, about 2 mm. broad; stamens 5, inserted
near the throat, the anthers ovate, acuminate, about 1 mm. long, subsessile; ovary
2-carpellary, hemisyncarpous, obovoid, about 2 mm. high, 1.5 mm. broad, the
ovules 2 in each locule on an axile placenta, the disc annular, about 1 mm.
high, the style linear, 3-5 mm. long, glabrous, the stigma-head cylindrical, about
1 mm. high, obscurely bilobed. FRUITS not seen.
[Vol. 43
344 ANNALS OF THE MISSOURI BOTANICAL GARDEN
Fig. 32. Rauvolfia praecox K. Sch. (Ule 6256)
On river banks in dense forests; flowering from July to October. Bolivia, Peru
and Western Brazil.
CoMMON NAME: Bolivia—A margo.
Rauvolfia praecox has been confused in the past with R. sellowi, ma
eastern Brazilian in its distribution. It can be distinguished by its long-petioled,
glistening, elliptic leaves and the terminal inflorescences with a profusion of
reddish-violet, slender flowers.
1956]
RAO—REVISION OF RAUVOLFIA 345
BRAZIL. AMAZONAS: near mouth of Rio Embira, tributary of Rio Taracua, lat 7.30°
s., long. 70.15? w., Krukoff 5018 (A, F, G, M, MO, P, S, US).
LIVIA. SANTA CRUZ: Sara, Buena Vista, banks of Rio Surutu, Steinbach 6536 (A,
F, M, MO, S, US), 656r (F, G, MO, S, UC, W).
PERU. LORETO: Iquitos, Ule 6256 (G, L).
Fig. 33. Rauvolfia sellowi Muell.-Arg. (Weir 173)
34. Rauvorri seLLown Muell.-Arg. in Mart. Fl. Bras. 61:34. 1860. (T.
Sellow 238)
[Vol. 43
346 ANNALS OF THE MISSOURI BOTANICAL GARDEN
Trees with branches verticillate, terete, the bark gray, longitudinally wrinkled,
the internodes short, the nodes with distinct pectinate glands confined to the leaf-
axils. LEAvEs in whorls of 4, long-petiolate, oblong-obovate to oblong-elliptic,
obtuse-acuminate, attenuate at the base, 6—20 cm. long, 3—6 cm. broad, membran-
aceous, the secondary veins several, arcuate, equally evident on both surfaces, the
upper surface greenish, the lower dull grayish green; petioles slender, 2-3 cm. long.
INFLORESCENCES terminal, ternate, many-flowered, corymbose; peduncles stout,
3-5 cm. long, 3- or 4-branched, glabrous, minutely bracteate, the pedicels slender,
3-5 mm. long, glabrous. FLOWERS relatively small; calyx deeply 5-lobed, the lobes
broadly ovate, acute, about 1.5 mm. long; corolla salverform, white (fide Sellow),
the tube stout, 5—7 mm. long, about 1.5 mm. in diameter at the base, glabrous
without, scantily villous within near the stamens, the lobes ovate, broadly acute
to obtuse, 2~3 mm. long, about 1.5 mm. broad; stamens 5, inserted near the throat,
the anthers ovate, acute, about 1.5 mm. long, subsessile; ovary 2-carpellary, apo-
carpous, subglobose, about 2 mm. high, 1.5 mm. in diameter, the ovules 2 in eac
locule on a ventral placenta, the disc annular, about 1 mm. high, the style linear,
3-4 mm. long, glabrous, the stigma-head cylindrical, about 1 mm. high, obscurely
bilobed. rnurrs apocarpous, drupelets ellipsoid, 12-16 mm. long, 7-10 mm. broad,
the stones distinctly rugose, the seed ellipsoid, albuminous, the testa membrana-
ceous, the albumen fleshy, the embryo about as long as the seed, erect, the cotyle-
dons ovate, obtuse, about 7 mm, long, 3 mm. broad, the hypocotyl terete, about
as long as the cotyledons.
In dense forests on hill slopes, flowering from September to December; ripe
fruits in February to March. Southeastern Brazil.
Rauvolfia sellowii is distinguished by its twigs with short internodes, the
oblong-elliptic membranaceous leaves, the much-branched many-flowered inflores-
cences, and the ellipsoid apocarpic fruits.
BRAZIL. MINAS GERAES: Cidade de Caldos, Henschen 866 (F, US). PARANA: Cap
bove Dusén 7631 (S). RIO DE JANEIRO: Rio de Janeiro, Glaziou 2915, 604. 5 (P),
SAO PA AULO: REN Ari F biman 7995 (UC), To 1457 &), Sao
E Mello & Mos (S). PRECISE LOCALITY UNKNOWN: rcbell 4678 (K),
Glaziou s. n. (P), Regnell 111866 P, S, UPS, US), Sellow s. n. (K), d 173 (K).
DUBIOUS SPECIES
Rauvolfia arborea Larrañaga, Escritos D. A. Larrañaga, Atlas 1: pl. 132. 1927
(Publ. Inst. Hist. Geog. Uruguay)
Rauvolfia bilabiata Larrañaga, Escritos D. A. Larrañaga, 2:85. 1923.
Neither the illustration of the first species nor the description of the second is
sufficient to refer the above two to any known species.
Rauvolfia oppositiflora Sessé & Moc. in La Naturaleza, Ser. 21:32. 1888. The
description is insufficient to refer this to any known species.
1956] a
RAO—REVISION OF RAUVOLFIA 347
Rauvolfia rhomhofiae Mgf. in Notizblatt 15:384—385. 1941. I have not been
able to examine any representative material of this species. From the original
description, I am inclined to treat it as synonymous with R. sanctorum Woodson.
EXCLUDED SPECIES
Rauvolfia dentata Tafalla, ex D. Don, in Edinb. New Phil. Jour. 10:237. 1831
= CITHAREXYLUM DENTATUM D. Don, Le 1831.
Rauvolfia flexuosa Ruiz & Pav. Fl. Peruv. 2:26. 1799 = CITHAREXYLUM
FLEXUOSUM (R. & P.) D. Don, in Edinb. New Phil. Jour. 10:237. 1831.
Rauvolfia glabra Cav. Icon. 3:50, tab. 207. 1794 = VALLESIA DICHOTOMA
Ruiz & Pav., Fl. Peruv. 2:26. 1799.
Rauvolfia laevigata Roem. & Schult. Syst. 4:805. 1819 = MALOUETIA JAS-
MINOIDES A, DC. in DC. Prodr. 8:379. 1844.
Rauvolfia longifolia A. DC. l. c. 338 = Tonpuzia LONGIFOLIA (A. DC.)
Met in Fedde, Rep. Spec. Nov. 20:112. 1924.
Rauvolfia lycioides Cav. in Anal. Cienc. Nat. 5:69. 1802. This is a species
of Citharexylum according to my observation of the type specimen from the Paris
Herbarium.
Rauvolfia macrophylla Ruiz & Pav. Fl. Peruv. 2:26. 1799 = CITHAREXYLUM
RETUSUM D. Don, in Edinb. New Phil. Jour. 10:237. 1831.
Rauvolfia oppositifolia Spreng. Neue Entdeck. 3:33. 1822 = TABERNAE-
MONTANA OPPOSITIFOLIA (Spreng.) Urb. Symb. Antill. 4:493. 1910.
Rauvolfia pubescens Roem. & Schult. Syst. 4:805. 1819 — CITHAREXYLUM
MOLLE Jacq., fide Index Kewensis.
Rauvolfia spinosa Cav. in Anal. Hist. Nat. 1:43. 1799; Icon. 6:16, tab 520.
1801. My observation of the type from the Madrid Herbarium indicates that this
is also a species of Citharexylum.
Rauvolfia stenophylla Donn. Sm. in Bot. Gaz. 44:115. 1907 = TONDUZIA
LONGIFOLIA (A. DC.) Mgf. in Fedde, Rep. Spec. Nov. 20:112. 1924.
Rauvolfia strempelioides Griseb. Cat. Pl. Cub. 170. 1866 = STREMPELIOPSIS
STREMPELIOIDES (Griseb.) Benth. in Benth. & Hook. Gen. Pl. 2:702. 1866.
INDEX TO EXSICCATAE
(sine numero) to unnumbered
talicized numerals refer to collectors’ numbers, s. n.
d to the species conserved in
lecti Merentheriesl numerals refer to the ibas assigne
this re
Abbott, W. L. s.m. (28). Allemáo, F. 974 (8).
Aguilar, M. H. 138 (3). Allen, P. He ais EC us
Alain, Br 8 (8 IO, 3319, 3673 Anderson, N. J.
o. 4075 (8); 3310, 3319, 3 Anis, ) D. NH |
Alexander, R. G ein. puoi E. ret 735 i 4816, 5860
ard, H. A. 14627 (28). (4): 63
E inis Ch. s.s. (28). Arson, —
348
Asplund, E. s.n., 5125 (3).
Autenour, —. 89627 (23).
Baker, C. F. 173, 215, 608, 667, 846,
5130 (3).
Baker, R. E. D. 1, 2421, 2640, 4963 (8).
Bangham, W. N. 333 (3).
1 (5).
EA K Zug 26262 (3).
jene, e 5337 (4).
Benzon, P. sn (28).
Bernoulli, C. S. & Cario, R. 1835 (3).
Bertero, C. - s. n. (3) ; s. n. (28).
n. (3).
Blanchet, J. S. 1007 (5); 2716, 2769 (6);
105, 211, 2314, 3468 (23); 2920 (24);
186 (29).
Blanner, —. 50 (28).
Bórgeson, F. 668, 23615, 27565 (4).
NEST t 1235; 23618 (4).
E ).
I (1); 12365, 15695,
21467 ( 2 1259 (8).
gem, AT A (28).
= ps eed y 21306: (28y;
NEN. L. & E. G. 28).
Britton, N. L. & S ond W. E. 2460
8
Britton, N. L., & E. G., & Earle, F. S. 9749
ud N. L. & E. G., Earle, F. S. & Gager,
C. S. 0p03 (3).
Britton, N. L. & E. G., & Kemp, J. F. $9
(28).
Britton, N. L. & E. G., & Marble, D. 401,
Sud 2 E 3 (28). ha
& Sha et A. 549
G); a ^s EN 96 (2
Britton, N. L. & dx Powel, —. I327I
3
Britton, N. L. & Cowell, J. F. 9976, —
12688 gu 254, 351 (4); 9087 (
Zen 28).
Britton, N. L., Cowell, Ze F., & Carlos de.
La Terre 13345 (19
ANNALS OF THE MISSOURI BOTANICAL GARDEN
[Vol. 43
Britton, N. L., Cowell, J. F., & Stewardson
Brath, M. "5048 (4); 4571, 6424, 6481
(3).
, F. S., & Gager, C. S.
(28
Britton, N. L. & Earle, F. S. 7562
Britton, N. L., Earle
ad sth ).
ritton, N. L., Earle, F. S., & Wilson, P
x 34 (3); 4599 (19).
i Gager, C. S. 7328 (19).
He s, W. E. 2716 (28).
Britton, N. L. Stevens, F. L., & Hess, W.
SS ^
Britton, N. L. & Wheeler, W. M. ei (4).
Britton, N. L. & Wilson, P. 32 306,
369 (8); STH: Hr gue 24864 Em
Broadway, W. E. 229 (3); sn
103, 109 (4); 2566, "2680, 0010, 9265
Burchell, W.J. 4678 (34).
e D 348 (3); 1654 (8).
Ger cai oe 3823 (18).
E
Car . » 552 (3).
Carvalho, O. de 6 (8).
ipt R. R. 984 (3); 74 (8); 900
11).
Greg Bro. 57 a):
Cesped
ark, O M.
erede Bro. 35
Combs, R. 31
E
, 3605, 4390 (18).
(3); 180 eg 245 (19);
Constantine, D. 2257 (24h
Conzatti, C. 2165, 3991 (3).
Send O.F. gr (3).
ook, O. F. & Doyle, C. B. 654 (3).
ee P. 202 (4); 516 Gi dë
Co
Cowell, e F. oos (28).
Sale R.Q (Astralia) 5428 (3).
dëse: O 30
H
854, 866 (3); Hm rug:
Curtiss, A. H.
aaepe B. E. & Persaud, A. C. s. t. (3)-
Deam, C. C. 6267 (3).
1956]
RAO—REVISION
EC —., Britton, N. L., & Brace, L. J.
28).
Dias, R. de 63 (8).
Diag, A. 5.1 (3):
Drouét, F. 2707 (8).
Ducke, A. s.n., 23 (8); 626, RB30117 (12);
74, 785, 917, 1118, 21 59F, 11391, 11395,
23050, RB3or15 (14); RB$36,
(15); 492, 11038 = RB13208, 16544 =
Dugand, A. 5
4822 (4); 528, 1025, 45. )
Dugand, A. & Bar G. 2307
Dugand, A. 8 Jaramillo, R. 3213, 3217,
3374 (4); 2748, 326
Dugand, A. & Peten, —. Ce ).
Dusén, P. 139 (7); 7631 (34).
Duss, P. 6, 1225, 2614 d 8, 2544 (30).
Echeverria, J. A. 284, 4166 (3).
Eggers, H. 15531 = 15568 (2); 5411 (3);
6461 (4 4); s. n., 183, 1900, 2518 (28);
74 (30).
Kate CA uw 138 (311,3
o. E. L. 348, 825, 944, 5679, 6301,
7963, 8535, 8575, 9760, 10452, 12465,
13419, 16892 (3); 5927, 16859 i
1733, 5013, 5939, 9552, 9085, 1530
(17); 3322, 354 4147, 6145, GE:
8, 9078 (18); 10922, 11842, 12583,
17069 (19); H.919, H.4427, 4440, 5145,
6112, 6325, 9324, 9390, 14294 (28).
Elias, Bro. 1543 (2); 1014, 14189 (4);
1076, 1425
Emrick, G. M. 186 (3).
Espiña, R. (4).
Eyerdam, W. E 434 (3).
eet J Serie: A. 8711 (3);
22524 (
Fairchild, D.
Favrat, —
Fendler, x 636, 2206 (3); 612 (4); 2347
32. (287.
Fennell, LL. 651 428).
Ferreyra, ke P ace 10085, 10949 (3);
)
Mexia, Y. 5208 (3
Fiebrig, K. 1278, 1350, 4034, is (7).
Fishlock, W. C. 75 (4); 430 (28).
Focke, W. O. 1287 (8).
Förström, —. Pes r807 (a)
Fogg,.]. M. 21751, 21757, 21759,
mes ind (3); 21775 (4); 21770
OF RAUVOLFIA 349
Freeman, W. G. 8879 (3).
Fróes, R. de L. 27307 (13).
Fuertes, M. 238 (3); 318 (28).
Ga "ofi H. EE (f):
Gucómeito, De PM (3 >).
1183 fy 639 (23); 1182
4); S-n., 1045, 2915, 6045 (34).
8. (4).
. 517 (4); 517 (28)?
—,. s.m. (3).
Goodding, L. N. 340 (3).
Goudot, M. ng, (3).
Greenman, J. M. & M. T. 5145, 5687,
5958 (3)
Hahn, L. s.n. (3); 1459, 1503 (4).
Harris, W. E, 5850, 9743, 11071 (28).
H nn 353.
Havard, V. (3).
Heller, A. A. 1186 au p (28).
Heller, Mr. & Mrs. A. A 3 (28)
Henschen, 866 34).
Heriberto, lo, 36, 3 54 (4).
Herrera, 41
Heyde, H. T. & Lux, E. 641 2 (3)
Hinton, G. B. 724, 3826, pros 12028
(3).
Hinton, G. E et al. 3582, 5799, 6018 (3).
Hiralal, —. s.n. (3).
EE Kë H. "674 (4).
H , W. H. & Howard, R. A. 4139
€ ?
Hoehne, F. C. 4280 (7); 3430, 7544 (8).
Holt, —. 545 (4 )
Holton, J. F. s.n. (4).
eng "Horesal "RBÓI277, RB62875 (23).
(
un. 5582 (8); 5942, 6104
(18); A 6501; 12140 (28); 11794
).
Howard, R. A. & E. S. 9760, 9770 (28).
Howard, —, Briggs, —, et al. 253, 392
(3).
Howell, J. T. 10155, 10257 (3).
Humboldt, A. von 69 (4).
Jack, 4661, ts pee 7297 0);
ees p (19); 49 ims 5154»
5427, 5685, 6735, Mee i
350
Jaeger, E. C. e (3).
Jahn, A. 179 SUR
aed $, de de 2706 (28).
Ji (3).
35 (4).
66, er? (3).
Johnston, J. R.
Jones, M. E.
Karsten, H. 77^ (3).
sv W. A. 8059 (3); 4560, 4570
“Ken A nL.A. 3:66 AÓ41 (3).
).
mith, A. C. 14161, 14191
(3); 15315, 15302, 29928 (22).
Klug, G. 35 (13).
i ( 8).
Faits; B. A. = DI 2047 (8); E
6698, 8262 2,
8871 (15); 12608/63 (29); 5018 d».
NT $1 (8); 121, 6448,
T (23); 7995 (3 4).
ara, O: V74 (4); 511 (28).
Kupper, W. 1424 (3).
Lanjouw, J. 1255 (14).
Leavenworth, W. C. cf (3).
Bra W. C. & Hoogstraal, H.
1352
Leon, E 138, 1337, 5201, 9563, 12366
(3); 7191, 9223, 0447 (8); 5208, 9246
(19); » 3696, 11863, 12143 (27).
ee
Leon & Charles, Drok, 14 (1
Leon & Clement, Bros. 23046 (1
Leon, Clement, & Alain, E^ 5 (18).
Leon, [ds , et al
Leon, » & Roca, M. QUE 7252 (8);
7712 ve 19).
Leon, Victorin, & Clement, Bros.
20785
M J. 4499 (2); 502 (3); 4286, 4287
Leonard, E. C. 872, 4145, 8914, 9912 (3);
3230, 3367, figs i e
Leo eis CX see n., 13705 (3);
274, 12452, rii 13290, 14275,
14826, 15590 (28).
M P. SA x
Lundell, C. L. 900, 2493, 3399, 4768 (3).
Lundell, C. L. & A. A. 1464 (3 ^ut o
Lyonnet, E. 2185, 2655 (3).
[Vol. 43
ANNALS OF THE MISSOURI BOTANICAL GARDEN
E een i Hs 30 (3);
Malme, G. A. Ze 267995, 2736 (7);
I
Martines- rM G. go (3).
Martius, K. F. P. von e (6); 908 (23).
den Ze Lu
Matuda, E. 16423, bs 16896, 31291
E Extra 5, 2687, 2734, 16273, 16297
dg W.R.& sag A D. 7657 (3).
xon, W. al. 7104, 7156, 7221,
7282, 7545, 7613, 7740 (3).
Mello, J. C. & Mosen, T. CT s. n. (34).
Mendes Magalhaes, —. 4.431 (23).
Merck & Co. s. n. (3);
n. (8).
Mexia, Y. 940 (3); pod T 6415 (13).
Meyer, T. 18.600 (7); 18.003 (20); 18.-
541 (26).
Mille, L 989 (39:
Miller, G. S 1626 (28
Miller, O. O. & TEE LO 43 (4).
266 (3).
Molina, A. 30, 3866, 3867 (3).
Molina, A. & Barkley, F. A. 1808256
eh. R. t 551
(5).
Mutnsción. W. C. 12371, 12601 (3).
Nash, G. V. 892 (28).
Nash, G. V. & Taylor, N. 1522 (3); 1525
(28).
Nelson, E. W. 1872 (3).
Netto, A. 40 (8).
Octavis Solis, —. 89 (3).
Comité C R. SE 5103 (3).
a, J. G. 4023, 4126, 4167, 4678;
pu 7210, mo 7372, 7482 (5).
Otto, E. 804
pe». M. Lx (3);
Palmer, E. s. n., 86, 106, 421, 1030, 1437»
1430 (3).
Palmer, W. died: A 840 (3).
Parkins, LR 17105 (28).
ie Bro. C34 EN
arce, —. S.M., Pie in s
nell, F. W. E 3); 4140 (
denis P. 2046 (3).
n. (3).
n. (4).
Pires, J. M. Ss 4021 (14).
1956]
RAO—REVISION
Pires, J. M. & Silva, N. T. 4784 (16).
Pittier, E. $, B
Pittier, H. I5 , 10282, 13329,
14038 (3); 5970. 7844, 8872,
14035 (4); 9717 (8); 9465,
Plée, A. 610 w; : n. (28).
13388,
11052,
11062
Prestoe, —. ).
Pringle, C. a 4580, 4861, 5068 (3).
Purdie, 40
Purpus, C. A. 2016, 5309, 6143, 7147 (3):
Questol, A.
Quinos, R. M. 789 (3).
Quentin, R. P. E (4); 726 (30).
228 (4).
Raimondi, A.
s. 4. (4); 73 CIDE 4
n. (30).
secker, A. E. 407 (4); 362 (28).
SE Jl. 333 (4); 428 (28).
Ridley, H. CONS Ye T. L & Ramage, G.
3
"S L sm. (12); s.m. (28); s."
9).
Rijgersmaa, H. E. s.n. (4).
Riley, L. A E 125
(2).
o, Jardim Botanico de 47482
(8); 21835; série 4).
itter, —,. 30).
Robert, A. s.m. (26).
Roig, J. T. 1733 (8); 65 (18); 3 (19).
Rojas, T. 175 (3
Rose, J. N. 1575, 14032, 14622, 14680,
dane (3); 3515, 4054 $9
Rose Painter, J. H., & Rose, J. S.
930 5 (3).
Rose, 1 N., Standley, P. C., & Russell, P.
G. 14280 As MÀ
Ross, G. B. s.s. (3).
Kee? w- Y. & H. E. 123, 124 (3).
MM. V. D
ugel, F. ech (3); 751 (27).
Rusby, H. H. 2388, 2389 (8).
r ie la s.n. (3); 414, 815 (19);
S. n.
Saint H A. de b? 197 (23).
Salas, G. 363 (3).
Sargent, F. H. 195 (4); 522 (28).
OF RAUVOLFIA 351
Schiffino, J. 126, 160 (28).
Schomburgk, Robt. $91 (12).
Schott, A. 431,851 (3); s. n. (4).
Sellow, RS ERA S
Sessé, M., Mociño, J. M., et al. 5066, 5067
(3); 675, 1449 (28).
Shafer, J. A. 173, 1647, 2847, 2853, 2886,
3 .
Shimek, B. & Smith, GL 230 is
sc? F. W. 74, 268 7M de (8); s. "t.
. IOIO O
Simmonds, H. G. 3).
gier P. II HE Ze 05, 2806 (4); 1193,
1193 bis, 2372, 3372; 3835, 5036, 5680
a
(28).
Skutch, x d 3556 (32).
Smith, Q L. sA, 127, 055 1336,
(3)
Smith, H. H. 1905, 1906 (2); 1648, 1649,
6018
1
Srivatsava, G. S. na (3)
Standish, M. s.n. [o
E 1304, 3806, 8078,
9376, 11306, Ga 14200,
8374, 20402, 20601, 20783,
21309, 21935,
68829, 73654,
78703, Josi (3); 20910, 120996, 6
Steg , P. C. & Cha ae ; Km 1.
Scaler p.c. diis a EH
Standley, P. Valerio, R. 46663
466689 (3)
Stanford, L. R. 6938 (3).
e ere, W. C. Se 9.
cda M. 16 (3).
ate ah 951, 981 (4); 702 (30).
S M. 6164 > 9994, 7002 (4).
: E 3).
2 (2 s
Steyermark, J. A. : x CH n ao
50757, 51431, 5021 ( ,
(4); 47806 (8); 60022: (10); 57428
(27).
Stork, H. E. 528, 3397 (3)-
352
Stork, H. E., & Perso, W. J., & Beetle,
A.A 8732
SC? E. & Horton, ©. A 8606 (3).
Sürngar, V. E.R
Swartz, O. s.m. (3); sn, EOM
Tamayo, F. T. 314 (3); 197 (8).
Taylor, N. 449 (28).
Thi — 1223 (3).
Thompson, J. B. 284, 298 (4); 98, 803,
$76. D).
Tonduz, A. po 13940 (3).
Triana, J. J.
Trinidad Bot. aes 3570 (3)5:2772.(8);
1931 (28).
Tucker, —. 040 (8).
Ule, E. 6805 (3); 7114 (9); 5174 (15);
6256 (33).
Underwood, L. M. & Earle, F. S. r30 (3).
Underwood, L. M. & Griggs, R. F. 628
(4); 460, 599 (28).
Vahl, M. 5209 (28).
Valerio, R. d DN 2320, 2399 (3).
Vitas EIL 28):
Van GER OR P 706, 908 (3); 673
6 (3).
Vogel, P. C. s (4); 1455 (8).
[Vol. 43 _
ANNALS OF THE MISSOURI BOTANICAL GARDEN
Von Jacquemont s.n
on Türckheim, H.
3485 (28).
Wawra, H.
A
Weberbauer, A. (21).
HA SS ans (26).
25).
1385 bis, 2692, 2831,
; 130, I37 (2).
Williams, L. O. 11189, 14070 (3); 10170
(4
Vilis L-O,
& Molina, A. 12470,
13266, 14680, 16760 (3).
Wilson, P. des (3); 1345 (3 E
len, & Seibert,
Wright, C 2942 : E ); 2943, 2944
(8); 1386, 2046 ds); 2947 pes ih,
397, 400, 1383, 1384? (28).
Wright, C., Perry, C. C., & Brummel, H.
219, 407 (28).
Wiillschaegel, H. e 342 (4).
Wydler, H. 290 (28).
Xavier, L. 103 (28).
Yuncker, T. G., et al. 8078, 8655 (3).
Zaandam, —. 6885 (14).
Zainum, J. B., Molina, A., & Barkley, F. A.
B.O.I
II5 (8) .
Zanna Gutia, —.
5272 (3).
1956]
RAO—REVISION OF RAUVOLFIA
ENUMERATION OF THE SPECIES
RAUVOLFIA
SECTION I.
SE
6. blescbeti A DG.
7. mollis S. Moore
8. ligustrina R. & S
SECTION IIl. MACROVOLFIA
SER AE
1
SERIES 4. ANGUSTIFOLIAE
type indicates accepted, preéxisting names;
^ E und subgeneric categories are in CAPIT
ini
SUBSERIES 1. CUBANAE
21. andina Mgf.
00
weddelliene- Muel. Ted.
27. steyermarkii Woo
2 :
34. sellowii Muel.-Arg.
SYSTEMATIC INDEX
Huercki ia
Ophio.
auv
Sect. ANISOPHYLLAE .——
arborea
bahiensis
biauriculata REO OIN OIN CU
boliviana — — — ——— ———
pueda NE EE
flexuosa — — —————
var. glabra —————————
353
italics indicates synonoms; bold face
ALS.
354
[Vol. 43, 1956]
ANNALS OF THE MISSOURI BOTANICAL GARDEN
petaphsda — — 5 ee 314
polyphylla ` — 306
var. connivens __ 306
var. divergens 306
aecox 343
bata 291
pubescens 347
purpurascens 340
Sect. RAUVOLFIA 281
rhonhofia 347
salicifolia TIR
sanctorum 325
sara piquen sis 342
schulei 215921
sellowii Baur.
sessilifolia 330
inosa _. 347
cel Se
stenophylla 347
steyermarki oe
strem pelioides C SA7
subpubescen 285
ternifo. a 299
Ser. TERNIFOLIAR ..— ——— 499
tetraphylla AE 7 A
Ser. ease EE 281
tomentos Eco ue ue
Klee 291
weddelliana —— — |; 5209 330
wood ix cR TL TI ARE
Rauwolfia ON
Rerwolüs — =a o o 279
Strempeliopsis strempelioides --------------—-- 47
Tabernaemontan eeN E 347
Tondurza longifolia — — .—— Sua 347
Vallesia dichotoma -ae = 347
ADDENDUM: To the list of Beie Species" on p.
1825.
Reuvolfia brasiliensis Spreng
1:834
346 add:
Due to insufficient ere both
A. DeCandolle and Mueller- Ate have. also treated this as a dubious specie
EXPLANATION OF PLATE 3
Historical illustrations of Rauvolfia (see page 255)
pu t.
Plate 236 from Burmann's edition of Plumier's 'Plant. Amer. Gen.'
IZ93
Fig. 2. Illustration of R. tomentosa from Jacquin (Obs. Bot. pl. 36. 1763).
Ehret's drawing of a plant in the Chelsea Garden on which Linnaeus based
Fig. 3
his R. tetraphylla.
Fig. 4. Patrick Browne's illustration of R. canescens in the Linnean Herbarium.
ANN. Mo. Bor. Ganp., Vor. 43, 1956 P 3
LATE
` qm A
$ G: y + RIA a
S Aaivelfie tomentosa
2
RAO—REVISION OF RAUVOLFIA
MISSOURI BOTANICAL GARDEN
STAFF
Emeritus Director
Grorce T. MOORE
Grorce B. VAN SCHAACK,
CARROLL W. DODGE, ;
Mycologist of Herbarium
Roamer E : JULIAN A. STEYERMARE,
ior ds m Honorary c M sociate
REDERICK G. YER.
HENRY N. ANDREWS, Dendrologist ;
Faléohotanist Arce F. TRYON,
Rorra M. TrYO R x
Assistant Gerster ‘of the Ion D. Dwyer,
Herbarium Research Associate
Hoer C. CUTLER, E Next C. HORNER,
Curator Museum of Librarian and Editor
Economic Piants of Publications
Oscar GLAESSNER,
Business Manag
Second. Vice-President
Eucene PETTUS
Leicester B. FAUST -
HewaY HrrcHcOCK ——— = A WESSEL SHAPLEIGH `
RORERT BOOKINGS Surm EE
-EX-OFFICIO MEMBERS.
STRA
Volume XLIII Number 4
E. Annals
mi | | ef the s; e
.. Missouri Botanical |
Annals
of the
Missouri Botanical Garden
A Quarterly Journal containing Scientific Contributions from the
Missouri Botanical Garden and the Henry Shaw School of Botany of
Washington University in affiliation with the Missouri Botancal
Garden.
Informaton
"The ANNALS OF THE Missourr BOTANICAL GARDEN appears four times
during the calendar year: February, May, September, and November. Four
numbers constitute a volume.
Subri ës E Zeng > per volume
Single Numbers 0 each
Contents of previous issues of the ANNALS or THE MISSOURI BOTANICAL
-GARDEN are listed in the Agricultural Index, published by the H. W. Wilson
Company. :
Annals
of the
Missouri Botanical Garden
Vol. 43 NOVEMBER, 1956 No. 4
A NEW FOSSIL PLANT FROM THE NEW ALBANY SHALE WITH SOME
COMMENTS ON THE ORIGIN OF LAND VASCULAR PLANTS*
PART 1. CROCALOPHYTON, A NEw TRANSITIONAL SEA-LAND PLANT
HENRY N. ANDREWS ano KAREN S. ALT**
Introduction:—
The Devonian-Mississippian black shales of the east-central states have yielded
in recent years a fossil flora that is remarkable in the unique and diversified nature
of the plants. Within this broader area the New Albany shale of Kentucky and
Indiana has produced an exceptional wealth of plant fossils. It is noteworthy that
SO great a variety of plant groups has been described since the collections from
this area and horizon are not abundant. Few, if any, of the plants composing the
flora are known with a completeness that is in any way satisfactory, yet in spite of
the fragmentary nature of the specimens, they reveal plants that are novel in their
morphology and anatomy and disclose major taxonomic groups hitherto unknown.
The additional fact that several competent paleobotanists have not found it pos-
sible to agree on the affinities of certain constituents of the flora is indicative of
its exotic nature.
It is the purpose of the present paper to record a distinctive and particularly
problematical new member of the flora from a locality in Kentucky. In view of
its perplexing taxonomic status a discussion follows (Part 2) dealing generally
with some of the problems of the evolution of land plants which seem to be
inextricably related to the particular problem of our fossil’s affinity.
The geology of the New Albany shale has been dealt with in the comprehensive
study by Campbell (1946) and the fossil plants of the area with which we are
Concerned have been described by Read and Campbell and by Hoskins and Cross.
It seems unnecessary to review these previous paleobotanical studies although we
——
* The publicatión-of this investigation was aided by a grant from the National Science Foundation.
** Henry Shaw School of Botany of Washington University. St. Louis, Mo.
——
Iss
ued December 22, 1956. (355)
[Vol. 43
356 ANNALS OF THE MISSOURI BOTANICAL GARDEN
have included in the "Literature Cited" section a fairly complete list of the con-
tributions on the subject.
The reader will perhaps agree, if from only a casual glance at the illustrations,
that a few additional words of introduction may be justified. Our study of this
fossil has proven to be a fascinating if somewhat precarious venture; fascinating
in that it is anatomically the most unique fossil that has ever come into this
laboratory, and precarious in that its affinities remain, at the conclusion of our
studies, quite uncertain. Suggestions are offered on later pages concerning its
possible position in the plant kingdom, but, quite admittedly, these are speculative
although we hope that these speculations may prove to be of some significance.
It was evident from an initial inspection of the fossils that the cellular organi-
zation was highly unique, so much so in fact that consideration was given to the
possibility that it was of animal origin. However, detailed studies of the cellular
structure have left no doubt as to its plant nature, and other botanists who have
been kind enough to examine our preparations have agreed on this point.
Description: —
The following description is based on two specimens that were collected several
years ago by Mr. Charles B. Read (U. S. Geological Survey) from the New Albany
shale at a locality near Boston, Kentucky. . We are indebted to Dr. S. H. Mamay,
who is in charge of the Geological Survey's Paleozoic plant collections, for kindly
placing the two specimens in our hands for study. The specimens and all figured
slides are now preserved in the collections of the U. S. National Museum.
Since the two specimens are very nearly identical in size and form and
the surfaces revealed, under low magnification, apparently identical anatomical
— aid
Tp SS
Qua
"m
GEES
-fig. 1. Puen profile through xt-fig. 2. Longitudinal section through
eran U.S.N.M. No. 40 Gu? showing Zog ern axis showing the stratified or-
surfaces from whic thin tions were ganization of the fossil. Strands are shown
ami in black.
1956]
ANDREWS AND ALT—CROCALOPHYTON READI 357
structure, only one (No. 40797) was used in the preparation of thin sections.
Later it proved to be necessary to make a longitudinal cut through the second
specimen (No. 40798) and a study of this left no doubt as to the validity of the
original supposition that both represent the same species.
The specimens are conical (pl 4, fig. 1) being about 12 cm. tall, and the
diameters of the more or less oval base measure approximately 5.5 by 11.5 cm.
Externally the specimens are of a light grayish color, irregularly but not deeply
pitted. The irregularly broken aspect of their basal structure suggests that we are
dealing with the terminal portion of a trunk or branch of some sort. It may be
noted, however, that the horizontal orientation of the supposedly conducting tissue
offers some reason to believe that the specimens represent complete or nearly com-
plete plants; this possibility will be discussed on a later page.
Text-fig. 1 presents in profile the plan that was followed in studying specimen
No. 40797. Numerous transverse and longitudinal sections were prepared from
the A, B, C and D blocks, sufficient to insure a representative coverage of the
internal structure.
Although the preservation of the specimens is excellent the original plant
material has been highly replaced by phosphate of the calcium-magnesium group.
This lack of a sufficient mineral-organic differential, combined with the fact
that the specimens are rather soft and porous, prevented the use of the peel tech-
nique. No difficulty was encountered, however, in preparing ground thin sections.
The blocks to be sectioned were allowed to stand in melted resin for a few minutes
before affixing to the slide. The only particular caution found to be necessary
Was very careful grinding in the last stages due to the softness of the petrifaction.
The general organization of the fossil, which has been given the new generic
name Crocalophyton is shown diagrammatically in text-figs. 2 and 3. The speci-
mens are composed of but two tissues, one consisting of ray-like strands made up
of transversely elongated cells, the strands being enclosed in a thick-walled, nearly
isodiametric parenchyma. Whether the term parenchyma is appropriate may
questioned; it is at least as fitting as any that we are aware of. The strands are
shown in black in both figures. Text-fig. 3 is taken from the lower surface of
the B1 block. The figure was prepared from four transverse sections taken from
B1A, B1B, B1C and B1D as shown in text-fig. 1. After each section was photo-
graphed, the strands were inked; the four photos were then pasted together and a
tracing prepared of the entire cross-section. Text-fig. 2 is a diagram of the ar-
rangment of the same tissue system in a median longitudinal surface; this was
prepared from a free-hand sketch of the surface as it appeared under a binocular
microscope. It may be noted that these figures are intended to show the approx-
imate organization of the larger strands and that the photos should be relied upon
for a precise portrayal of their structure and pattern of organization.
[Vol. 43
358 ANNALS OF THE MISSOURI BOTANICAL GARDEN
Text-fig. 3. General organization of the strands in transverse section prepared from the
basal surface of the B 1 block.
The Strands:—
The term strand is applied to the highly unique ray-like tissue system
of the fossil. The cells composing the strands are transversely elongate
and pitted; they are presumed to have functioned as a radially con-
ducting system and are tentatively thought to be a primitive tracheidal
or sub-tracheidal cell type.
In their organization the strands, as shown in text-fig. 3, tend to radiate from
the central part of the specimen toward the periphery; this is, however, only a
very general tendency. It will help to clarify the nature of this curious tissue if
we glance next at fig. 3, which is a photograph of a representative portion of the
cross-section. It shows an area of approximately 1.0 X 1.5 cm. considerably
enlarged. A comparison of this with text-fig. 3 reveals the fact that the latter
does not include the numerous smaller strands that occur in many parts of the
specimen.
There are few strand terminations in the central part of the fossil although
occasionally the smaller ones end blindly. They form for the most part a continuous
and hiehly complex anastomosing system; a few continue to the extreme periphery
of the fossil and some end abruptly within a few millimeters of the periphery.
The strands vary from one to nine cells wide and are vertically elongate sheet
or ray-like structures. One must, as in the case of wood rays, observe tangential
and radial sections in order to acquire an understanding of their 5-dimensional
structure. While they present a superficial similarity to the wood rays of a conifer
or dicotyledon it is evident from text-fig. 3 and fig. 5 that they are by no means
comparably oriented,
Referring to fig. 3 (pl. 4) it will be noted that two lines have been drawn in
the lower right portion of the photo. When a longitudinal section is prepared in
the “A-A” plane (referred to as tangential) the rays appear as shown in fig. 4 (an
1956]
ANDREWS AND ALT——CROCALOPHYTON READI 359
appreciably higher magnification than fig. 3) ; and longitudinal sections prepared
in the “B-B” plane (referred to as radial) reveal the rays as shown in fig. 5.
The strands are about 1 cm. high and, as shown in the longitudinal profile
(text-fig. 2), they are distinctly stratified. Thus a transverse section taken
between the strand areas will reveal no strands. Most of them terminate blindly
although a few continue to the periphery of the specimen (text-fig. 3). The tor-
tuous pattern of the strands is emphasized in fig. 2 (and parts of fig. 3) where
they present a circular pattern in transverse view.
There is no externally delimiting tissue. The only evidence which suggests
that the original surface of the fossil is intact is the close similarity of the two
specimens. It is of course possible that some outer tissue or tissues was lost prior
to fossilization.
Detailed Structure of the Strands:—
Any transverse section (figs. 3, 6, 7) clearly reveals the variation in the width
of the strands which may take place within a few millimeters due to the anasto-
mosing pattern of this tissue and to the frequent intercalation of parenchyma cells.
Text-fig. 4. Representative portions from
several strands showing shape of strand cells.
One of the most striking characteristics of the strands is the órganization of the
cells in very nearly perfect vertical rows; this is clearly shown in tangential
sections (fig. 4). The cells are transversely elongate, their length appearing to be
extremely variable. In tangential view the strand cells are approximately square,
being 38 p in diameter, while their length, as observed in cross-section (figs. 6, 7),
varies between 105 and 230 p. Some cells, however, have been followed for a
considerably greater distance without any evidence ef terminations, but where
they have been observed they are for the most part transverse. Representative
Portions of strands are shown in text-fig. 4 where figures A, B and E were drawn
[Vol 43
360 ANNALS OF THE MISSOURI BOTANICAL GARDEN
from strands in transverse section and figures C and D from radial sections. While
some of the cells possess a tapering end, transverse or only slightly oblique end
walls are more usual.
he pitting in the strand cells is particularly interesting. In our initial suite
of slides many of the cells displayed obscure striations that were suggestive of
some sort of wall sculpturing but it was not until several radial sections were
examined with considerable care that clearly defined pits were found. Knowing
then what to look for they were found in several slides and we are reasonably
confident that the pitting is uniform on both the transverse and radial walls of
most or all of the strand cells. In fig. 5a and 5b are shown portions of two par-
ticularly well-preserved strand cells in radial view, and the structures which are
interpreted as pits appear in two rows wherever they have been observed with
clarity. Text-fig. 5 shows pitting in cells from two selected areas. No indication
|
Gan O
05005 8
a
o0000
od EE 20
0009 00000000
ooo Oo000o
900 0000065
000000 OO
= 300 5060 GO
xt-fig. 5. Pitting in the "radial"
S bei SÉ strand cells.
of a border has been noted and the slightly angular outline renders them somewhat
distinct from the bordered pits of most vascular plants.
The Parencbyma:—
The only cell type other than the strand cells is the tissue enclosing the strand
cells which has been referred to as parenchyma. This consists of cells which are
variable only within rather narrow limits and are for the most part isodiametric;
rarely are these cells more than twice as long as broad. A representative portion
of the parenchyma is shown in fig. 10. The transverse diameter of the parenchyma
cells (fig. 10), based on measurements of 140 cells, was found to vary between
1956]
ANDREWS AND ALT—CROCALOPHYTON READI 361
45 and 122 y, with an average of 80 p; in longitudinal view (fig. 9) their length,
based on measurements of 70 cells, varies from 52 to 165 p, with an average of
100 p.
The above figures are based on what is termed, as a matter of convenience,
"ordinary" parenchyma, referring to the relatively large areas where the paren-
chyma is not associated intimately with the strands (fig. 3) or with the purely
parenchymatous “inter-strand” portions of the specimen (text-fig. 2). The paren-
chyma between closely associated strands consists of cells that are slightly elongated
vertically and arranged in regular rows, assuming an organization comparable with
that of the strand cells. Fig. 11 shows a representative portion of such paren-
chyma between two strands appearing at the extreme left and right edges of the
photo. Parenchyma cells in such areas also may show a tendency to be elongated
transversely; this is evident between some of the strands illustrated in fig. 3.
Aside from these structural variations, which certainly are not extreme when
compared with the generally bizarre characteristics of the fossil, there is no evidence
to suggest any appreciable differences in the functional nature of the tissue.
The walls of the parenchyma cells are rather thick, being approximately 6-7 p.
This character may appear exaggerated due to the difficulty of distinguishing, in
the photos, between the actual cell wall and the mineral layer in the periphery
of the lumen.
Although we have stated that the parenchyma and strands constitute the only
two cell types, one other structure may possibly be distinct, It may be noted that
all of the photos which include any appreciable quantity of the fossil (figs. 3, 4,
8) display open areas or apparent breaks. In a very few instances we have observed
structures which suggest that some sort of secretory areas may be present. It is
apparent, for example in fig. 12, that the central cells are in an advanced stage of
disintegration but whether this cavity is a secretory structure or simply decay
cannot be determined for sure. In only two or three cases we have observed that
the parenchyma cells are radially aligned (fig. 13) in such areas. These are so
few that it is problematical as to whether any significance can be attached to them.
Tbe Ontogeny of Crocalophyton:— ;
The arrangement of the strand cells and closely associated parenchyma in very
nearly perfect vertical rows (figs. 4, 11) seems to imply meristematic activity of
some sort. Judging from the characteristic alignment of these cells they were
formed by an apical meristem which laid down, successively, the strand and inter-
strand layers or, at an early stage in the plant’s development, several meristematic
areas were differentiated. Whether the strands may be termed “xylary” is certainly
debatable. The cells are perhaps best termed “pre-tracheidal”; they are elongate
and pitted and probably served as conducting elements but the organization and
orientation of the strands can only be said to be unique. ;
If the argument presented in Part 2 is in any way valid the organization of the
fossil is not as surprising as may at first appear. Meristematic activities are perhaps
[Vol. 43
362 ANNALS OF THE MISSOURI BOTANICAL GARDEN
more varied in plants than is generally realized, and it is more than likely that the
algae, from which we suppose land vascular plants arose, experimented rather widely
in this way and in some cases succeeded.
Without necessarily implying a brown algal ancestry for any of the dominant
pteridophytic groups of the Devonian it is difficult to study living representatives,
such as Pterygophora with its stipe of radially aligned cells, “annual rings," and
its trumpet cells (Smith, 1939), without visualizing tendencies that were used to
advantage in the creation of a land flora.
With reference to the habit of land vascular plants it is evident that xylary
systems evolved along many different lines. Of particular interest here is Schizo-
podium davidi (Harris, 1929) from the Devonian of Australia. It is described as
having an outer wood zone "intermediate between primary and secondary wood",
the tracheids being cubical, vertically elongate, radially elongate or tangentially
elongate; and apparently not formed by a clearly defined cambial meristem. This
"outer xylem" may be considered a fumbling attempt (probably one of many) of
the plant to produce secondary wood. Crocalophyton may be looked upon as a
plant in which a unique, and apparently unsuccessful, mode of meristematic activ-
ity made its appearance. It is perhaps most aptly summarized in the words of one
botanist who examined our illustrations, as a "kind of foolish experiment" that was
involved in the evolution of land plants.
Crocalophyton readi! gen. et sp. nov.
Diagnosis of tbe Genus and Type Species: Conical-shaped structure, about 12
cm. tall with a roughly oval base measuring approximately 5.5 X 11.5 cm.;
stratified in a plane perpendicular to the long axis and consisting of strand-
parenchyma bands alternating with parenchyma bands; strands about 1 cm. high,
generally radiating from the center but forming a highly irregular system fre-
quently branching and anastomosing and contorted to the extent of forming a
loop, or series of loops, 1-9 cells wide (transverse), with the cells arranged in
vertical rows; strand cells horizontally elongated with mostly transverse end walls
and two rows of apparently simple pits in the longitudinal walls.
Type specimen: United States National Museum, No. 40797.
Locality: near Boston, Kentucky.
Horizon: New Albany shale; Upper Devonian or lowermost Mississippian.
Affinities: On the basis of our own observations and the opinions of specialists
familiar with the various cryptogamic groups, Crocalophyton shows no evident
relationship to any known thallophyte, bryophyte or pteridophyte. Since its affin-
On the assumption that this was a plant that lived ina h a transitional between strictly
aquatic or dry land, the be _ of the generic name is taken f Greek krokale, meaning
the
h or seashore; the s c name is in recognition of Charles B. Read's studies of the New Albany
shale plants.
1956)
ANDREWS AND ALT—CROCALOPHYTON READI 363
ities are so very vague we have felt that nothing is to be gained at the moment by
setting up any new taxon such as a family or order. A more detailed consideration
of the possible relationships of the fossil is included in Part 2.
Part 2. Some COMMENTS ON THE ORIGIN OF LAND VASCULAR PLANTS
AND THE TAXONOMIC PosITION OF CROCALOPHYTON
HENRY N. ANDREWS
In his introduction to a consideration of the problems of Jurassic paleobotany
Harris (1947) made the following comment:
t of the “problems” of paleobotany are those questions which we feel we would like to
be able to answer e: cannot because our knowledge is incomplete. Such problems are often
ephemeral. In time some are answered; some are left aside by progress because they cease to
bei interesting, or are ru realized to be meaningless;
I would supplement this with a paleobotanical proverb that has been attributed to
D. H. Scott, although most paleobotanists must have felt at one time or another,
that: “fossil plants frequently create more problems than they solve". This is not
intended as a facetious comment; it points to the great complexity and diversity
of organization that existed in many groups and we are forcefully reminded on
occasion of the necessity of altering our concepts of the lines of racial origin and
development within a group.
The problem of the origin of vascular plants has been with us for a long time;
it is not one of the ephemeral ones and if it is solved, at least in part, it will have
proved to have been a multi-faceted problem involving development along several
(or many) morphological lines. Many of us have become accustomed to look
upon Rhynia as the primitive land plant; there is now concrete evidence to suggest
that it is a primitive vascular plant representing a particular line of pteridophytic
evolution. In the following pages I should like to discuss what seems to me to
the nature of the problem; this is essentially a review although it may possibly
assist in clarifying the issues that are involved. The discussion is also aimed specifi-
cally at the question as to just why Crocalophyton does not conform to any tax-
onomic category in the plant kingdom.
.. The structure of Crocalophyton is quite unlike that of any known alga. The
possibility that it might be a member of the Phaeophyta was considered, but the
organization of the strands could be correlated in no way with the anatomy of any
brown alga. Several competent algologists have examined the illustrations and
have readily agreed that the plant has no place in the algae. As to the vascular
cryptogams there is equally little to choose from. Crocalophyton is unique in
the ray-like organization of the strands with their radially elongated, pitted cells
arranged in vertical rows. The pits show no evidence of a border, and the shape
of the cells is not closely comparable with that of a tracheid.
Thus, lacking any clues that might suggest affinities with a known group of
Plants it can be considered only on its own merits.
[Vol. 43
364 ANNALS OF THE MISSOURI BOTANICAL GARDEN
On the supposition that the strands constitute a conducting tissue of sorts a
guess may be made as to the plant's mode of life. The stratification of the strands,
being separated by rather broad bands of distinctly non-conducting tissue (the
parenchyma), combined with the radial elongation of the strand cells, would seem
to imply that whatever transport capacity the strands served was primarily trans-
verse rather than longitudinal. It is also unlikely that such a plant could have
attained any great height if it existed in a strict land habitat. If we next assume
that it lived submerged at a depth of as much as several fathoms, as do some of the
Phaeophyta, the trunk might have attained a considerable length but in such a
habitat it is difficult to understand why so profusely developed a conducting system
was needed. Since plants do seemingly strange things that do not always conform
with our understanding of their "needs", this possibility need not be entirely
ruled out.
If we postulate next an intermediate habitat, approximately in the tidal zone,”
a possible case for the strands may be made. A relatively short but stout trunk
attached to the rocks between low and high tide marks, and perhaps even somewhat
beyond the latter, lacking a root system or any other specialized organ of water
intake, would necessarily have to depend upon the absorption of water by the
surface of the trunk. Assuming a fairly strong absorption capacity for the exterior
surface the strand system could have readily conducted the water throughout the
trunk.
The assumption that Crocalophyton is a primitive semi-land plant, yet can be
assigned to no living or fossil group, would seem to call for further explanation.
Two general problems may have some real bearing on the subject. The first is the
gap that exists between the earliest land vascular plants and the algae from which
they are presumed to have evolved. The second, and I believe correlative, problem
is the evolutionary pattern that exists in any major group of plants, Briefly,
the concept that I wish to explore is that any large group (lycopods, articulates,
“ferns”, psilophytes, etc.) is highly polyphyletic and the transitional members
between it and the group(s) above it, if it so evolved, may be expected to be
correspondingly varied.
As a starting point, a few comments might be made on the origin of the algae
as well as their pre-Paleozoic and early Paleozoic record. The earliest record of
structurally preserved thallophytes is that reported by Tyler and Barghoorn (1954)
wherein they describe fossils that are tentatively assigned to the blue green algae
and “simple fungi” from rocks in southern Ontario that are dated as being in the
ZE of 1300 million years and possibly much older. These authors note that,
“Since the pre-Cambrian flora with which we are dealing comes from near the base
of the Gunflint formation, it seems likely that age may approach 2 billion years.”
? It has been called to my attention that the present discussion implies an origin of land vascular
plants from marine algae. I wish to acknowledge this criticism and indicate that the evolution of
D $b . + tionally
hina plants from plants of fresh water habitats or waters of varying salinity is not inten
1956]
ANDREWS AND ALT—CROCALOPHYTON READI 365
Another recent contribution suggests an even greater age for the algae. In
1941 Macgregor described supposed algal remains from Rhodesia in rocks which
have been dated as "at least 2,600 million years" and possibly in excess of 2,700
million years. Since they are not structurally preserved I feel that a considerable
element of doubt surrounds these Rhodesian algae; so far as I am aware, the Tyler
and Barghoorn report gives us the oldest authentic date of a structurally preserved
plant on the earth.
Before returning to the algae, for the purpose of my argument, a few words on
the earliest records of vascular plants will be interpolated here. The earliest un-
questioned record of such a flora is the mid-Silurian one (Baragwanathia, etc.)
described by Cookson and Lang from Australia. It is evident that these plants
were separated by a broad gulf of evolution from any algal ancestors.
Recently Krishtofovich (1953) has reported a supposed lycopodiaceous plant
from the Middle Cambrian of the Aldan Mountain range in Siberia. The plant
remains were associated with trilobites (believed to correspond to the Paradoxides
forchhammeri zone of Western Europe) and according to the author, “Conse-
quently the geological age of the find constitutes no doubt whatever.”
Krishtofovich’s Aldanophyton antiquissimum is represented by shoots up to 13
mm. wide and 8.5 cm. long which are covered with microphyllous leaves up to
9 mm. in length. "In places a thin rod-conducting bundle may be traced as far
as the base of the enations (leaves)”. Comparison is drawn with other early
lycopods such as Drepanophycus and Baragwanathia although sporangia were not
found in Aldanaphyton.
There have been several accounts of spores from the Cambrian which are pur-
ported to be those of land vascular plants. Several Indian investigators (Jacob
et al., 1952, 53a, '53b) have described spores and wood fragments from the Middle
and Upper Cambrian of Kashmir, Spiti, and other Indian localities. As many as 43
different spore types are recorded including those referable to Equisetalean plants,
“other primitive pteridophytes” and the Pteridospermae. These authors are insistent
that their preparations are free of contamination and, although I am prepared to
accept the existence of vascular plants in pre-Silurian times, the occurrence of the
Pteridospermae in the Cambrian is a matter that will require more substantial proof.
Naumova (1949) reports a considerable assemblage of spores from the lower
Cambrian blue clay of the Estonia-Latvia-Lithuania area (details of locality are not
Since the approximate date at which vascular plants first appeared has some
bearing on this discussion the high degree of complexity that already existed in the
lowermost Mississippian and Devonian seems significant.
3 Although I have had access to the original publication it is in Russian with no summary SCH
other language; I am indebted to Dr. Harlan P. Banks, who kindly supplied me with an English
translation.
[Vol. 43
366 ANNALS OF THE MISSOURI BOTANICAL GARDEN
Largely as a result of Florin's important contributions, we are particularly well
informed on the evolution of the cordaite-conifer line and certain associated groups.
With reference to the origin of this assemblage Florin (1949) notes:
The Ginkgoinae, Cordaitinae, Coniferac and Taxinae undoubtedly belong to the same
natural plant group of higher order, . . . but they constitute parallel evolutionary lines which
probably were already separated from each other in Upper Devonian or Lower Carboniferous
times. At all events, a clear differentiation can be seen as far back as the available fossil
records go. [pp. 101-102].
Judging from its stem anatomy the well-known and widely distributed Devonian
Callixylon is in all probability a coniferophyte and possibly a rather highly special-
ized member of the group; the fact that it was a large forest tree would suggest
that the coniferophytes may date back to an earlier period than the Devonian.
It is also pertinent to note the early appearance of heterosporous ferns, specifi-
cally Archaeopteris latifolia (Arnold, 1939) from the Upper Devonian (possibly
upper Chadakoin) of Pennsylvania, and Stauropteris burntislandica (Surange,
1952) from the Lower Carboniferous (Calciferous Sandstone series) of Pettycur,
Scotland. Just how these plants fit into the general stream of "fern" evolution is
problematical but there can be little doubt that they represent the culmination of
a long period of evolution of land vascular plants.
It is perhaps not surprising to find that the angiosperms are not to be excluded
from this general retrenchment in time of so many major groups. Under the new
binomial Sanmiguelia lewisi, Brown (1956) has reported leaves from the Dolores
formation (Middle to Late Triassic) of Colorado. They are alternate, large,
pleated, monocotyledonous, and closely resemble those of a palm. If this apparent
relationship is correct Sanmiguelia is the earliest angiosperm on record and suggests
a Paleozoic origin for the group.
In view of the evidence, direct and indirect, cited above I do not find it difficult
to accept land vascular plants in the Cambrian as a possibility or even a probability.
Next to be considered is the status of the algae during the approximate time
when plants were becoming established on the land. A few records based on fairly
well-preserved plants indicate the presence of several modern groups of algae in the
Devonian. These include smaller algae referable to the Chroococcaceae, Ulotricha-
ceae and Desmidiaceae from the Onondaga chert of New York (Bashnagel, 1942);
Charophyta from the Lower Devonian Downtonian beds (Croft, 1952) of west
Podolia (borders of Poland and Russia) ; fossils referable to the brown and red algae
from New York (Fry and Banks, 1955) ; and the remarkable fungi and algae from
the Rhynie chert (Kidston and Lang, 1921). There is abundant evidence that such
diversity, comparable with that of today, existed long before the Devonian—Fenton
(1946); Fenton and Fenton (1939); Johnson (in Smith, 1951); Pia (in Hirmer,
1927); Walcott (1919), and numerous other references cited by them.
The vast period of time that the algae existed prior to the advent of land
plants, even assuming their existence in the Cambrian, is somewhat startling. In a
1956]
ANDREWS AND ALT—CROCALOPHYTON READI 367
chart* recently prepared by the U. S. Geological Survey the beginning of Cambrian
time dates back 520 million years and the mid-Silurian about 340 million years.
Accepting the mid-Silurian "Baragwanatbia flora” as the oldest authentic record
of vascular plants and subtracting this (340 million years) from the figures of
Holmes or Tyler and Barghoorn cited above we find the algae existed from 960 to
1360 million years before the advent of land vascular plants. If we accept Krishto-
fovich's report of a Siberian lycopod from the Cambrian the figures are decreased by
another 100 to 200 million years, but we are still faced with the apparent possi-
bility that the algae existed for one billion years before migration to the land was
accomplished.
Unless land vascular plants did evolve much earlier than mid-Silurian times,
and by “much earlier” I imply at least several hundred million years, the obvious
and baffling question is why was the transition from an aquatic to a land habitat
so long delayed? This query is not a new one, and I allude to it only to present
two theories that may constitute a fresh approach whether or not they
contribute to the solution: One, that there existed prior to Silurian (or Cam-
brian?) times environmental conditions that rendered the transition difficult or
impossible. Two, that there was a great diversity of plant form involved in the
transition and that plants of the Rhynia type represent a successful arrival in but
one of many different lines that made the try.
As to the environmental factor, I have been intrigued with a suggestion by
Rachel Carson in “The Sea Around Us’. Assuming a closer proximity of the Moon
to the Earth in pre-Paleozoic times she postulates that the action of the tides may
have been vastly greater than at present creating a littoral zone, surrounding the
oceans, of such turbulence as to render the landward migration impossible. There
are obvious difficulties to accepting this as a significant explanation but it is at
least thought-provoking.
The second factor mentioned above, namely, the diversity of plant form that
may have been involved in the sea-land transition, may have a real bearing on the
taxonomic position of Crocalophyton.
Evolution in certain major groups, such as the lycopods or articulates, has been
a process of extreme polyphylesis once the basic morphology of the group was
established. The guess may be hazarded for larger groups, such as the conifero-
phytes or angiosperms, that they were polyphyletic from the start. The early land
plants (excluding clearly defined representatives of the lycopods or articulates)
present a highly complex assemblage; some may be justifiably classified as psilo-
phytes while others do not fit with any degree of comfort into any major category
and seem to imply a highly polyphyletic origin from the algae.
Few writers of text-books now portray the “family tree” of the plant kingdom
4 Prepared from the Report of the National Research Council, Committee on the Measurement of
Geologic Time, 1949-50. iis
5 See p. 158 of the original Oxford University Press edition.
[Vol. 43
368 ANNALS OF THE MISSOURI BOTANICAL GARDEN
as a great central trunk from which secondary branches progressively arise. Such
diagrammatic representations of evolution have passed out of favor because the
trunk of the tree failed to materialize, and I think it is clear that this failure was
simply due to the fact that the trunk never existed. Although I suspect that this
thesis could be successfully argued for any major group I will confine myself to
plants with which I have some special interest and which bear on the problem of
Crocalophyton.
It is becoming apparent, even in non-paleobotanical circles, that the term
"psilophyte" cannot be defined with any degree of precision. From the seemingly
simple (one might say classical) example of Rhynia a considerable diversity of
morphology is now known; this has been dealt with briefly but effectively by Miss
Leclercq (1954). The complexity in stelar anatomy that exists in Devonian and
lowermost Mississippian plants is perhaps not so well known. I refer particularly
to the stelar pattern in Rhynia, Asteroxylon and the assemblage of more compli-
cated forms including Pietzschia, Steloxylon, Siderella, Cladoxylon, Xenocladia,
and Periastron.
The problem of relationships among certain of these Devonian and Mississippian
fossils is a perplexing one, but, since in some stems only anatomy is known while in
others our information is more or less limited to the gross morphology, any postu-
lated relationships are obviously tentative. It is, however, instructive to contrast
the stelar anatomy of plants of the Rhynia type with the more complex polystelic
fossils such as Steloxylon and Xenocladia. The fragment of the stem of Xeno-
cladia described by Arnold (1952a) from the Middle Devonian of New York
measures approximately 1 X 5 cm. in cross-section (suggesting a stem of about 10
cm. diameter) and includes more than 40 steles. Several fragmentary specimens of
Steloxylon have been collected from the New Albany shale, the largest of which are
about 3 X 6 cm. and are composed of several dozens of steles. The contrast
between plants of this sort and those possessing slender monostelic stems presents
a deep and broad chasm to be spanned by any evolutionary bridge. It is possible
that they may have originated from an earlier, indeed much earlier, common land
plant ancestor but since there is no evidence to support this it seems at least equally
reasonable to postulate separate ancestry prior to establishment of the land habit.
Taking size and form relationships into consideration, if an early land plant
were originally large we might expect to find either a large stelar unit or severa
centers of stelar organization.
Prototaxites is an example of an early (possibly land) plant with longitudinally
aligned conducting cells throughout the trunk. This does not in any way imply
a brown algal ancestry for the groups that were successful in accommodating
themselves to the land habitat, but I believe it does present potentialities for a
conducting system wholly different from the Rbynia type.
Crocalophyton affords evidence that a plant of large diameter (as compared
with the Rhynia type) did exist and it apparently possessed an extensive apical
1956]
ANDREWS AND ALT—CROCALOPHYTON READI 369
meristem of a somewhat indecisive character. Conducting cells that are “almost
tracheids” were formed but in a very irregular fashion and apparently not con-
stantly, as is evinced by the vertical stratification of "vascular" and non-vascular
tissues. Briefly, it may be concluded that Crocalophyton is an early land plant
wholly different from the classical Rhynia.
We are accustomed to look upon Rhynia as “typical” or as a central type of the
psilophytes; at least it seems to occupy this position in most general accounts and
texts. I believe it occupies this position chiefly because of the excellence and com-
pleteness of the preservation. What we do know to be a fact is that the early
vascular plants were, by Devonian times and probably earlier, a highly complex
assemblage. It would seem to follow that these plants were highly polyphyletic;
they were radiating out in many directions a few of which became definable as
lycopods, articulates, coenopterids, etc. There can be no doubt that we have only
a fragmentary picture of this polyphyletic plan; discoveries of the future may or
may not clarify the path followed by the more successful groups of the Carbonif-
erous but they will certainly add to the complexity of the polyphylesis that is
already evident.
It seems clear that once a distinctive pattern of organization is established it
may radiate in many directions, and there may or may not be a conspicuous channel
which leads to another basic pattern. It is evident that the articulates and ferns
(and I should think to a somewhat lesser degree the lycopods) diversified rapidly
in many directions, although it would seem to be more difficult to define what we
mean by a fern than an articulate because of a greater ang rot: in the former
group.
The degree of morphological diversity that has existed in the algae is enormous,
and it is very likely that they were highly diversified in pre-Cambrian times.
point, therefore, that I have attempted to lead to is that in all probability several
(or many) groups of algae participated in the attempt to conquer the land. May
we possibly explain the diversification of morphology and anatomy that exists in
the early lycopods, articulates, the “psilophytes”, as well as the many that “do not
fit" as representing numerous lines of evolution from the algae rather than just
three or four? I believe this viewpoint receives support from other "groups
plants that defy classification. I allude here to fossils such as Profosalvinia and
Prototaxites,
Arnold (1954) notes:
ition = äere? deen in the plant kingdom is unknown, but it had evolved
being conn
The posi
vel [8] c rable t t of the lower bryophytes. . . ot cria ea
xai Protosalvinia during Devonian time was a
an
have underlined the word level. There subtle distinction between classing 4 plant as
SE of the Bryophyta and saying that it has d that level.
[Vol. 43
370 ANNALS OF THE MISSOURI BOTANICAL GARDEN
With reference to Profofaxites, the relations of this curious plant have recently
been reviewed by Arnold (1952b) in a contribution which includes a description
of a beautifully preserved new species. Not only is the taxonomic position of
Prototaxites very much in doubt but its habitat is as well; suggestions of various
authors place it anywhere from the deep sea to a land habitat.
As to Crocalophyton I can only conclude that it represents a curious effort, one
of a great many, in the struggle to evolve a vascular flora. Very possibly it or its
immediate descendants reached a dead end; at any rate it has no known contem-
poraries with which it may be closely compared.
Acknowled gement:—
Because of the exceptional problems involved in the matters of anatomical
interpretation. and the establishment of natural affinities we have called upon
authorities in several fields for advice. Their suggestions have been most helpful
and of not a little comfort although the authors assume responsibility for the
tentative conclusions that have been reached. We are particularly indebted to:
James M. Schopf, Charles J. Felix, Gilbert M. Smith, G. F. Papenfuss, and Johannes
Proskauer. Special thanks are due Sergius H. Mamay for kindly placing the
specimens in our hands for study and for assistance during the course of the
investigation.
Literature Cited: —
Arnold, C. " Aog Petrified wood in the New Albany shale. Science 70:581—
———————, (1939). Observations on fossil Pim from the Devonian of eastern Nd America. IV.
mut Univ. a Mus. Paleont. 5:272-314.
E Ibid. VI. Xenocladia che Arnold. Ibid. 9:297-309.
,(1952b). A ren of Prototaxites from the Kettle Point black shale of Ontario. Palae-
ontographics 93B:45—56.
» (1954) Fosil. sporocarps of the genus Protosalvinia Dawson, with special reference to P.
furcata (Dawson) comb. nov. Svensk Bot. Tidskrt. 48:292—300.
Bonae RA A. (19 M M microfossils from the Onondaga chert of central New York. Buffalo
Soc t. Sci, Bull. 173:
Brown, Rus Went vie Palmlike plants from the Dolores formation (Triassic) in southwestern
Colorado. U. S. Geol. Survey, Prof. P. Paper 274-H:205—
Campbell, Guy (1946). New Albany shale. Geol. Soc. Amer. Bull. 57:829—908.
Croft, W. H. (1952). A new perei (Charophyta) from the Downtonian of Podolia. Bull.
Brit. Mus. br Hist., Geol. 1:1
Cross, A. T., and J. H. Hoskins ( iS E Devonian-Mississippian transition flora of ms
United States. Compte Rendu Trois. Conger. Strat. et Geol. Carb MUN Heerlen, pp. 113-122
Darrah, W. C. (1937). Spores of Cambrian plants. Science 86:154—15
Elkins, Sa uo G. R. Wieland (1914). Cordaitean wood from A Indiana black shale. Amer.
ci :68—78
Fenton, C. L. (1946). Algae of the Pre-Cambrian and early Paleczoic. Amer. Midl. Nat. 36:259-
» and M. A. Fenton (1939). Pre Cambrian and Paleozoic algae. Bull. Geol. Soc. Amer.
50:89—126. : ^
Florin, R. ie i^ The morphology of Trichopitys heteromorpha Saporta, . . . Acta Horti Bergiani
Fry, W. L., and H. P. Banks (1955). Three new genera of algae from the Upper Devonian of New
York. Jour Paleont d 37-44.
Ghosh, A. K., and A e (1955). Did vascular plants exist in Cambrian times? Nat. Inst. Sci.
India (New Deli), Bull 7:298-303.
1956]
ANDREWS AND ALT—CROCALOPHYTON READI 371
Harris, T. M. (1929). poeseos davidi Ke ie e nov.—a new type of stem from the Devonian
ocks of Australia. Trans. Roy. Soc. Lon :395-4
————, (1947). The prob Kaes of pardos plc Bol. €— Geol. Portugal 6:1—32.
Hirmer, M. (1927). Handbuch der Palaeobotanik.
Holmes, A. (1954). The oldest dated minerals of the 0 shield. Nature 173:612-6
Hoskins, J. H., and A. T. Cross (1951). is structure and classification of four plants Fon the
New Albany shale Amer. Midl. Nat. 46:684—716.
The petrification flora of the Devonian-Mississippian black shale. The
——H
Pal bereet Sé 215-238.
coh, 2 C. Jacob, and R. N: —M atit: Evidence for the existence of vascular land
plants in the Cambrian. Current
(1953b). MAE nd tracheids ng vascular plants from the Vind-
ben. system, India: The advent of vascular plants. Nature 172:166—
Kidston, R., and W. H. Lang (1921). On Old Red sandstone Ens slowing structure, from the
atm Pen bed, Aberdeenshire. Part V. The Thallophytes, etc. Trans. Roy. Soc. Edinburgh
Krishtofovich,. A. (1953). estu of Lycopodium (Lycopodiaceae) in the Cambrian deposits of
iberia. E cad. Sci. U. S. SS R. 919:1377—1379. (Russian text)
Ladd, H. S., and R Se: 956). Fossils lift the veil of time. Nat. Geog. Mag. 109:363-386.
een. e (1954). ga aa psilophytes a starting or a resulting point? tnt Bot. Tidskr. 48:
Macgregor, e? M. diei A pre-Cambrian algal limestone in southern Rhodesia. Trans. Geol. Soc.
South Africa 43:9-15
Tee C. N. (1949). _ (Spore of the Lower Cambrian.—in Russian]. Acad. Sci. U.R.S.S, Bull.
r. No. 4, pp. 4
Read, c. B. (1935). A Kee flora from Kentucky. Jour, Po 10:215—2
» (1936). The flora of the New bad shale. Part tichnia Letzte a new repre-
eol. Surv. Prof. ewe res H:149-
1937 bue fora of de New nnda shale. Part 2. The ed Hr and their rela-
tionships. U. S. Geol. Surv. Prof. Paper 186-E:81-91.
* and G. C. Campbell (1939). Preliminary account of the New Albany shale flora. Amer.
1 Nat. 21:435-453
Smith PE e oe: The comparative histology of some of the Laminariales. Amer. Jour. Bot.
6:571—5
Smith, = = SÉ : pi Manual - Phycology. Chronica Bot. Co. 375 pp. Waltham, Mass.
Tyler, S. A., and E Barghoorn (1954). Occurre e =F structurally preserved plants in Pre-
mbrian ay 3 the C Ca dn n shield. Science 119:6
o K. R. (1952). e morphology of ee qoia Zeng P. Bertrand and its mega-
rangium Se fateris R. Scott. Phil. London 237B:73-91.
Valeo C. D. (1919). Cambrian geology and eent IV. E 5. Middle Cambrian algae.
mithson. Misc. Coll. 675:217-260.
[Vol. 43, 1956]
372 ANNALS OF THE MISSOURI BOTANICAL GARDEN
EXPLANATION OF PLATE
PLATE 4
Crocalophyton readi Andrews and Alt
Fig. 1. Specimen No. 40798 (U.S.N.M.), approximately natural size.
Fig. 2. Detail of transverse section showing looped form of strands. 23X. Slide
B1C,b1.
All figured slides are preserved in the Paleobotanical collections of the Hee
States National Museum. Slide numbers noted herein indicate the position from which they
were prepared as indicated in text-fig. 1
Ann. Mo. Bor. Garb., Vor. 43, 1956 PLATE 4
qe
.
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ANDREWS AND ALT— CROCALOPHYTON READI
on
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Ann. Mo. Bor. GARD., Vor. 43
[Vol. 43, 1956]
ANDREWS AND ALT—CROCALOPHYTON READI 373
EXPLANATION OF PLATE
PLATE 5
Crocalophyton readi Andrews and Alt
ig. 3. Transverse view through a representative portion of a strand zone. Line A-A
indica cates orientation of “tangential” section (see fig. 4); line B-B indicates orientation of
“radial” section (see fig. 5). 14X. Slide B1C,b1.
[Vol. 43, 1956]
374 ANNALS OF THE MISSOURI BOTANICAL GARDEN
EXPLANATION op PLATE
PLATE 6
Crocalopbyton readi Andrews and Alt
Fig. 4. Strands in tangential view. 40X. Slide A2-2, s2.
PLATE 6
Ann. Mo. Bor. Garp., Vor. 43, 1956
yor,
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ANDREWS axp ALT — CROCALOPHYTON READI
PLATE 7
ANN. Mo. Bor. Garp., Vor. 43, 1956
2
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y e ` x A)
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ANDREWS ano ALT — CROCALOPHYTON READI
[Vol. 43, 1956]
ANDREWS AND ALT—CROCALOPHYTON READI 375
EXPLANATION OF PLATE
PLATE 7
Crocalophyton readi Andrews and Alt
Fig. 5. Strands in radial view. 32X. Slide A2-1, st.
ag 5A, 5B. Pitting in radial walls of strand cells. About 600X. Slide A2-1, s2.
ES, 6, 7. Detail of strands in transverse section. 75X: fig. 6, Slide B1-B, b1; fig. 7,
Slide. ree tl.
[Vol. 43, 1956]
376 ANNALS OF THE MISSOURI BOTANICAL GARDEN
EXPLANATION OF PLATE
PLATE 8
Crocalopbyton readi Andrews and Alt
Fig. Longitudinal view showing occasional distortion of the normal “radial” align-
ment of strands. 34X. Slide A2-2, s2.
Aun. Mo. Bor. Garp., Vor. 43, 1956 PLATE 8
a 5 antes
A
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ANDREWS ano ALT — CROCALOPHYTON READI
PLATE 9
1956
,
ANN. Mo. Bor. GARD., Vor. 43
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ANDREWS anb ALT — CROCALOPHYTON READI
[Vol. 43, 1956]
ANDREWS AND ALT—CROCALOPHYTON READI 377
EXPLANATION OF PLATE
PLATE 9
Crocalopbyton readi Andrews and Alt
Fig. 9. Parenchyma in longitudinal section. 100X. Slide A2-2, t2.
Fig. 10. Parenchyma in transverse section. 100X. Slide A2-2, t2.
Fig. 11. Parenchyma associated with strands; tangential section. 100X. Slide Dy, s1.
[Vol. 43, 1956]
378 ANNALS OF THE MISSOURI BOTANICAL GARDEN
EXPLANATION OF PLATE
PLATE 10
Crocalopbyton readi Andrews and Alt
Areas possibly of a secretory nature. For further explanation see text.
Fig. 12. 77X. Slide DY, bi.
Fig. 13. 27X. Slide B1-B, b2.
ANN. Mo. Bor. Garp., Vor. 43, 1956
PLATE 10
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ANDREWS anv ALT — CROCALOPHYTON REA DI
A NOTE ON THE NODAL ANATOMY OF
ANKYROPTERIS GLABRA BAXTER
HENRY N. ANDREWS*
Considerable interest has been centered in the Carboniferous genus Ankyropteris
Since it presents, so far as I am aware, the earliest known instance of axillary
branching. A comprehensive understanding of the genus may be gained from the
works of Scott (1912), Holden (1930), Read (1938), Baxter (1951) and others
cited in these papers. It is the purpose of this note to supplement the description
of A. glabra Baxter (1951) with particular reference to the nodal anatomy.
The specimen of A. glabra described here was found in the same collection of
coal balls from which Baxter obtained the type specimen,— the Indiana No. 5 coal
about 10 miles north of Booneville, Indiana. This species was delimited from pre-
viously described species of Ankyropteris on the basis of the lack of multicellular
bairs and the origin of the axillary branch from the stem above the point of
departure of the leaf trace. When compared with conceivably related species I
believe the difference is actually more clear cut. In A. grayi Scott, for example,
the stele is much more angular, the central mixed pith is more conspicuous, and the
trace at the point of departure from the stem stele is triangular, being quite in
contrast to the more or less round (in transverse section) trace of A. glabra. Sim-
ilar differences clearly delimit A. glabra from A. corrugata (Holden, 1930).
On the basis of the type specimen Baxter pointed out that in A. glabra the
petiole ( phyllophore) trace departs from the stem before the axillary stele, the
atter remaining attached to the stem stele for another centimeter. In contrast
to this, in A. grayi a single triangular trace departs from the stem stele and then
divides into two, one trace becoming differentiated into the petiole trace and the
other into the axillary branch trace.
The specimen under consideration consists of a stem about 12 cm. long bearing
the basal portion of two petioles and associated branches. The internodal distance
is 7 cm., corresponding to that cited for the type specimen. Since the anatomy of
the two nodes differs somewhat from each other as well as from the type specimen
it may be convenient to consider them separately.
Node 1.—The node is identified by the departure from the stem stele of a
strand or “common trace” (pl. 11, figs. 1, 2) that is of essentially the same or-
ganization as the stem stele itself, differing only in its smaller size. It is not until
this is distinctly separate (fig. 3) that it shows evidence of differentiating into a
leaf trace and an axillary branch stele. In fig. 4 the two are separate, the leaf trace
being a slender, tangentially elongated strand and the axillary branch trace being
more or less circular in transverse section. The leaf trace next divides into two
essentially equal strands one of which passes off to the right, as shown in fig. 5,
* Henry Shaw School of Botany of Washington University. St. Louis, Mo.
956.
Issued December 22, 1 (379)
; [Vol. 43
380 ANNALS OF THE MISSOURI BOTANICAL GARDEN
and the other presumably develops into the main petiole strand, although preserva-
tion beyond this point is very poor. The significance of this early division of the
petiole trace is not clear. It may possibly be interpreted as a dichotomy of the
rachis, as is known in certain of the sphenopterid fronds and in Telangium affine.
Node 2.—The departure of the common trace at first follows the same pattern
described above; that is, it separates from the stem stele and then becomes differ-
entiated into an axillary branch stele and petiole trace (fig. 6). The latter, starting
as a tangentially elongated bar, develops into the characteristic H-shaped Ankyrop-
teris leaf (phyllophore) trace (fig. 7).
It is then apparent that the xylary nodal structure presented by this specimen
compares more closely with that described for A. grayi Scott than for A. glabra
Baxter. Since the specimen described here was obtained from the same coal ball
collection from which Baxter obtained A. glabra and the two compare closely in
every other way, there was apparently a certain degree of variation in the nodal
anatomy. Since this difference, although of considerable anatomical interest, is not
sufficient to warrant a distinct specific entity, it seems desirable to emend Baxter's
specific diagnosis:
ANKYROPTERIS GLABRA Baxter, emend. Andrews.
Stem oval in transverse section, approximately 14 X 9 mm.; stele 5-lobed,
consisting of very narrow discontinuous rays of small tracheidal cells surrounded
by much larger tracheids. Cortex a narrow inner zone of thick-walled cells and a
broad outer zone of thin-walled parenchyma; aphlebiae numerous; surface of plant
glabrous. Petiole (phyllophore) trace of the di-upsilon type. Axillary branch
stele arising either from the stem above the point of departure of the petiole trace
or from the division of a common trace.
Origin: Identical with that cited for the type specimen by Baxter (1951, p. 440).
Literature Cited:—
Baxter, Robert W. (1951). Amkyropteris glabra, a new American species of the Zygopteridaceae.
Amer. Jour. Bot. 38:440—452.
Holden, H. S. (1930). On the structure and affinities of Ankyropteris corrugata. Phil. Trans. Roy.
. Lond. 218B:79-114,
Read, C. B. (1938). A new fern from the Johns Valley shale of Oklahoma. Amer. Jour. Bot. 25:
335-338.
Scott, D. H. (1912). On a Palaeozoic fern, the Zygopteris Grayi of Williamson. Ann. Bot. 26:39-
69.
EXPLANATION OF PLATE 11
Ankyropteris glabra Baxter
_ Figs. 1-5. Stages in the departure of a petiole trace and axillary branch in Node 1.
Fig. 6. Stem stele, branch stele, and trace of Node 2. Fig. 7. Petiole trace at a slightly
higher level than shown in fig. 6. Peel preparations as follows: fig. 1, 888 B-t16; fig. 2
B-t4; fig. 3, C-b2; fig. 4, C-b6; fig. 5, C-b10; fig. 6, G-b6; fig. 7, G-b13.
Ann. Mo. Bor. Garp., Vor. 43, 1956 PLATE 11
TE
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ANDREWS — ANKYROPTERIS GLABRA
SOME LICHENS OF TROPICAL AFRICA. II. USNEA
CARROLL WILLIAM DODGE*
Through the kindness of Sir Edward Salisbury, Director of the Royal Botanic
Gardens at Kew, I have had an opportunity to study a large accumulation of
unnamed Tropical and South African lichens, of which nearly half belong in the
genus Usmea. At about the same time I received a smaller collection from the
East African Herbarium at Nairobi, Kenya, through the kindness of Dr. Bernard
Verdcourt. I also wish to express my appreciation of the courtesies of Mr.
Hamilton, then Acting Curator of the Herbarium of the Art Galleries of the
Glasgow Corporation, for permission to study types in the Stirton Herbarium, and
of Dr. John Ramsbottom and the late Miss Annie Lorrain Smith at the British
Museum, during my stay in these institutions while a John Simon Guggenheim
Memorial Foundation fellow in 1930. I am also indebted to Dr. I. Mackenzie Lamb
or permission to study specimens at the Farlow Herbarium, and to Dr. Ove Alm-
born for a few African specimens.
Most of the species previously reported by Motyka! were represented in these
collections as well as several previously undescribed species. On microscopic study,
I have transferred a few of Motyka's species to other sections or subsections and
have rewritten my key? to include the new species. Wherever there might be
ambiguity, as when a species is somewhat intermediate between sections, I have
included it in both. Since some of the South African species also occur in Tropical
Africa and the present collections contain some material from South Africa and
Madagascar, I have also included species reported by Motyka from those regions,
i. e. all species of the African continent and of the adjacent islands south of 15°
N. latitude.
Where I have seen specimens, the colors are recorded from comparison with
Ridgway (Color standards and color nomenclature. 44 pp. 53 pl. Washington,
D. C. 1912). Measurements of the cortex, algal layer, medulla, and axis should
nOt be taken too literally, as proportions of these parts are more constant in a
given species than the absolute measurements, It should also be remembered that
the thickness of the medulla as given by Motyka includes the algal layer as well
as the medulla proper. In groups with a thick medulla, the thickness of the algal
————
1 ` ` ; i k i 1936-3
VE Unocsin Miniont ad Lacum Tana er Sema a R. Pichi-
anno 1937 lectae. cree = 83-404, GE E E ee EE Milos
Florist. Geobot. 12:25—37. 1954.
novae vel minus cognitae generis Usnea [1.]. Frag. S
D Carroll W. Some lichens of Tropical Africa [I]. Ann. Mo. Bot. Gard. 40:271—401.
E
Sermolli
9545.
* Missouri Botanical Garden, St. Louis, Mo.
———
Issued December 22, 1956.
(381)
[Vol. 43
382 ANNALS OF THE MISSOURI BOTANICAL GARDEN
layer does not make much difference in the final ratios of the medulla to cortex
or axis, but it does in a few species with a thin medulla, as the algal layer shows
less variation in thickness from species to species than the other structures. As
far as possible transverse sections of a branch were made just above the first
dichotomy or branch, or, if an articulate species, from the middle of the first joint
above the first branch. If the basal portion of the plant were missing, I sectioned
the basal end of the portion available. As apothecia are often few and apothecial
structures are quite constant throughout the genus, I have seldom sectioned the
apothecia.
The species numbers in the following key are those used in the present mono-
graph to be continued in the next number of the Annars (Vol. 44, No. 1), and
the new taxa herein described are numbered as in the key.
KEY TO USNEA IN AFRICA SOUTH OF 15°
. Algae osa — grayish green, sometimes drying yellowish, rigid,
s 250-1150 u in diameter; medulla compact, usually thinner euer
ROCCELLINAE z
8
DH
ie EE o o EE
1. Algae Trebou eg
2. Bs Geer nee a e spinules may grow out im situ as short pseudoramuli
EE 3
2. Thallus clearly ramulose and fertile; exciple reticulate-rugose; axis hollow... 7
3. Apothecia present, i marinn cilia rare, distant; surface of the thallus ote aan 4
3 A pochi absen EE 5
4. Apo pem 0 mm. in diameter, almost eciliate; thallus not spinulose, up to 8 cm. tall;
uua. Erde S Khoder . cervicornis Dodge
4. Ads 3—5 (—6) mm. in diameter, cilia rare, short, subdistant; alte spinulose-
isidiose SÉ small papillae; axis solid; corticole; S. Sudan and K EUR
Phage Steiner
5. Axis — Meine not eroded; thallus up to 8 cm. tall; soredia SO te, Vue on
n pes t appearing farinose after iida a are shed; conical Uganda, Ze
tal orediosula Müll. Arg. in Motyka
zg
Axis very slighely ee surface longitudinally erode E and so esoe angular, compose
»
of interwoven strands es thick-walled hyphae; thallus 3—4 cm. tall; soralia
» farinose, sotedia dark fuscous;: sario ii ensi tie
5. Axis lacerate with many small de ics surface smooth; torte internodes short below and
above, longer in mid-portion, tips recurved, surface minutely verrucose, each
verruca producing a single isidium; both een and corticole; Kenya an
(on ar seruis E dep MC IMMUNE ET 4. U. nutans Dodge
te
Ka
. Axis hallow, loosely stuffed with brown hyphae; thallus branched near the base, ultimat
nches 7 cm. long, densely isidiose-spinulose throughout; mass pube.
EE pute e Dodge
6. deer es or csi yellowing; thallus sparingly dichotomous, 4 cm. d
; Southwest Africa U. Me eoensis (Vainio) Darb.
6. Medalla K E Ka isses a dichotomous, 3 cm. tall, 0.5—0.7 n diameter
uthwes U. seniors n (Vainio) Darb.
7. Aporhecia up to ge mm. in decet enen ESCH along the ridges; geraden
owded, 2 mm. long; disc white nose; ramuli dense t aleng
a: dini TIN mc? a large cavity; oP we Us a ee gees Dodge
ro hohe 5-7 (-10) mm. in diameter; exciple not ra ramulose; marginal e not een d,
long; disc buify brown, n muli irregularly dis
» i : ot prui
m. hee axis 0.75 mm. in diameter, poc nic A narrow; saxicole; K
Tan Liechte nsteinti ” Steiner
8. Axis holes or sans stuffed in larger branches thecia ciliate, rarely eciliate;
thallus smooth or papillate, not aie TAN Seats i isidiose. e Subg. EUMITRIA 4
8. Axis solid, rarely somewhat lacerate or with a very narrow ek cavity.
....Subg. EUUSNEA 2
1956]
DODGE—LICHENS OF TROPICAL AFRICA. Il. USNEA 383
9. Thallus distinctly angled or at least with papillae in de rows 10
3. e: s terete, not an ngled nor with papillae in dense Fos, EE 11
lus olive green, 2- to ege Sen? d; cortex continuous, larger ramuli smooth; soredia
isidiose along ridges beu the rare dins coastal Kenya, Tanganyika.
E ci d ee ea I JU. cristata Motyka
10. me — te olive below, shading through Isabella color to chamois above;
reolat KS — ramuli foveolate, sparsely papillate below; EE "à
Sé qa mm. in diameter, exciple smooth or papillate with geen id ech mou
RN 8. boragéeng e
11. Cavity narrow, or axis lacerate and cavity more or less stuffed with Zeg SE
1l. Cavity very wide and distinct, usua ally not sti Ee is
12. Thallus ESCH piter: eg a pia irregular m inc SCH ES branches
perpendicular, strai ee: r irre deu flexuous, tall, 2 mm. in diameter,
tapering pisci ro to gen eg slightly tide deren Angola. 2;
Vel witschiana EE
gulos
iooth; .. S Dosis ord HUM AAA E Marc im t
Lx Medalla. rose; ramuli long, slender, perpendicular, dense; apothecia
iameter, teca alau: sec? flesh-color, marginal cilia few an nd e geg
Bechuanaland, LCERATINAR), eseu U. MN Vainio
13. pud whites press 4—5 mm. ees thick, sparse to dense, appres mea to the branches;
ote b to 10 mm. in diameter, exciple and margin REP Kenya, Tangan-
. U. Liecbtensteinii Steiner
de
ROCCELLINAE A a des
14. Thallus cams sorediose. ....... AAA O NP
di A E
15. Medulla pe qeda iod sely filled; thallus 20 cm. long; Congo, Kenya........-.--- U. elata MM.
15. Medulla white, cavity densely filled with yellow hyphae; thallus 30 cm. long; ss pns.
16. Soredia granulose, isidiose from E tubercles over the whole plant.............. ria Motyka
16. Soredia on capitiform tubercles, mm. in diameter, mostly on the Zeie: branches.
ee ad vdd EE U. s f. bisorediata Morro
17. Thallus fertile, eg Sgr Ge EE,
12 Thallus aale aire EE »
18. Branches sli ira inflated; apothecia 7-10 (-15) mm. in ne iets exciple —_
papillate to e gl cilia rare; disc pruinose; medulla rose; Cameroons
be frm y SUR
18. po not dd: apothecia up to mm.
ucose and venose-rugulose; deos cilia dst: de cinnamon; medulla white,
ees ros S o 2. medio-efricans Dodge
18. Branches not inflated; apothecia 1-2 mm. in diameter; exciple convex, smooth, mar-
ginal cilia dos; disc carneous, hie pena m medulla white; i is de nana Dodge
19, Thallus. i3 .2 mm. in diameter, r e below, nude above; St. Helena,
iode and south to Réunion............—.—-- et U. E Ine ^Zahlbr.
19. Thallus 2 sm. in diameter, ramulose throughout... dis
in reenish ashy, la p e on tubercles, especially on the ramu
U. B Bailey 1 “Morya
20. Thallus remaini
3
20. Thallus Les Mars ei wn to russet t with paler ramuli
tuberc ingle minute isidia, not sorediose; teoria
21. Thallus og or Posi spoil flaccid, base n ot blackened. pice diste cicer ATAE 22
21. Thallus terete or an ually more rigid; base often Machine EE :
22. Tha
—
-
ite to pale S KR d
22. Thallus darker colored, 9 cm. or less tall, somewhat fruti re but rat de accid....24
23. Thallus white, green-punctate, 14 cm. long (incomplete), 1 1.5 mm. in diameter, summits
es laceous; apothecia 2 mm. in diameter, lateral on ts branches, estat A
Africa (AMABIL €: e de Uo s Mauer ee U. primitiva s
23. Thallus Ee clay color, partly somewhat fuscous, 20-30 cm. long, not MAN
h thecia and soredia unknown [ue
immi oe eo aie "versicolor Motyka
24. Thallus “fatt E 3c cm. 1. tall, 2 mm. in n diameter; aposhecia =p? to 2 E mm. in diam
densely ciliate; Ethiopia... . U. complanata (Müll. re Moryka
24. Thallus not distinctly flattened; SET EE EE
o...
[Vol. 43
384 ANNALS OF THE MISSOURI BOTANICAL GARDEN
25. Thallus up to 12 cm. tall, 2 mm. in diameter, acutely rugose; Ethiopia to Tanganyika.
16. U. corrugata wäi
25. Thallus Pe cm. tall, not acutely rugose
26. Branch bey ely -— ose; sor redia in farinose eege Ethiopia to Comoro Islands
(see also f. ech cilia Motyka) U. pulverulenta (Müll. Arg.) Moryka
26. Branches sparsely eg irregularly ramulose; b. idiom on low ridges, cortex =
t€ branches not areolate; French Guinea, Angola to S. Africa, pers acu an
Islands J. leprosa UM
27. Thallus Sëch with pseudocyphellae but not papillate, inflated, (10—) 2 cm. lon
ery la ART facul dE 28
27. Thallus M or, if articulate, without pseudocyphellae, often fruticose, papillate,
and ramul
` Thala up to 60 c cm. lon . in diameter, pale stramineous; joints 4—10 2 o
m. long, only slightly s ramuli 10 mm. long; isidiose on nee join
others; Eritrea to Tanganyika 18. U. flav Vescehs Wen
28$; TB db (10—) 25-60 cm. lo: ong, e ramulose or etai an occasional ENZ
29. Leite" ze elevated, praia uous; thall gem e
29. Pseud penis not elevated, rather inconspicuous; hallas usually fuscescent in the
arium 31
30. Thallus foveolate, 40-60 cm. lon, mm. in diameter; joints 5 (—10) mm. long;
alon SS? idges; pale di *cephalodia" often present; — Somaliland
19. U. praelonga Stirton
30. Thallus Se up to 40 cm. long, 2 mm. in diameter, joints 0.3-20 mm = is
apothecia 5 (-10) mm. in diameter, marginal cilia few; en aen Leone to
>
U. psendocyphellata Motyka
31. Thallus up to 30 cm. long, 2.5 mm. in diameter, relatively Cer? Geen up to 7
1
long, curved; Sierra Leone to St. Helena 21: Ui pecli « Moryka
sf. es 10-40 cm. lon 8» flaccid, joints 10-30 mm. long, nearly straight
. Thallus smooth or only s slightly deformed eg
o». Thallus foveate to 34
. Thallus 15-25 cm. rat ze -4 mm. in diameter, stramineous or pale stramineo-fuscous;
ecia 5-7 mm. in ta exciple smooth, marginal cilia dense cres long
. Africa and Madagas U. fle Ka Stirton
33. Thallus up to 60 cm. long, 1 up to mm. in diameter, lurid fuscous, summits long, capilla-
ceous; apothecia rare, 2-4 mm. in diameter, exciple lacunose, a cilia pes
but long; Kenya = Nyasaland, d Angola U. vesiculata Motyka
33. Thallus 20 cm. long, 1 mm. in diameter, sepia, summits cream bu E joints soto ef
inflated; RE and soredia unknown; Tanganyika to S. Africa ren: NE
AMOENAE) Gen "Motyka
33. Thallus KSE 10-15 cm. long, 0.5 mm. in diameter, summits emo a s Ses
ose; Ethiopia to Tanganyika as: U. - inuta Motyka
This boas to scrobiculate on ien? pa (25-30 X 4-5 mm.), smooth o
maller joints (5-10 X 1-1.4 mm.) with small ice of isidioee sor soria on "p. ie er
kenger iuri bud. ramuli 5 mm. gm on eg branches; corticole;
opia and Kenya 24 o ve? Sex arii Dodge
iameter,
oo foveolate, pe fuscous, up to 20 cm. long,
apillaceous; soredia pale sulfur eer N in ee d perar Kenya, ; Tan
a and Con 25. U. terrestris "Motyka
34. Thallus partly Seles to rugose on larger joints, not on pa 20 cm. long; jo:
o 10 X 2 mm. alternating with short joints 0.3 mm. in diameter e SE
uscous; no read quer no apothecia nor Sr “known; Kenya,
ganyika ( RUBIGIN: abrir SCENS Stein) Motyka
Tan
34. Thallus longitudinally rugose, lurid fuscous, up to Piae to 4 mm
diam ameter, summits pes M, more densely had ed ire ipod eii 5m
in diam eter, marginal cilia few; Et hiopia 26. U. rugosa a Moerka :
` Thallus e either persistently stramineous, yellow soon becoming fuscous in the herbar-
ium; cortex thin, papery, or, if thicker, very Keess gc H wale uiris and lax
(o often relatively thinner and more com the STRAMINEAE,
- Thallus ashy green, rarely red or, if stramineous, ‘hee papillate or Pe er Berens
and very late fuscous in the herbari SE
36. Thallus ridi iin ; elongate, rather flaccid, mostly more than 20 cm. long.
STRAMINEAE 37
36. Thallus short, under 15 cm. long, fruticose and usually more rigid..............—.... GLABRATAE 46
1956]
DODGE—LICHENS OF TROPICAL AFRICA. IIl. USNEA 385
37. Thallus samih pale yellow or pale ee subarticulate, usually eramulose; parte
ery ly fertile in U. gracilis Ach OENAE 39
37. Thallus st stramineous when fresh, soon Seen in the herbarium, abundantly E bu 58
+ Thallu 41
Thallus cialis rugose to ig ae sulcate ATAE 43
39. has weg pale fuscous in the herbar in diameter; gun bs See? the
in Senn K—; Ethiopia to epe ig "Madagascar to Rév
SR: Zeg Ach,
39. Thallus keng stramineous, 0.6 mm. cr less in diameter; medulla dier yide
s, K red; Ethiopia, Kenya and Uganda nioana s zai
39. Thallus. E P rd in the herbarium, 1' mm. or more in diameter, more oar summit:
aceous, medulla K— or yellow...........
40. Ramuli p to small di in upper portions of the branches; joints Server 2
ength, to 1.3 mm. in diameter, many inflated, clearly annula - me
yellow; ri Helena and Fernando Dosser io cle ch
40. Ramuli very rare or absent, not in clusters; thallus 1 mm. in diam Je jo
1-3 (-10 m. long, not constricted nor siet less annulate; medulla
Kenya to S. "Afri EE ap UE moniliformis docta
41. Thallus rose or copper-rose, medulla K—, 30 cm. long, 2-3 (—4) mm. in diameter; i
arly ramulose; no apothecia nor soredia known; saxiole Ethiopi iU. saxatilis Motyka
41. Thallus bicai fuscous, sorediose, medulla x » alle ot ARS AD A AO
42. Tha lus 1 m. long, 0.8 mm. i di tacts joints not constricted at crack
pw opi diated Ste thick, gea continuous in cracks, K yeilow then
ITY Dery, BEE E U. dasypogoides Nyl
42. su a Ze cm. long, 2.5 mm. in diameter, soft, sober d articula Es
surface indistinctly and minutely papillate; medulla K y then red; oom
- Rodriguez, Comoro, — and S i. Ve tue . U. stram E Arg.
42. Thallus 30—35 (—60?) mm . long, es up to 2 mm. in diameter below,
capillaceous above, indistinctly geste dite with oblon ridses, eem
lla ese apothecia rare, 2-3 mm. in cene omg slightly e
th few marginal cilia; soredia on ramuli; Zeg Veg Ro anda and jme
EEN exasperata (Mill. zen a
43. Thallus Rasy Ve vi eg and CHERCHE are? Se cm mm. in diamet
tely on some joints, eire y tips capillaceous, Rudy
apothecia lateral on secondary
br CANT ramuli pur but irregularly dis
; Egypt be Ethiopia p vk Kenya
1 m.
capillaceous; a rare and i regul rly on e ¿e
i i geg ve
43. Thallus more than 55 cm. long, about 1.5 mm. in diameter, longitudinally Zeg
ow, thin, tops farinose, not su ulcate between ridges; r: uE irre edt y disposed, up
o 30 mm. long, tuberculate, forming isidiose Seat
EE EE
the
U. fusca Motyka
34. U. derer p
43. Thallus more angled and -— sulcate between ridges on larger branches. POUR UM
43. Thallus art rticulate, r -fusc nating joints 2 mm. and very short; 0. j mm.
oy , some joints or foveate, others smooth; no apothecia soredia
a, Uganda, rs (RUBIGINEAE) +-.--0+-+--7 9s U. erubescens (Stein) Motyka
ig restado thallus over 40 em. long, ac acutely angled and
Ze rugose, dud de foveate between es;
mm. long. i u , thick below with obtuse tips; apot
known; isidiose soredia on tubercles od some ramuli; S ve TOES
44. Branches 1.3 mm. or less in diameter; apothecia about 10 n diameter...
45. Branches 1.3 mm. in eter, arpa up to 60 cm. cte eu pa and sulcate;
ii Sg base thick, tips acute; summits ts densely
ramuli irregularly cuore d g, base t > d: pare ci:
Ettel
EE nidi dd
dic. U a to
45. Braha 0.5 mm. in erc? er, Bac ieu 13 cm. long, ay diet Le
uli disposed along the whole thallus, 3-10 mm =. itorm;
angan ipe o Sout 3 Africa............ Se, 1 U. flactida Oral. o ign e Monks
3 nedulla lax, Soe eg
- Thallu "ssburicilee bebe ramulose; med x ëng e ramis me d
[Vol. 43
386 ANNALS OF THE MISSOURI BOTANICAL GARDEN
46. I with abundant spinules; medulla lax, fertile (also sorediose in U. zs
tirt ton) CABRIDAE 56
46. Thallus persistently yellow-stramineous, sorediate and sparingly fertile; SSC thick
46. Thallus palei GE yellow (becoming fuscous in U. incrassata), sterile (fertile in U.
s [LONGISSIMAE ] Bel ch m sg be referred here); medulla dense under
Seng becoming lax near the axis; eramulose, SSEOLEUCAE 59
47. Thallus eine m. long, minutely verrucose, sonieries mes with reticulate ridges, are er js
nde appearance; eramulose (sometimes isidiose soredia develop im sifu bo
dense clusters of pseudoramuli); terricole; Cameroons..................--- 39 U. erricola das
47. Thallus vie 15 cm. tall, very indistinctly papillate, papillae cylindric; ramuli tae
long, very slender; Tanganyika, Uganda and N. Rh SE
40. U. on iip Glo? Arg.) Motyka
47. Thallus up to 15 cm. tall, with distinct rugi but nearly terete; ramuli irregularly disposed,
variable in length and thicker; Tanganyika, Sidi and Mada eiit
41
U. usambarensis =>
47. lo Vues Aes much under 10 cm
thallus 2.5 cm. tall, esu branched, persistently pale stramineous,
dulla K— ; SSC isidiose in minute soralia; South Africa....... U. delicata Vainio
a Your Seege dense 49
Eramulose.
49. “Thales 7-10 cm. tall, 2 (-2.5) mm. in diameter, fuscous; moe K red; soredia white
o ye sii in tubercles; Congo to Cape of Good Hope............. : vob nn Laurer
49. Thallus. KA to iS tall, 1-1.5 mm. in diameter, old gold to Tsabelline, sedili Ky
Set. then fuscescent (red fuscous); soredia isidiose; ien cif
Unida os Africa; Madagascar (U. pete Motyka, p. p.)
3 0% jg brun Vainio
49. Tu d cm. tall 1 mm. in diameter, fuscous; medulla K—; soredia in ven
rcles Ven to Tanganyika 4 riw na Mocyks
50. Thala 7e m. diameter, rose to coppe nals aed K
tinctly yellowish fuscescent; soredia in slightly elevated oblong soralia; A Afric
ES asii “Motyka
Madag:
50. Thallus up to To cm
tall, 1.5 (- 2) mm. in diameter, ashy green to taw Lo
.medulla K—; soredia io in LE hemispheric soralia, Kr ming "Rino
eroons, Angola, and Uganda y; ie umanii i Motyka
51. Branches vom cylindrie, Ken ee inflated; D di. up to
diameter; Uga 46. U. s Mice Morii
51. Branches GEES bed, more or less closely EE
52; AM = indy pus gros ossly papi leie branches foveate and dE apotheci:
spem r; Mauritius, Réunion, and South Africa........... U. fusc Seral Motyka
Thallus s eerste papillate; spore = eh Jarger KEE in 7 iech Ste iod ud 53
53. tz 3—5 cm. tall, olive fuscous in the herbarium; Apos JE. n diam
xciple Ser marginal cilia Go ; Cameroons to South Africa..... i U. su ect Steiner
S cou. go
2 mm. in diameter, way olive wi ie cinnamon- “buff, See ram
11-12 mm. in diameter, exciple minutely verrucose with sad ds dal
an rm concrescent axis and ramuli; marginal cilia distant, Ade
a 49
53. Thallus. > cm. tall, 2
. ble prod es Dodge
Tan
53. Thallus SE cm. "t eg fuscous in the herbarium; apothecia 10-15 m
mm. in = — sa Dag Gute ha ^ recat
54. SEA upt
the
hoi DOLO abundan allus . tall, u
in diameter, dark few. ramuli E rare des irregularly dispos os
Uganda GER (acta) KE
54 pothecia up to 10 mm. in diameter
55, Thallus Ru to 10 cm. tall, 2 mm. in diameter; exciple smooth to spinulose; Tanganyika
Comoro Islands.. U. blepbarea Motyka
55. Thallus. de 7 cm. long, 1 mm. in diameter; exciple smooth to fo Sa marginal < ilia
distant, 5 mm. long; Fernando Po, Congo, South Africa, and Madag
e UL. ld Stirton
55. ims ` cm. tall, 1 mm. in diameter; exciple smoot th, ra d marginal cilia
merous, co Soe thick, almost inflated; Ethiopia to Tan a.
51-U. dits MOMS f. perstrigosa Motyka
1956]
DODGE—LICHENS OF TROPICAL AFRICA. IIl. USNEA 387
56. Thallus 9—30 cm. long, 1.5 mm. in diameter; tito a dispersed over most of
othe
the branches, spine s no ; apothecia 7 m n diameter, exciple smooth to
foveolate with short dade. ipd cilia fv, pres Gees o Cape of Good
Hope $2 U. sulle Stirton
sorediose; apothecia eg 47
57. ae about 30 cm. long, n diameter; spinules ET SECH perpendicular,
variously curved, the Mais "brc apothecia bur n diameter; exci n
ned smooth, marginal cilia dense, variously curved An branched, u up to
; Tanganyika to et of Good Hope and inicie ats te te 3, U. ëch Steiner
s7: Ca boul 5 cm. tall, 1.2 mm. in diameter; ramuli 3 mm. long; Goen to K
esia and Se Marques Sa erspinosa Motyka
Rho
58. Thallus about ër cm. tall, branches up to 2 mm. in diameter, sfr to
ile; An Motyka
58. Thallus S to 8 cm. tall, 1.5-2 mm. in diameter; ramuli rare; speci 3
diameter; Madagascar S Mauritius e Piko Motyka
58. — cm. tall, 0.5-1.2 mm. in parias ierra — inline branches
mulate sir ; apothecia m. in diameter; Madagascar........... soleuca Laien
59. Both fertile and sorediose (see no. pe above, U. gricors eb and U. stead
tyka).
59. Sorediose only. 60
59. Fertile e ilius up to 60 cm. long, about 1 mm. in diameter, etat with ën
short branches or ramuli eua. edits y holdfasts, chamois to deep colon
buff; apothecia 3-5 mm. in diameter, St smooth, eciliate p
a
SE maniensis Dodge
60. Eure z: cm. long, persistently osseoleuco-stramineous, 0.6—0.8 mm. in diam
Me. Motyki
60. Thallus e re cm, Gage long; primary branch 3 mm. m dismetet, secondary
oe 1-) 1 m. in diameter, becoming fuscous in dii petens Zei lla
the ke g eee Tee
61. Thallus die dicati un r late becomin E iud cou: the herbarium; medulla thick, hu ta
thin; irregularly branched, secun Sr Che or sub-smooth, apices vehi
thick Ll ee EE ITICAE 62
61. Thallus eres never fuscous; — relati "r thin, compact; cortex thick, often
horny (so gogo? scaling and or less ecorticate) ; branching more + regular. E2143
62. Thallus. Mike ut pseud d oet an ado ege? PE E E ADOCARPEAE 64
62. Thallus with PER RO SP oe Ge Een
t ra:
64. Tila, ir To um GE rate, rather rigid below, abia subrugose
and subfoveate on smaller branches; sterile; Eritrea (BARBATAE, C pr
EE 7. U. syriaca ceder
dione in U. Ledien M Asis
tuberculate yer papill
U. fila sap Wen
mm
EEN
64.
65. Thallus mm. in di inter becoming ferr óo-fuscow
st Ne ramulose; soredia PER Jan Ca 2 o vg ud d
65. Thallus 1 -2 mm. in diameter, olive-green, verruculose-papilla ate; apothecia u
verruculose, sorediose; marginal ge? few or
E ito Eius D SUD M ue aui e NUN EH U. par ese Moka
65. To e - mm. in diameter, m ashy stramineous, foveolate, ok pa —
minute, cylindric; eramulose; Cape of Good Hope. U. cornea Motyka
66. Exciple smooth « or slightly verrucose or ruguleeg- EE 7
66. Exciple RT Werer tt 68
66. Exciple Sage pA pies 10 mm. in diameter; thallus 7 cm. tall, citrine dra
to re olive buff; ramuli close, the shorter 1-3 mm. long, smooth, the gres T
o mm, long, Ee E in tuberculate soralia; Cameroons to
Ue apa 13 Motyka
n— Had
67. pare der 10 (-1 5) mm. in diameter; thallus u up to 15 cm. tall, scpia mag ro
herbarium; -— subdistant on larger branches, closer on oder cis,
E mm. long, t a smooth, the longer somewhat a but
not
sorediate; Prema to Angel, seenen U. bispida Motyka
[Vol. 43
388 ANNALS OF THE MISSOURI BOTANICAL GARDEN
67. CR 5 mm. in diameter; thallus 10 cm. tall, Saccardo's umber and buffy brown;
wood-brown to avellaneous, subdistant, 5 ez er a Eritrea to South
As 61. U. ochrophora Stzbgr. in oe
67. ee up to 9 mm. in eme Se about 5 cm. tall, buy clive to yellow
e; eramus Sudan to Tan 62. U. Rer pnr Dodge
68. Thallus about ng, warm sepia Verona brown; ramuli 1-10 mm. te? ag
shorter M the dree subescala but not cies igi» 5—
in diameter; papillae gece tiform; dees rea to Tanganyika....... ts rien Motyka
68. Thallus 5 cm. tall, light brownish-olive to Isabelline; ramuli iq disposed, 2—
ong, smooth; ene up to Gs mm. in diameter; Sierra Leone, Angola a nd
4. U. sentata Motyka
69. Thallus o pale fuscous, white-spotted, cortex hard; ee D (2-15) m
er; esorediose; ec pia and Sudan albomaculata Motyka
69. Thallus becoming fusco-sepia, o mall pseudocyphellae white, cortex Bik apot othec
Bareer eeh sulfur yellow on slenderer branches; Ethio;
U ABAT Moyka
69. E dirty ashy-greenish becoming pale fuscous or yellowish, = small pseudo-
yphellae AEA cortex soft; apothecia 10 mm. in diameter; Ethio
obtusata Motyka
70. Ramuli dense, pale fusc 51b. U. SS e pesii pasel Motyka
70. Ramuli less dense, eeng mias: branches yellowish-fuscous.
51a. U. obtusata f. — M
St Thallus olive green; branches up to 2.5 mm. in diameter € thick Sot
isidiose in punctiform Se Cape Verde, Camercont; C
TE pila Steiner
71. Thallus sepiaceous fuscous or stramin eo-yellow; medulla up to 3000 D thick 72
72. Branches 2-3 (-5) mm. thick, papillae in low ridges; soredia in ridges at summits
67
din, estaba Ethiopia U. Flotowii "Zahlbr.
72. Branches 2 mm. eter, stramineo-yellow, verrucae hemispheric, SCH distrib-
uted or in dien Zen soredia single, ovoid-spinuliform in Se uliform soralia,
white; Madagasc po aia Motyka
72. Branches 2 mm. in seg Gs vicarii papillate, SG CNA topi flattened;
à un ose; apothecia 10 mm. in diameter, exciple farinose-spot rd ia and
E EE 68 erpentaria Motyka
73. Miel: A thin, white, o ue thallus fruticulose or, if over 20 cm. ap ees cortex
reddish or reddish-macul
73. Medulla thicker, Gage rose or, E white, thallus over 25 cm. long, pendent; cortex
not reddish nor reddish maculate
74. Thallus a usually densely ramulose, 3 er eg tall, fruticose............... DENSIROSTRES
74. Thallus red or reddish-maculate, -fruticose or pendents soou an a RUBIGINEAE
75. Thallus saxicole, caespitose, 5 cm. tall, unbranched or or geg së only at the base 76
75. Thallus corticole, sparingly branched above the b. 79
76. Sparingly ramulose, branches few, Fe pliye ies green, rather lee ur sum-
mits incurved; rpm. to South Afri U. capensis Motyka
76. ge ramulose; b ranches several times s dichotomous at base, tips wi ramuli black-
ecia rare, mm. in diameter, esorediose; Southwest Africa, Tangan-
iac Melina: er Mauritius Gestion? with U. transvaalensis).
E U. pulvinata dins
76. Eramulose; axis solid or slightly lacerate, surface longitudinally e eed im ewhat
vom composed of interwoven strands of thick-walled hyphae; soralia farinose,
> dark O A EE 27
2 Medulla . K= or slightly e thallus sparingly ere Se cm. tall, 1 mm
r; Southwest A a (Vainio) Darb.
77. Medulla K red; thallus et dichotomous, 3 cm. tall, 0. i 7 mm
Ost Anh uu U M (Vainio) pus
78. Sterile and sore dices tcp i E EIN RA TED Ee TEPORE, Ee
78. Apothecia present..... puri
79. Thallus about 5 cm. L pale glaucous green, rigid, 1.5 mm. in diameter, closely sym-
ially br cd, ramuli dense, 1-2 (-5) mm. dica redia isidiose aeg
soralia in upper p. = a as from low ere ansia apis i
Aid iio oksa U. myrioclada (Müll. As y
1956]
DODGE—LICHENS OF TROPICAL AFRICA. Il. USNEA 389
79. Tolles ege? 8 cm. tall, pale stramineous, m. in diameter, divergently branched;
muli dense but somewhat ir Ke "aiser nd isidiose in "P tuber
culifo rm fons gc roded and Spam Madagascar... ambigua Motyka
79. Thallus. de cm. long, 0 m. in diameter below, ego nute SCH tubercles
muli ME disco 10-30 mm. ‘Tong; Togo, Donde Po, and i" Helena.
72. v pamm Motyka
80. Soredia dM covering tubercles, appearing spinulose when tuber cles crowded
an wën Bee Se pe are "vet thallus 5-6 cm. tall, between gg olive =
and tea is m r; ramuli 2-5 (-5 i MU cip ome den
sober i s er gege in WE See still immature) ; denied x Us putet p
80. Soredia absent
81. Thallus 8-10 cm. tall, 2 mm. in diameter; ramuli not abundant; papillae verruciform;
apothecia up to 20 mm. in diameter; Kenya to Cape of Good Hope.
9
8
eer Motyka
5. U. pict
81. Thallus up to 7 cm. dro 1 (-1.8) mm. in diameter; ramuli dense but fragile, eir
Ca 0.2 . from the branch and thus appearing very des “papilla
d. haben Dodge
€
s cia LI E in diameter; Keny
81. Thallus r cm. tall, in diameter, dei y covered with spinules 0. E
easily rn "off pus the thallus pooR: pue e; Mem m s mm.
diameter; Congo, Angola, Kenya and N Cs forie Weck
82. Thallus pendent, pae m. long, flaccid, ipdn
82. Thallus erect, 4—12 c RS rigid
83. Thallus densely ramulose, ess in n tubercles on upper portions, isidiose, olive Les in-
distinctly A e in diameter; Kenya, S. Africa and Réu
EA AE AE U. horridula (Mal. E Motyka
83. Thallus EE ramulose, res soredia in foster on ds low rtion
Senn ole Reis -maculat in diameter, acutely pesar e
5
ta ws of concrescent ze pilliform Err Kenya Pm 78. U. Dalei ëss
83. Thallus sto or agen ráre ramuli... concen
hallus m. long, 1 mm. in Ce soredia isidiose; thallus reddish nemah
. spilota Stirton
red-spo Cape of (e Hope...
84. Thales 20 cm. long, 2 mm. in di
th Foe eolate and rugose; fairy an
WË a erubescens (Stein) Motyka
Pm...
85. gen E reddish fuscous spotted with greenish fuscidulous,
nches about 1.5 mm. in diameter; apothecia a small, Mines ierit rugulose,
aie f eK n d ika, Gasen Islands a EN agasc
olive fuscous; Kenya, Tanganyi yrs n d Monks
Geesse eege EE KEE A
—€— 2 md idit
e o ad ee PH m rtm mt
e Thallus 10-12 cm. ges EEN gi
Thallus only 4-6 cm. tall..........—..———ámn007 Eege ER 88
87. Thallus u cm. Ria olive ash y green, deep red-spot Se ^ ` pe n di X" tim
oad fari ali th f part o the thallus; medula —;
ge P U. bicolorata s Motyka
d eroons an ind Reng PRA A A
87. Thallus Ma cm. tall, russet below shading to Mibi t above, up to 0.9
eter; soralia small with one to a few isidia, s es conuent and sene S e
fa diced after the E are shed; Ges EE U. U. spilotoides Dodge
88. Tm. 6 cm. tall, red or orange latericious, 1.5 mm. in in dia E n
es on des more br. ramuli; medulla ek e
Page y. parce n e) Monks
e; medulla Kenner
uds rió
89. "T Il enish s STRE 2.5 Gate? ter, tips e Mic Metis eege
reis seis yi E ..83. U. Meyeri (coim Motyka
as EE
89. Thallus hy go oodd 6 mm. in diameter, tips Ce not blackened; och sa xicole
corticole; Sudan to S. Africa and aes ET I EARE maculata Stirton
90. Thallus. SE or or cioe terete, more vias “1 mm. in
diamete dip uto branched, pale green (coppery fuscous in U. — T RR
90. Thallus "in cross ción; al at | re ER longitudinally Sen, more
yo angular diameter, unfrequently branched; green, ra rarely subfuscous.........-- GONIODES 96
"e terete or only very erai "longitudinally rugose, rarely sa er grayish
partly decortica NGISSIMAE 98
>
pavaberesesucevercesesecsossososresres
[Vol. 43
390 ANNALS OF THE MISSOURI BOTANICAL GARDEN
91. Medulla rose; thallus 5-8 cm. tall
91. Medulla white —— 25 cm. or more long, pendent, sparsely and irregularly papillate or
ul at
92. Axis with a narrow cavity; thallus 1 mm. in diameter; ramuli dense, 2 mm. long;
apothecia 7 mm. in diameter, See rugose; Kenya to Bechuaneland.
85. U. acanthera Vainio
92. Axis Rer thallus 1.2—1.3 mm. in diameter; ramuli rather close, 5-10 (215) m
sterile; isidia abundant, often single c or in q groups on verrucae, mostly
=Š ramuli - small branches; Morocco and Sierra Leone....... 6 U. marocana —
93. Thallus. 25 « cm. long, becoming owe Slain? N (-1.5 wi mm. in diameter; ramuli
shor diens Wei St. H U abi Motyka
93. Thallus P cms ore long
mm. in diameter, exciple smooth to subfoveolate, subglaucous;
dia r; ramuli dense, the larger branched; tubercles large;
94. deles apochesa I
de «in
ni Kc Cap Vert, Ss? Angola, Tanganyika, and S. Africa.
87
U. amplissima Stirton
9
94, € medulla K, at Sg one reddening or fuscesc
95. Thallus mm. in diameter, cortex areolate, uneven, tu
ae d on tubercles isidiose, finally farinose; medul Cap
ood Hope and Mauritius U. distensa Stirton
95. Thallus i mm. Se diameter, ded s often subseriate; soredia isidiose, cl Rt
ucae, e: medulla = ern reddening; Congo and Ken ng o Cape of G
car and Mauritius ontorii ncs
es
Hope , Ma romo:
96. Thallus 4 dee emp in transverse section, about 30 cm. long, ay green pee
olive buff, t
eem eed tdm i
la K red; e
-2 mm. in diameter, flattened at eem axils and ther
ide; zeien about 5 mm. long, I backs of ridges, up to 10 m shee near Se
summits; apothecia o 15 mm. in diameter, exciple ragore-reticulat and sub-
Gezmec eem pe to Cape of Good Hope and Madagasc
90. U. goniodes Stirton
97
96. Thallus with more — € than 50 cm. long
97. Ridges high and more or less alate. 98
97. Ridges lo lust e ra 99
98. TM 50 cm. long, 1 . in i diameter, pale olivaceous green, — Peer og
mm
ctured; almost order. Kenya, Uganda, ure pem dam
c E ou E U. E boreotler (Vainio) Moyli
98. Thallus up to 100 cm. € —3 mm. ai diameter, eyed sh green He d weg
sulcate between with minute white tubercles and papillae; ipie iis 15 mm. long
dtu more or less bheifad, giros ceed ate € sp inulose; Con,
T J. goniodes v. menn DeSmet & ee
99. Fertile; ramuli up to 40 mm. long; Bas Congo
99. Sterile; ramuli 5-10 (-20) m m. long; axis 1-2 mm. in diameter
100. Thallus 200 cm. long, 5 mm. in diameter, dirty ashy green, with tubercles a? Ze
papil ae; ramuli 20-40 mm. long, slender, very fragile; Sue 10 mm.
— ques uneven; Congo. U ipa Mn
100, Thallus - or more long, 1-2 mm. in diameter, grayish green; ridges corticate o
ith a Zen isidiose Sen plane eme the ridges with isidiose tubercles; la ul
bed 10-20 (-40) mm. long, 0.4—0.6 mm. in diameter, more or less isidiose-
tuberculate; true ramuli slenderer, 0.2-0.3 mm. i e
apothecia up to 15 mm. in dia meter; exciple smooth; marginal cilia few or
disc chalky Shyla Cong 92. U. Fe EH Duvign.
101. Thallus 100-110 cm. long, 1.6 mm. in eue ameter, dark olive buff; ramuli 4-10 (-20)
. long; s Reie nen on tubercles; Niseni: French Guinea Jesu ali anda,
Tanganyika, Transvaal. eg Motyka
101. Thallus 70 cm. or more deg P? i m. in diameter, grayish green, summits ema;
A acked with idos tubercles along the cracks; ramuli GE
ong, isidiose- See ate; Soe: s Feier
102. Geer well developed, on Semea branches, continuous or arole, Green scalin Gë
X soon scaling of o the EE ed
Cort
103. Mie axes " iudi in places, Ld diieuie longitudinally ru ru Sen EE
103. Main axes nearly terete, at least not flattened except in the axils -..105
1956]
DODGE—LICHENS OF TROPICAL AFRICA. II. USNEA 391
mm. in diameter, flattened in places t e, da
; ramuli irregularly mm (e
104. er in 100 cm. long, 0
e buff; medulla DCH rose, K reddening;
pes ase, tips often blackened; SE 3-4 m n diameter, plane; exciple
cl ghtly impressed; marginal cilia about 12, Weer in length; eg carneous
ensely — Sierra Leone to Cameroons eig iei Motyka
104. Thallus or more long, 1-2 mm. in diameter, slightly GH in place
irregula d ad oper pieni Stunt cked; medulla s s ydiovi
li close, . long, the longer sparsely isidiose; apot mm. in
ameter, concave; e SN with cracked, Steng cortex; ma ch EN few
y short; disc chalky white; Congo........-.--- Alsteeniana ia Daven:
105. Thallus. efe ashy, more than 20 cm. long, n diameter; ramuli sparse, irregu-
arly disposed, 10—30 mm bor: white tubercles, each bearing a ien de medulla
erna Helena...7 de A yngei Win
whi te, K inte
105. tage stramineous gree
ooth;
than 60 cm. long,
ff; ramuli 10-20 mm. dE apothecia 3-5 mm. in saei aptare sm
Tanganyika and Nyasaland................... e 96. ies Dodge
106. T more than 80 cm. eg g, 1.2-1.5 mm. in diameter, terete, yellow stramin ou
muli 10 (-20) mm. long, cu SN geng cia 2-4 mm. in dia e exci iple
ooth, eciliate; Zare r and Maude eere tn eburnea Motyka
106. Thallus sry green, 50 cm. or more long, 0. in di er; cortex ar es
ari ose from beicon de scars. of fallen dem
thecia
y smooth to very slightly pit tted; marginal c ilia
98. savanarum dec
ong,
in dreemen exciple near
few d Cái EES B
Thallus only bos " Een, longo ouai Ee
ne Kë about n diameter, terete; ramuli m mm. long, the longer
anched, s subpapillate bearing minute Wee axis Suh yellow hyphae: M Mada-
gascar we? lag: EE igena Motyka
107. Thallus about .in ditis terete with a Negeri low ridges p iving a
contorte P rance); ramuli irregula r, 10— subhelically twisted,
tuberculate; Së not seen; axis s wit th dark is zen. e age n Good Hope,
Ma agascar, Mauritius, and Réunion... . U. contorta Jatta
hië IE branches 0.5-1 mm. in donee ramuli 10-20 (-30) mm. has UT MEENTE c MSN 109
mary branches 0.4-0.6 mm. in diameter; ramuli 3-10 (-15) mm. Jong, nent 10
nches 1 mm. in diameter; ramuli 10-20 (-30) mm.
S. Rhod
101. U. € Motyka
i». That ls Los buff where corticate, $n
g, close; medulla K bright dell? Congo, Sudan to
EE branches 0.61 mm. in diameter, surface ru
109. Thallus "par green or grayish yellow, branches 0.6-1 m
ostly decorticate with spinules up to 3 m i ier d uli 10-20 mm. long, jon
er dies Gr Dog "steil eG
Se eee oie SO eS U. elata Duvign.
ate,
Fiet simple or forked, gni long
buff, — wholly decori
—8 mm.
110. Pei ema keel corticate; ramuli deep olive
.4-0.6 mm. iameter; ramuli 3-10 (-15) mm. long, close; apotheci
in diameter, sis ane; exciple smooth t o sublacunose, giereg cilia few, ‘a mm.
long; disc carneous, lightly pruinose; miden to S. Africa, — redu
EEN U. richodeoides Vainio
110. TM d apis green; axils and base of ramuli somewhat ix tte taies
até, tartareous; ra uli 3-10 mm. long, close, tips black; ci jar
St mm. in diameter, concave; exciple rugulose; marginal cilia abundant, up t
A — nn
ameter; cortex areolate
m. long; dise white; g
cm. or more long, 0.5 mm. in di
sling e sometimes farinose from coalescent scars of fallen isidia; ramuli
—5 mm. long, dense lateral branches tuberculate-isidiose; renes ES
m. in — sech nearly smooth to slightly pitted; C cilia few
CU. sevenarem Davies.
iiie ON
[Vol. 43
392 ANNALS OF THE MISSOURI BOTANICAL GARDEN
1. Usnea cervicornis Dodge, sp. nov.
Type: Southern Rhodesia, Makoni Inyanga District, summit of Forest-Hill
Kop, 1700 m., on shaded rock face, Frederick Eyles 827, at Kew. |
Thallus erectus, ad 8 cm. altitudine, 2 mm. diametro, teres, ad axillas com-
planatus, olivaceo-alutaceus; cortex areolatus, subfoveolatus, minute et irregulariter
papillatus, papillis apicibus nigratis; eramulosus. Apothecia lateralia, 8 mm. di-
ametro, margine crenato, eciliato aut cum 1-2 ciliis ad 8 mm. longitudine, ca. 0.5
mm. diametro, subinflatis, apicibus obtusis; discus brunneo-alutaceus, epruinosus.
4. Usnea nutans Dodge, sp. nov.
Type: Kenya, Northern Frontier Province, Mt. Kulal, 1935 m., on exposed
rock, P. R. O. Bally 5686, at Kew.
Thallus erectus vel decumbens, rigidus, ca. 8 cm. altitudine, cinnamomeo-
alutaceus ad obscure olivaceo-alutaceus in ramis ultimis; hapteron 5 mm. diametro,
cum decem vel pluribus ramis perpendiculariter dichotomis, curvatis, apicibus re-
curvis nutantibusque, teretibus, laevibus; cortex inferne annulatus subareolatusve,
superne minute verrucosus; isidia singula; eramulosus, rami ultimi nutantes, laeves,
apicibus nigris. Apothecia non visae.
5. Usnea Dodomae Dodge, sp. nov.
Type: Tanganyika, Dodoma Prov., Mt. Mpwapwa, 1610—1935 m., saxicole,
P. J. Greenway 2432, in East African Herb.
Thallus erectus vel decumbens, 10 cm. altitudine, rigidus in virgultis densis ex
gompho communi flavo, brunneo-olivaceus, Isabellinusve ad melleus in ramis ulti-
mis; rami 1.5 mm. diametro ad basim nigricantem ad 2 mm. diametro sub di-
chotomo primo, rami ultimi ad 7 cm. longitudine, flexuosi, eramulosi; cortex
inferne fractus et subareolatus, superne laevis; isidia spinulosa fere totam plantam
tegentia. Apothecia non visa.
6. Usnea pulvinulata Dodge, sp. nov.
Type: Uganda, Kigezi, Kasatoro forest, 1935-2560 m., on trees, I. R. Dale L39,
at Kew.
Thallus fruticosus, erectus, ad 5 cm. altitudine, ramis 1 mm. diametro, obscure
olivaceo-alutaceus, dichotome vel sympodialiter ramosus, internodiis brevibus, pul-
vinulum hemisphericum formans; cortex inferne nigricans, annulatim fractus sub-
areolatusve, superne laevis; densissime ramulosus, ramuli acuminati, 0.5-3 mm.
longitudine, recti; medulla inferne rubra.
Apothecia subterminalia, ad 15 mm. diametro, primum cupulata dein subplana;
excipulum tenuiter scrobiculatum, rugis ramulosis; cilia marginalia caespitosa, Ca.
2 mm. longitudine, apicibus acuminatis; discus laevis, juventute dense albo-
pruinosus dein rufus vel subaurantiacus.
1956]
DODGE—LICHENS OF TROPICAL AFRICA. II, USNEA 393
8. Usnea subcristata Dodge, sp. nov.
Type: Tanganyika, Kondoa District, Swaga-swaga Hill, summit, 1775 m., on
shrubs, chiefly Trichalisia cacondensis, B. D. Burtt $97.
Thallus rigidus, fruticosus, erectus, 7-8 cm. altitudine, inferne pallide brunneo-
olivaceus, superne Isabellinus vel melleus; rami basi 2 mm. diametro ad 3 mm.
expansi dein ad 1 mm. sub apotheciis, tenuescentes, angulati, papillati; cortex sub-
areolatus; rami secundarii 1-1.5 mm. diametro, axi principali similes; ramuli 1-15
mm. longitudine, minores sublaeves, majores foveolati, apicibus rotundatis.
Apothecia terminalia, plana, ad 20 mm. diametro, excipulo laevi vel papillato
radialibus cum rugis; cilia marginalia subdistantia in duobus ordinibus disposita,
extera crassa, dichotoma, intera 1-2 mm. longitudine, apicibus acutis; discus
vinaceo-alutaceus, pruinosus.
10. Usnea nana Dodge, sp. nov.
Type: Uganda, Kigezi, Kasatoro, Kabale River, 2256 m., corticole, I. R. Dale
L43 p. p. min.
Thallus fruticosus, erectus, ad 4 cm. altitudine, olivaceus, dichotome ramosus,
teres, ad 1 mm. diametro superne tenuescens, dense ramulosus; ramuli 1-2 (—4)
mm. longitudine, curvati, apicibus obtusis, raro dichotomi.
Apothecia sessilia in ramis tenuibus secundariis, cupulata, 1 (-2) mm. diametro,
excipulo laevi; cilia marginalia caespitose radiantia, 1 mm. longitudine; discus
concavus, carneus, albo-pruinosus.
12. Usnea medio-africana Dodge, sp. nov.
Type: Uganda, Kigezi, Mafuga, 2400 m., on trees, I. R. Dale L41, at Kew.
Thallus erectus, subrigidus, circa 5 cm. altitudine, 1.5 mm. diametro, super
basim semel aut bis dichotome ramosus, obscure brunneus, ramulis olivaceo-
alutaceis; teres vel subrugosus vel subfoveolatus; ramuli densi, 5-7 mm. longi-
tudine, fragillimi, laeves, apicibus acutis nigricantibus.
Apothecia subterminalia, ad 20 mm. diametro, plana; excipulum foveatum,
minute verrucosum, radiantibus cum rugis ex ramulis concrescentibus formatis;
cilia marginalia longitudine variabili, ad 8 mm. longitudine; discus cinnamomeus,
leviter albo-pruinosus.
13. Usnea brunnescens Dodge, sp. nov.
Type: Uganda, Kigezi, Mafuga, 2400 m., on trees, I. R. Dale L41, at Kew.
Thallus erectus, fruticosus, rigidus, ad 10 cm. altitudine, fusco-brunneus,
ramulis cinnamomeo-alutaceis; basi bis vel ter dichotomus, ca. 2 mm. diametro, ad
0.3 mm. tenuescens in apicibus obtusis; cortex inferne impressus subareolatusque,
sublaevis, superne dense verrucoso-papillatus; ramuli 1-3 mm. longitudine, in
ordinibus verticalibus, majores sparse verrucosi, singulis cum isidiis.
[Vol. 43
394 ANNALS OF THE MISSOURI BOTANICAL GARDEN
24. USNEA ARTICULATA (L.) Hoffm. subsp. aethiopica Dodge, subsp. nov.
Type: Ethiopia, below Debra Erki, on fir, Schimper 18, 21 Oct. 1850. det U.
articulata v. intestiniformis Ach. by Müller Argau at Kew.
Thallus pendens, flaccidus, plus quam 25 cm. longitudine, brunneus; basis 1
mm. diametro, 8 mm. longitudine, annulatus, non nigricans; dichotome ramosus,
ramis articulatis, articulis irregularibus, curvatis, cucumiforibus, 4-5 mm. di-
ametro, 25-30 mm. longitudine, foveatis vel scrobiculatis, pseudocyphellatis, vel
subrectis, 1-1.4 mm. diametro, 5-10 mm. longitudine minutis cum pseudocyphellis;
rami terminales, tenues, fere non articulati, dense ramulosi, ramulis 5 mm. longi-
tudine isidio-sorediosis.
34. UsNEA cameroonensis Dodge, sp. nov.
Type: Cameroons, Miss Cheeseman L3, at Kew.
Thallus pendens, flaccidus, plus quam 55 cm. longitudine, axes principales brun-
nei vcl cinnamomei, ramulis olivaceo-alutaceis; articulati, dichotome ramosi; axillis
complanatis, articulis ca. 10 mm. longitudine, 1 (-1.5) mm. diametro, non
conspicue inflatis, longitudinaliter rugosis, rugis tenuibus acutis, dorsis fractis
farinosisque; ramuli irregulariter dispositi, ad 30 mm. longitudine, basi subarticu-
lati inflatique, ad apicem acutum tenuescentes, tuberculati deformatique; soredia
isidiosa. Apothecia non visa.
36a. USNEA DECIPIENS Motyka var. Rhodesiana Dodge, var. nov.
Type: Southern Rhodesia, Matopos District, top of high granite hill, 1610 m.,
on tree, Frederick Eyles 1023, at Kew.
Thallus pendens, subflaccidus, plus quam 30 cm. longitudine, Isabellinus, in-
frequenter dichotome ramosus, nec articulatus nec annulatus, longitudinaliter et
subreticulate rugosus, rugis subacutis, dorso fracto, inter rugas sulcatus, 1 mm.
diametro, superne tenuescens sed sine summitatibus capillaceis; ramuli caespitosi,
3-15 mm. longitudine; tubercula farinosa sorediosa.
39. Usnea terricola Dodge, sp. nov.
Type: Cameroons, Mamoquilles, 2450 m., terricole among mosses and Sfereo-
caulon sp., Miss Cheeseman 25, at Kew.
Thallus fruticosus, subrigidus, 8-10 cm. altitudine, cinnamomeo-alutaceus,
axibus principalibus irregulariter nigro-maculatis, summitatibus obscure olivaceo-
alutaceis; basis nigra, dense dichotoma, ramis inferne 1 mm. diametro in medio ad
2-3 mm. dilatatis, dein superne ad 0.5 mm. tenuescentibus; articulatis, articulis 5-
20 (-30) mm. longitudine, apicibus constrictis, basalibus sublaevibus, minutis cum
pseudocyphellis; cortex fractus, rugis sulphureo-farinosis, isidiosis cum sorediis;
superne articulis plus minusve minute verrucosis, rugis plus elevatis et subreticu-
latis, soraliis majoribus elevatis, sorediis isidiosis. Apothecia non visa.
1956]
DODGE—LICHENS OF TROPICAL AFRICA. II. USNEA 395
49. Usnea blepharoides Dodge, sp. nov.
Type: Tanganyika, Kilimanjaro, at upper limit of forest, 3000 m., G. Geilinger
4417, at Kew.
Thallus fruticosus, pauci-ramosus, prostratus vel suberectus, ca. 9 cm. longi-
tudine, subarticulatus, inferne 2 mm. diametro, ad 1 mm. diametro sub apotheciis
tenuescens, subverrucosus, axibus fulvo-olivaceis, ramulis ciliisque cinnamomeo-
alutaceis; ramuli subdistantes, 4-5 (-10) mm. longitudine, curvati, basi subarticu-
lati inflatique, sparse papillati, apicibus acuminatis. Apothecia sessilia, geniculata,
subplana; excipulum minute verrucosum; cilia marginalia distantia, ad 10 mm.
longitudine, acuminata; discus planus, carneus, tenuiter albo-pruinosus.
62. Usnea subfoveata Dodge, sp. nov.
Type: Uganda, Bunyoro, Busingoro, on bark of Jacaranda, I. R. Dale Lóg.
Thallus fruticosus, erectus, rigidus, 5 cm. altitudine, flavo-ochraceus, dense di-
chotomus, articulatus, articulis 1-1.5 mm. diametro, 10 mm. longitudine, apicibus
constrictis; rami secundarii teretes, subfoveati, papillati in rugis; isidia singula vel
bina in apicibus papillarum; eramulosus. Apothecia subterminalia, ad 9 mm. di-
ametro, cupulata, dein plana; excipulum laeve vel verrucosum et subimpressum;
cilia marginalia distantia, ad 1.5 mm. longitudine, medio inflata, apicibus subacutis;
discus concavus dein planus, ochraceo-alutaceus, tenuiter albo-pruinosus.
73. Usnea Gossweileri Dodge, sp. nov.
Type: Angola: Cuanza Sul, Amboim, Capir near the Carloaongo River, 1000
m., on dead trees, J. Gossweiler 9991.
allus erectus, fruticosus, 5-6 cm. altitudine,
podialiter, superne dichotome ramosus; rami ca. 1 mm. diametro, annulati, dense
verrucosi-tuberculati, teretes; soralia super tuberculos insidentia, isidiosis cum
sorediis; ramuli 2-3 (-5) mm. longitudine, irregulariter dispositi, breviores laeves,
Apothecia rara, sessilia, 3 mm. diametro; excipulum
1-2 mm. longitudine, conica; discus carneus
olivaceo-alutaceus, inferne sym-
longiores minute tuberculati.
laeve; cilia marginalia subdistantia,
albo-pruinosus.
76. Usnea isabellina Dodge, sp. nov.
Type: Kenya, Northern Frontier Province, near Buna, summit of Ajao hill,
1030 m., on bushes, I. R. Dale L77.
us erectus, rigidus, 7 cm. altitudine, isabellinus; rami 1-1.8 mm. diametro,
dichotome ramosi, teretes; cortex laevis, annulatim fractus subareolatusque; ramuli
densi fragiles, 2-4 mm. longitudine, crassi, subverrucosi sed non sorediosi; apicibus
subnigricantibus; summitates ad 4 cm. longitudine, tenues. Apothecia rara, ca.
5 mm. diametro, lateraliter sessilia, plana vel subconvexa; excipulum laeve, sub-
rugosumve; cilia marginalia distantia, fragilia, longitudine variabilia, plus quam
3 mm. longitudine, ramulis similia; discus avellaneus, tenuiter albo-pruinosus.
[Vol. 43
396 ANNALS OF THE MISSOURI BOTANICAL GARDEN
78. Usnea Dalei Dodge, sp. nov.
Type: Kenya, Northern Frontier Province, near Buna, summit of Ajao hill,
1030 m., on bushes, I. R. Dale L70, at Kew.
Thallus pendens, subrigidus, plus quam 16 cm. longitudine, inferne aurantiaco-
cinnamomeus, superne obscure olivaceo-alutaceus, roseo-maculatus; basi subnigri-
cans annulatusque; dichotome ramosus, axibus principalibus ca. 1 mm. diametro
ad apicibus tenuescentibus, acute angulatus ex tuberculis papilliformibus concres-
centibus, inter rugis sulcatis, dense ramulosus in rugis corticatis; ramuli 4 (-15)
mm. longitudine, curvati flexuosive, tuberculati, cum spinulis paucis; summitates
tenues, non ramosae nec capillaceae.
82. Usnea spilotoides Dodge, sp. nov.
Type: Kenya, Chyulu hills, H. D. van Someren, in E. African Herb.
Thallus procumbens aut pendens, subrigidus, ca. 10 cm. longitudine, fulvus,
ramulis pallide olivaceo-alutaceis; sympodialiter ramosus, ad 0.9 mm. diametro, ad
summitates subcapillaceas tenuescens; inferne articulatus, articulis subinflatis,
apicibus constrictis, longitudine variabili, minute albo-verrucosus, sorediis isidiosis;
ramuli subdistantes, ca. 1 mm. longitudine, laeves, apicibus acutis; summitates
fragiles, ad 20 mm. longitudine, apicibus acuminatis. Apothecia non visa.
83a. UsNEA MEYERI var. chondroclada (Steiner) Dodge, comb. nov.
Usnea chondroclada Steiner, Verhandl. Zool. Bot. Ges. 53:228. 1903.
96. UsNEA amaniensis Dodge, sp. nov.
Type: Tanganyika, East Usambara, Amani-Maramba, 1030 m., growing in
festoons on the topmost branches of tall evergreen rain-forest trees, P. J. Greenway
4151, ex Herb. E. African Agr. Res. Sta., Amani.
Thallus prostratus, ramis tenuioribus pendentibus, rigidus, plus quam 60 cm.
longitudine, flavo-alutaceus, basi nigrescens, dichotome ramosus, 1 mm. diametro,
superne ad summitates capillaceas tenuescens, teres, laevis; cortex annulatim fractus;
ramuli 10-20 mm. longitudine, tenuissimi, recti vel curvati, fragiles, apicibus
acuminati minute tuberculati. Apothecia in ramulis sessilia, cupulata dein fere
plana, eciliata; excipulum laeve; discus fuscus, albopruinosus.
(To be continued)
GENERAL INDEX TO VOLUME XLIII
Ers scientific names of pua and the final members of new combinations are printed
bold face type; synonyms and page
prs and all other jurati in peii
numbers having reference to figures and plates, in
type
A
ipo Tropical, Some lichens of, II, Usnea,
HE EE antiquissimum, 3
Algae, evolution of vascular Su from,
364
,
3
Alt, Karen S, Henry N. Andrews and: A
>
of land vascular plants. P
es a new transitional. sea- d Ta
35
nie: nodal, of — glabra
Baxter, A note on the,
Anasazi sites, 194; best: dedu in, 2
transitional sea-land plant, 355; Part 2
me comments on the origin gen land
vascular plants an
tion of Crocalophyton, 363;
the medal anatomy of puse midi e zeit
Baxter,
Ang MM early record of, 366
Ankyropteris, 379; corrugata, 379; glabra,
M ei 350i grayi, 379
Ano 3
Archaeological sites: beans found in, 194,
hice latifolia, 366
Arrhenechthites, 74; — 79;
Le 77; egre , 78; mixta,
uin 78; tomentella, 24,95
AME qua eead ees , 304
TE Erichthitoid speciés of Senecio
Seiren climatic regions of, 112; geo-
logic history of, 115
Australian and New Zealand species of
Pittosporum, 87
B
Baragwanathia,
Basketmakers, nails of the, 194
Bean types: black-eyed, 190; common, 191,
4 90; kidney
: bri 192, 214, 251
48, 2
preparation of, 195; charts Moni vari-
ation in, 220—23
Belcher, Robert O.: A revision of the genus
s tites (Compositae), with inquiries
Mere an Arrhenechthites, 1
Bios
ag a eh 365
C
Eé: ges aia 30
Callix 366
Ca sibi. planta, 365
Citharexylum dentatum, 347; flexuosum,
347; molle, 347; retusum
Sé
e eech Senecio and Ar-
m
O
2
QRE,
Er: a
et
ES
E
ET
o
N
M
et
The Australian and
New Zealand species of Pittosporum,
Cordaite-conifer line, evolution e ded 366
Couma pentaphylla, 314
Crassocephalum valerianaefolium, 26
Croc gt phyton, A new transitional sea-lan
raid "fom & = Baga cone shale, 355;
Conifers phytes, evolution of the, 3
Cooper, Robert
oo
N
Crocalopkytón D radii, 36 2 Sag
Cyrtosipbonia, 279
D
Daylength, effect of, on habit of beans, 197
evonian-Mississippian shales, fossil flora of,
4,
Dodge, C Med William xm lichens of
tropical Africa. II, Usnea,
(397)
398
E
Economic uses: of species of prm.
d > species ipe juvet, dd
Ejot
uon Aden pte 365
Erechthites, 1
gt me “(Composiae), A revision of the
en ies into Senecio and
pad esta tin See ; index to species of,
so Sieten of Senecio in Austral-
<2 239
Zen dr iod recalar plants, 363
F
Ferns, heterosporous, early occurrence of,
Fossil plant, a new, from the New Albany
1,33)
shale
G
Gynura aspera, 20; malasica, 20; papuan
53; rosea, 26; zeylanica var. palates: 20
H
Hohokam culture in the Southwest, 194;
beans of the, 21
Hybridizacion 1 in Rauvolfia, 264
K
Kaplan, Lawrence: The arre beans of
the prehistoric Southwest, 189
Kentucky, a new fossil Wa from, 355
Ł
Land vascular plants, origin of, 355, 363
Lichens of tropical Africa. II. i ea, 381
dere
Lycopodiaceous Steeg eis 365
M
Malouetia jasminoides, 347
Marianthus procumbens, 182
5
the Southwest, 194;
ans of the, 207, 218, 229
N a
Navajo beans, 223, 233, 236
[Vol. 43
ANNALS OF THE MISSOURI BOTANICAL GARDEN
Neoceis, 10; carduifolia,
14; micr rocepbala, 61;
mentosa,
New LC mg shale, a new fossil plant from,
355
19; bieracifolia,
rigidula, 14; to-
New — and Australian species of Pit-
tospo
Nodal. pese of Ankyropteris glabra
Baxter, 379
O
Ophioxylon, 279
P
sonó A new fossil plant from the
w Albany shale, with some comments
on ida gem e de vascular para ce
A note on al anatomy of A
ropteris g Sibi Baires 379
Phaseolus, 189; acutifolius, 192, var. lati-
folius, 192, 206, 207; coccineus, 192, 206
207; lunatus, 192, 207; vulgaris, 191,
206, 207, subsp. aborigineus, 191, subsp.
ioliseaies, 191
Phylogeny of Pittosporum, 98
Piman beans, 22
Pittosporaceae, distribution of,
Pittosporoides, 121; umbellata,
Pittosporum: Aiscroliim mey a Zealand
species of, 87; interrelations of species of,
117; index of species, 187; index of ver-
nacular names, 187; Gase of, 121
Pomaderris microcarpa, 1
Prehistoric Southwest, d der beans of
e, 189
rotein, beans as a source of, 196
Prototaxites, 368
Pretopbytm evolution of, 363
Pteridospermae in Cambrian time, 565
FR bieracifolia, 14
R
Rao, Aragula Sathyanarayana: A revision of
Rauvolfia, with particular reference to
phy, 268; index to species, 353; relation-
pue 273; variation and hybridi
264,2
Rauvolfia v x medicinal use of, 267
Rauwolfia, 2
Rhynia, vg
Rio Grande Pueblo beans, 223, 233
1956]
S
Sanmiguelia lewisii, 366
izopodium diese 362
Sea-land plant, a new transitional, 355
Senec ecio and ebe E
Senecio, Erechthitoid species of, in Austra-
lia, e index to species, 85
Senecio, 10
Senecioninae key to the, 8
Serpentin
Soncbus Mond. 19; brasiliensis, 19; eryth-
ropappus, Za laevi 13; occiden tabs, 19
Southwest, culture ry of,
ay si dine Cultivated ius of the,
189
Stauropteris burntislandica, 366
Strempeliopsis strempelioides, 347
T
Tabernaemontana oppositifolia, 347
Tonduzia longifolia,
Trichilia monopbylla, 134
U
Usnea in Africa south of 15? N., 381; key
to, 382; new taxa, 392
399
amaniensis, 396; art culata
; subcristata, 393;
subfoveata, 395; terricola, 39
V
Vallesia dichotoma, 34
ariation: x sch bod) beans, 229-2390; in
Rauvolfia,
Vascular ge: And: E of, 563
Verde Valley beans, 2
Vernacular names of ide of Pittosporum,
187
X
Xenocladia, 368
x
Yuman beans, 223
Z
Zuni beans, 222, 236, 238
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